U.S. Department of the Interior

U.S. Geological Survey

Scientific Investigations Report 2016–5038

Alaska Arctic Marine Fish Ecology Catalog

Prepared in cooperation with Bureau of Ocean Energy Management, 

Environmental Studies Program (OCS Study, BOEM 2016-048)
Cover: Photographs of various fish studied for this report. Background photograph 

shows Arctic icebergs and ice floes. Photograph from iStockTM, dated March 23, 2011.
Alaska Arctic Marine Fish Ecology Catalog

By Lyman K. Thorsteinson and Milton S. Love, editors

Prepared in cooperation with Bureau of Ocean Energy Management, 

Environmental Studies Program (OCS Study, BOEM 2016-048)

Scientific Investigations Report 2016–5038

U.S. Department of the Interior

U.S. Geological Survey
U.S. Department of the Interior

SALLY JEWELL, Secretary

U.S. Geological Survey

Suzette M. Kimball, Director

U.S. Geological Survey, Reston, Virginia: 2016

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Suggested citation:

Thorsteinson, L.K., and Love, M.S., eds., 2016, Alaska Arctic marine fish ecology catalog: U.S. Geological Survey 

Scientific Investigations Report 2016-5038 (OCS Study, BOEM 2016-048), 768 p., http://dx.doi.org/10.3133/

sir20165038.

Chapter within report:

Love, M.S., Elder, Nancy, Mecklenburg, C.W., Thorsteinson, L.K., and Mecklenburg, T.A., 2016, Alaska Arctic marine 

fish species accounts, in Thorsteinson, L.K., and Love, M.S., eds., Alaska Arctic marine fish ecology catalog: U.S. 

Geological Survey Scientific Investigations Report 2016–5038 (OCS Study, BOEM 2016-048) p. 41–48, http://dx.doi.

org/10.3133/sir20165038.

ISSN 2328-0328 (online)
iii

Acknowledgments

This study was supported by the U.S. Department of the Interior, Bureau of Ocean Energy 

Management, Environmental Studies Program, Alaska OCS Region, through Interagency 

Agreement Number M09PG00005 with the U.S. Geological Survey (USGS). We appreciate the 

program planning advice and project reviews by fishery specialists in the Bureau of Ocean 

Energy Management and all expert technical reviews provided during the USGS-managed peer 

review process. We extend special appeciation to the USGS Tacoma Publishing Service Center 

for their editorial, production, and publication assistance. 
iv

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v

Contents

Acknowledgments........................................................................................................................................iii

Chapter 1. Alaska Arctic Marine Fish Ecology Catalog—Chukchi and Beaufort Seas

By Lyman K. Thorsteinson......................................................................................................1

Chapter 2. Alaska Arctic Marine Fish Inventory

By Lyman K. Thorsteinson....................................................................................................23

Chapter 3. Alaska Arctic Marine Fish Species Accounts

By Milton S. Love, Nancy Elder, Catherine W. Mecklenburg, 

Lyman K. Thorsteinson, and T. Anthony Mecklenburg....................................................41

Pacific and Arctic Lampreys.......................................................................................49

Pacific Lamprey (Entosphenus tridentatus)....................................................49

Arctic Lamprey (Lethenteron camtschaticum)...............................................55

Spotted Spiny Dogfish to Bering Cisco .....................................................................60

Spotted Spiny Dogfish (Squalus suckleyi).......................................................60

Arctic Skate (Amblyraja hyperborea)...............................................................66

Pacific Herring (Clupea pallasii)........................................................................70

Pond Smelt (Hypomesus olidus)........................................................................78

Pacific Capelin (Mallotus catervarius).............................................................83

Arctic Smelt (Osmerus dentex) ........................................................................91

Arctic Cisco (Coregonus autumnalis)...............................................................98

Bering Cisco (Coregonus laurettae)...............................................................107

Broad Whitefish to Dolly Varden..............................................................................112

Broad Whitefish (Coregonus nasus) .............................................................112

Humpback Whitefish (Coregonus pidschian) ..............................................121

Least Cisco (Coregonus sardinella)................................................................128

Pink Salmon (Oncorhynchus gorbuscha) .....................................................136

Chum Salmon (Oncorhynchus keta) ..............................................................142

Coho Salmon (Oncorhynchus kisutch) ..........................................................148

Sockeye Salmon (Oncorhynchus nerka) ......................................................153

Chinook Salmon (Oncorhynchus tshawytscha) ...........................................158

Dolly Varden (Salvelinus malma).....................................................................163

Inconnu and Glacier Lanternfish..............................................................................172

Inconnu (Stenodus leucichthys) ....................................................................172

Glacier Lanternfish (Benthosema glaciale)...................................................179

Ice Cod to Pacific Cod................................................................................................183

Ice Cod (Arctogadus glacialis) .......................................................................183

Arctic Cod (Boreogadus saida).......................................................................189

Saffron Cod (Eleginus gracilis) .......................................................................201

Walleye Pollock (Gadus chalcogrammus) ....................................................209

Pacific Cod (Gadus macrocephalus) .............................................................218

Threespine Stickleback to Antlered Sculpin..........................................................226

Threespine Stickleback (Gasterosteus aculeatus) .....................................226

Ninespine Stickleback (Pungitius pungitius) ...............................................233

Whitespotted Greenling (Hexagrammos stelleri) ........................................239

Okhotsk Hookear Sculpin (Artediellus ochotensis) ....................................245
vi

Contents—Continued

Chapter 3. Alaska Arctic Marine Fish Species Accounts—Continued

Threespine Stickleback to Antlered Sculpin—Continued

Hamecon (Artediellus scaber) ........................................................................249

Antlered Sculpin (Enophrys diceraus) ..........................................................254

Arctic Staghorn Sculpin to Ribbed Sculpin............................................................259

Arctic Staghorn Sculpin (Gymnocanthus tricuspis) ...................................259

Butterfly Sculpin (Hemilepidotus papilio)......................................................265

Twohorn Sculpin (Icelus bicornis) ..................................................................270

Spatulate Sculpin (Icelus spatula) .................................................................274

Belligerent Sculpin (Megalocottus platycephalus) .....................................279

Brightbelly Sculpin (Microcottus sellaris).....................................................283

Plain Sculpin (Myoxocephalus jaok) .............................................................287

Great Sculpin (Myoxocephalus polyacanthocephalus) .............................292

Fourhorn Sculpin (Myoxocephalus quadricornis) .......................................298

Arctic Sculpin (Myoxocephalus scorpioides)...............................................305

Shorthorn Sculpin (Myoxocephalus scorpius) ............................................310

Hairhead Sculpin (Trichocottus brashnikovi) ...............................................316

Bigeye Sculpin (Triglops nybelini) ..................................................................320

Ribbed Sculpin (Triglops pingelii) ...................................................................324

Crested Sculpin to Leatherfin Lumpsucker............................................................330

Crested Sculpin (Blepsias bilobus).................................................................330

Eyeshade Sculpin (Nautichthys pribilovius) .................................................335

Polar Sculpin (Cottunculus microps) .............................................................339

Smoothcheek Sculpin (Eurymen gyrinus) .....................................................343

Alligatorfish (Aspidophoroides monopterygius)...........................................347

Arctic Alligatorfish (Aspidophoroides olrikii) ...............................................353

Fourhorn Poacher (Hypsagonus quadricornis) ...........................................358

Atlantic Poacher (Leptagonus decagonus) ..................................................363

Bering Poacher (Occella dodecaedron) .......................................................368

Tubenose Poacher (Pallasina barbata) .........................................................373

Veteran Poacher (Podothecus veternus) .....................................................378

Pimpled Lumpsucker (Eumicrotremus andriashevi) ...................................383

Leatherfin Lumpsucker (Eumicrotremus derjugini) .....................................387

Sea Tadpole to Polar Eelpout....................................................................................391

Sea Tadpole (Careproctus reinhardti) ...........................................................391

Nebulous Snailfish (Liparis bathyarcticus) ..................................................396

Gelatinous Seasnail (Liparis fabricii) .............................................................400

Variegated Snailfish (Liparis gibbus) .............................................................405

Kelp Snailfish (Liparis tunicatus) ....................................................................410

Black Seasnail (Paraliparis bathybius) .........................................................415

Halfbarred Pout (Gymnelus hemifasciatus) .................................................420

Fish Doctor (Gymnelus viridis) ........................................................................424

Adolf’s Eelpout (Lycodes adolfi) .....................................................................429
vii

Contents—Continued

Chapter 3. Alaska Arctic Marine Fish Species Accounts—Continued

Sea Tadpole to Polar Eelpout—Continued

Doubleline Eelpout (Lycodes eudipleurostictus) .........................................433

Glacial Eelpout (Lycodes frigidus) .................................................................437

Shulupaoluk (Lycodes jugoricus) ...................................................................441

White Sea Eelpout (Lycodes marisalbi) ........................................................445

Saddled Eelpout (Lycodes mucosus) ............................................................449

Wattled Eelpout (Lycodes palearis)................................................................453

Polar Eelpout (Lycodes polaris) ......................................................................458

Marbled Eelpout to Banded Gunnel ........................................................................463

Marbled Eelpout (Lycodes raridens) ..............................................................463

Arctic Eelpout (Lycodes reticulatus) ..............................................................467

Threespot Eelpout (Lycodes rossi) ................................................................471

Archer Eelpout (Lycodes sagittarius).............................................................475

Longear Eelpout (Lycodes seminudus) .........................................................479

Scalebelly Eelpout (Lycodes squamiventer) ................................................483

Estuarine Eelpout (Lycodes turneri) ..............................................................487

Blackline Prickleback (Acantholumpenus mackayi) ..................................491

Stout Eelblenny (Anisarchus medius) ...........................................................497

Bearded Warbonnet (Chirolophis snyderi) ...................................................502

Fourline Snakeblenny (Eumesogrammus praecisus) .................................506

Daubed Shanny (Leptoclinus maculatus) .....................................................510

Slender Eelblenny (Lumpenus fabricii) .........................................................515

Snake Prickleback (Lumpenus sagitta) .........................................................520

Arctic Shanny (Stichaeus punctatus) ...........................................................525

Banded Gunnel (Pholis fasciata) ....................................................................530

Northern Wolffish to Greenland Halibut .................................................................534

Northern Wolffish (Anarhichas denticulatus) ..............................................534

Bering Wolffish (Anarhichas orientalis) ........................................................538

Prowfish (Zaprora silenus) ..............................................................................543

Arctic Sand Lance (Ammodytes hexapterus) ..............................................548

Bering Flounder (Hippoglossoides robustus) ...............................................556

Pacific Halibut (Hippoglossus stenolepis) ....................................................561

Yellowfin Sole (Limanda aspera) ....................................................................569

Longhead Dab (Limanda proboscidea) .........................................................577

Sakhalin Sole (Limanda sakhalinensis) .........................................................582

Arctic Flounder (Liopsetta glacialis) ..............................................................586

Starry Flounder (Platichthys stellatus) ..........................................................592

Alaska Plaice (Pleuronectes quadrituberculatus) ......................................600

Greenland Halibut (Reinhardtius hippoglossoides) ....................................607

Chapter 4. Synthesis of Arctic Alaska Marine Fish Ecology

By Lyman K. Thorsteinson and Milton S. Love ...............................................................617
viii

Contents—Continued

Chapter 5. Arctic Climate Change—A Tale of Two Cods

By Deborah A. Reusser, Melanie L. Frazier, Rebecca A. Loiselle, Henry Lee II, 

and Lyman K. Thorsteinson................................................................................................659

Chapter 6. Conservation of Arctic Alaska’s Marine Fish Resources

By Lyman K. Thorsteinson..................................................................................................679

Chapter 7. Glossary of Ecological Terms...........................................................................................691

Chapter 8. References Cited ................................................................................................................699

Appendixes ................................................................................................................................................739

Appendix A. Ongoing and Recently Completed Fishery Studies Funded by Bureau of 

Ocean Energy Management in Arctic Outer Continental Shelf 

Lease Areas ........................................................................................................739

Appendix B. Age-At-Size and Length-Weight Relationships for Arctic 

Marine Fishes .....................................................................................................741

Appendix C. Models Evaluated for Simulating Effects of Climate Change on the 

Distributions and Abundances of Arctic Cod and Saffron Cod in the 

Eastern Bering Sea ............................................................................................763
ix

Conversion Factors

[Inch/Pound to International System of Units]

Multiply By To obtain

Length

foot (ft) 0.3048 meter (m)

mile (mi) 1.609 kilometer (km)

[International Systems of Units to Inch/Pound]

Multiply By To obtain

Length

centimeter (cm) 0.3937 inch (in.)

millimeter (mm) 0.03937 inch (in.)

meter (m) 3.281 foot (ft) 

kilometer (km) 0.6214 mile (mi)

Area

square meter (m2

) 0.0002471 acre 

square kilometer (km2

) 247.1 acre

hectare (ha) 0.003861 square mile (mi2

) 

square kilometer (km2

) 0.3861 square mile (mi2)

Flow rate

cubic kilometer per year (km3

/yr) 0.000811 acre-foot per year (acre-ft/yr) 

kilometer per day (km/d) 0.6214 mile per day (mi/d)

Mass

gram (g) 0.03527 ounce, avoirdupois (oz)

kilogram (kg) 2.205 pound avoirdupois (lb)

Density

kilogram per hectare (kg/ha) 0.06242 pound per cubic foot (lb/ft3

) 

gram carbon per square meter

(g C/m3

)

62.4220 pound per cubic foot (lb/ft3

) 

Temperature in degrees Celsius (°C) may be converted to degrees Fahrenheit (°F) as follows:

°F = (1.8 × °C) + 32. 

Temperature in degrees Fahrenheit (°F) may be converted to degrees Celsius (°C) as follows: 

°C = (°F – 32) / 1.8.

Datum

Horizontal coordinate information is referenced to the North American Datum of 1983 (NAD 83).
x

Abbreviations

ACW Alaska Coastal Water

ADFG Alaska Department of Fish and Game

AFGP antifreeze blycoprotein

AFP antifreeze protein

AFS American Fisheries Society

AFS-ASIH American Fisheries Society-American Society of Ichthyologists and Herpetologists

AFSC Alaska Fisheries Science Center

AI Aleutian Islands

AIC Akaike information criterion

AMAP Arctic Monitoring and Assessment Programme

AO Arctic Oscillation

AW Anadyr Water

AOOS Alaska Ocean Observing System

ArcOD Arctic Ocean Diversity

BSW Bering Sea Water

BOEM Bureau of Ocean Energy Management

C carbon

Cl- chloride

CO2 carbon dioxide

CoML Census of Marine Life

CPUE catch-per-unit-effort

DBO Distributed Biological Observatory DIDSON. Dual Frequency Identification Sonar

DOI U.S. Department of the Interior

EBSA Ecological and Biologically Sensitive Area

eDNA environmental DNA

EIS Environmental Impact Statement

EEZ U.S. Exclusive Economic Zone

EBS Eastern Bering Sea 

EPA U.S. Environmental Protection Agency

FL fork length

GAM Generalized Additive Model

GI ganodosomal index

GOA Gulf of Alaska

IARPC Interagency Arctic Research Policy Commission

IDW Inverse Distance Weighted interpolation

ITIS Integrated Taxonomic Information System

LME large marine ecosystem

M natural mortality

Ma million years

MCA Metabolic Cold Adaptation
xi

MMS Minerals Management Service

Na

+ sodium

NEPA National Environmental Policy Act

NOAA National Oceanic and Atmospheric Administration

NPFMC North Pacific Fisheries Management Council

NPRB North Pacific Research Board

NSF National Science Foundation

OA ocean acidification

OBIS Ocean Biogeographic Information System

OCS Outer Continental Shelf

OR1 Online Resource 1

psu practical salinity unit

RACE Research Assessment and Conservation Engineering

rm intrinsic rate of increase

RUSALCA Russian-American Long-term Census of the Arctic

SDM species distribution model

SL standard length

SST sea surface temperature T Trophic Index

TL total length

tm age at first maturity

UAF University of Alaska, Fairbanks

USCGC U.S. Coast Guard Cutter

USGS U.S. Geological Survey

VBGM von Bertalanffy growth model

WFRC USGS Western Fisheries Research Center

YOY young-of-the-year

Abbreviations—Continued

Abstract

The marine fishes in waters of the United States north of 

the Bering Strait have received new and increased scientific 

attention over the past decade (2005–15) in conjunction with 

frontier qualities of the region and societal concerns about 

the effects of Arctic climate change. Commercial fisheries 

are negligible in the Chukchi and Beaufort Seas, but many 

marine species have important traditional and cultural values 

to Alaska Native residents. Although baseline conditions 

are rapidly changing, effective decisions about research and 

monitoring investments must be based on reliable information 

and plausible future scenarios. For the first time, this 

synthesis presents a comprehensive evaluation of the marine 

fish fauna from both seas in a single reference. Although 

many unknowns and uncertainties remain in the scientific 

understanding, information presented here is foundational with 

respect to understanding marine ecosystems and addressing 

dual missions of the U.S. Department of the Interior for 

energy development and resource conservation. This chapter 

describes the Department’s information need with respect 

to planned offshore oil and gas development, provides an 

overview of the Alaska Arctic Marine Fish Ecology Catalog, 

and presents regional background information, synthesis 

methodologies, and definitions for commonly used terms and 

concepts throughout the report. 

Introduction

Accurate natural resource inventories and knowledge 

of baseline conditions for Arctic resources and ecosystems 

are essential for estimating effects of the U.S. Department of 

the Interior (DOI) offshore oil and gas-leasing program. The 

exploration and development of these offshore resources is 

managed by the DOI Bureau of Ocean Energy Management 

(BOEM). This document, Alaska Arctic Marine Fish Ecology 

Catalog, addresses a specific need of BOEM to inform its 

decision making with respect to the fisheries ecosystems in 

Arctic Outer Continental Shelf (OCS) Planning Areas. This 

report is a compilation of species accounts of the marine 

fishes known from United States sectors of the Chukchi and 

Beaufort Seas. Certain diadromous (for example, Pacific 

salmon, char, and whitefishes) species are treated as marine 

fishes (McDowall, 1987) because much of their life cycle 

is in marine and brackish environments. This synthesis of 

information is meant to provide current information and 

understanding of this fauna and its relative vulnerability to 

changing Arctic conditions.

The National Environmental Policy Act (NEPA) requires 

the BOEM to prepare an Environmental Impact Statement 

(EIS) for each offshore oil and gas lease sale area offered 

through the DOI leasing program. The EISs provide an 

assessment of the potential environmental effects associated 

with major development proposals and communicate this 

information to decision-makers and the broader public (Wood, 

2008). In complying with the NEPA, the BOEM relies on 

the best available science to evaluate potential effects on 

regional ecosystems and living resources. Our objectives 

were to review and synthesize current fish biology, ecology, 

and fisheries information to assist BOEM’s NEPA analysts in 

assessing environmental effects from planned future offshore 

oil and gas development on the Arctic OCS. 

BOEM’s NEPA analysts require detailed information 

about the biodiversity, life history, and population ecology 

of regional biota to assess adversity of impact in the EIS 

process. For Arctic marine fishes, impact is evaluated in 

terms of potential declines in abundance or changes in 

geographic distribution and recovery of populations to preimpact status (thresholds of significance analysis, see Miner 

and Rivasplata, 1994, and Musick, 1999). To illustrate, 

adverse effects would require three of more generations for a 

population to recover (Bureau of Ocean Energy Management 

Regulation and Enforcement, 2011). The population 

understanding required for a robust EIS analysis is similar 

to what is required to assess Essential Fish Habitat under 

provisions of the Magnuson-Stevens Fishery Conservation and 

Management Act. 

Chapter 1. Alaska Arctic Marine Fish Ecology Catalog—

Chukchi and Beaufort Seas

By Lyman K. Thorsteinson1

1

U.S. Geological Survey.
2 Alaska Arctic Marine Fish Ecology Catalog

The compilation of species information provides BOEM’s 

NEPA analysts and others with a single, authoritative scientific 

reference about the biology and ecology of marine fishes in 

the United States Chukchi and Beaufort Seas. Biodiversity 

is broadly assessed with respect to current understanding of 

(1) classification and taxonomy, (2) abundance and geographic 

distribution, and (3) life history and population ecology. 

A focus of this study has been on new information and 

discoveries about this fauna since the publication of Fishes of 

Alaska (Mecklenburg and others, 2002). As such, the species 

checklist, geographic distribution maps, and depth profiles 

represent new biodiversity products to science. This synthesis 

is unique because it is based on confirmed species occurrences 

in United States sectors of the Chukchi and Beaufort Seas. 

Confirmation is an important process because it assures the 

reliability of resource information in management areas of 

direct concern to decision makers. For most species, the link 

between occurrence and abundance remains tenuous given the 

nature of sampling conducted to date. As such, quantitative 

aspects of the dynamics of fish populations and their 

interactions with the Arctic marine environment are hindered 

by inadequacies of existing data and relative lack of Alaskan 

records. The emphasis on population dynamics and ecologic 

relationships in the species accounts provides an important 

basis for assessment of outstanding needs.

Although knowledge about the marine fish fauna in 

the Alaska Arctic is among the poorest in the state, it has 

been slowly improving over time. However, much life 

history information presented in this report was acquired 

from scientific observations outside Alaska. As such, basic 

taxonomic science and population understanding is needed 

to support modern assessments and potential fisheries in the 

Arctic high seas. The goals of this report are to present the 

most current information about what is known about the 

marine fishes in the Arctic area of the United States with a 

special focus on geographic distributions, vertical structure, 

abundance, and life history parameters of key populations. 

The section, “Outline of Species Accounts” in chapter 3 serves 

as a users’ guide to information presented in the individual 

species descriptions.

Study Area

The primary geographical boundaries encompass Arctic 

OCS Planning Areas (Bureau of Ocean Energy Management 

(2012) in the U.S. Chukchi and Beaufort Seas (hereafter 

Chukchi and Beaufort Seas, fig. 1.1) and the U.S. Exclusive 

Economic Zone (EEZ) including the Arctic Management Area. 

The eastern boundary of the Beaufort Sea EEZ extends to the 

north and offshore of the Alaska-Canada Border. The Chukchi 

Sea extends from Point Barrow in Alaska and the Beaufort Sea 

in the east to the United States–Russia Maritime Boundary in 

the west. The Bering Strait forms the southern boundary of 

the Chukchi Sea. Some chapters, especially those addressing 

species descriptions (chapter 3) and discussing species 

diversity and possible geographic origins (chapter 4), include 

comparative information from adjacent seas and broader basin 

and ocean considerations, respectively.
Chapter 1 3

Figure 1.1. Bureau of Ocean Energy Management administrative boundaries for Outer Continental 

Shelf oil and gas leasing in the Chukchi and Beaufort Seas, Alaska.

tac14-5222_fig01-01

200

200

50

200

50

50

50

50

Noatak R

Kobuk R

Selawik R

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Great

Bear

Lake

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)
4 Alaska Arctic Marine Fish Ecology Catalog

Environmental Setting 

Physical and biological components of the Arctic 

marine environment as they relate to marine regional 

descriptions of fish habitats and their use by fish and other 

biota were described by DeGange and Thorsteinson (2011)2. 

The descriptions of physical oceanography, substrates, and 

biological productivity supporting marine fish habitats and 

populations are primarily based on this review. Information 

about the distribution of Arctic marine birds and mammals, 

potential competitors and predators of marine fishes, 

is not reported here but is available in DeGange and 

Thorsteinson (2011).

Regional Oceanography

Large-scale features of the Chukchi and Beaufort Seas 

oceanography were reviewed by Weingartner and others 

(2008). Surface circulation in the Beaufort Sea is dominated 

by the southern edge of the perpetual clockwise Beaufort Gyre 

of the Canadian Basin (fig. 1.2). The subsurface Beaufort 

Undercurrent Atlantic water masses and flows in the opposite 

direction, to the east, over the shelf (fig. 1.2). Bering Sea 

waters generally follow topography, moving north across 

the Chukchi Sea shelf and to the east over the shelf edge and 

slope (fig. 1.2). Currents in the shallower waters of the inner 

Beaufort Sea Shelf (fig. 1.2) primarily are wind driven and, 

thus, can flow either east or west. Because the predominant 

wind direction during the summer ice-free season is from the 

east, near-shore flow generally is from east to west. 

Oceanographers have observed regional, seasonal, and 

interannual variability in water mass properties in the Chukchi 

and Beaufort Seas. Generally, the temperature and salinity 

characteristics of the major water masses are: 

• Alaska Coastal Water—warm (2–13 °C), low salinity 

(to 32.2 practical salinity units [psu]); 

• Bering Sea Water—warm (>0.0 °C); well-mixed and 

moderate salinity (about 32.5 psu; 

• Anadyr Water—cold (<0.0 °C), marine salinities 

(32.8–33.2 psu); 

• Siberian coastal waters—cold (<0.0 °C), low salinity 

(24 psu); 

• Atlantic Water (at depths > 220 m)—warm (>0 °C), 

highly salinity (>34 psu); and 

• Beaufort Gyre—cold (<0.0 °C), moderate to high 

salinity (31–34 psu). 

In the Chukchi Sea, the summer water masses are cold in the 

east and warm in the west. Similarly, warm brackish waters 

(5–10 °C, <15 psu) occur adjacent to Alaska’s north coast 

during summer (to about 10 km off the coast) and cold marine 

waters (<1 °C, 28 psu) farther offshore.

Under persistent east winds, bottom marine water 

can move onshore, where it is forced to the surface. This 

upwelling of marine water can cause some otherwise brackish 

and warm areas along the coast to become colder and more 

saline. This replacement of brackish with marine waters results 

in the transport of warmer, less saline waters offshore.

The Chukchi Sea receives water that flows northward 

through the Bering Strait, driven by the 0.5 m drop in sea level 

between the Aleutian Basin of the Bering Sea and the Arctic 

Ocean (fig. 1.2). Coachman and others (1975) provide a good 

overview of the northward movement of Bering Sea waters 

into the Chukchi Sea. Three distinct water masses, each of 

different origin, move northward through the Bering Strait. 

Anadyr Water, cold, high salinity, nutrient-laden oceanic 

water that originates along the slope of the Bering Sea Shelf, 

flows northward through Anadyr Strait, west of St. Lawrence 

Island and into the central Chukchi Sea (fig. 1.2). As much 

as 72 percent of the water transported through the Bering 

Strait in the summer may come through Anadyr Strait. 

Alaska Coastal Water originates in the Gulf of Alaska. This 

low salinity, seasonally warm water hugs the Alaska coast 

as it transits the Bering Sea into the Chukchi Sea. Alaska 

Coastal Water is influenced by freshwater run-off from major 

rivers in western Alaska. Bering Shelf Water is the resident 

water mass of the central shelf region south of St. Lawrence 

Island. This water mass is intermediate with respect to its 

hydrographic properties when compared to Anadyr Water and 

Alaska Coastal Water, is advected northward on both sides of 

St. Lawrence Island, and then flows through the Bering Strait 

where it mixes with the other water masses. These waters are 

an important source of plankton and carbon in the Chukchi 

and Beaufort Seas, and influence the seasonal distribution and 

abundance of marine biota and migration behaviors of many 

species (Piatt and Springer, 2003; Hopcroft and others, 2008; 

Weingartner and others, 2008; Crawford and others, 2012). 

The deep waters offshore in the northern Chukchi Sea also are 

a potentially important source of nutrient-rich waters. 

2

This review of large-scale physical oceanography was guided by DeGange 

and Thorsteinson (2011) who presented information previously reported in 

Minerals Management Service (2008) and Weingartner and others (2008). The 

summaries of primary and secondary levels of ecosystem production are from 

Bluhm and others (2008), Hopcroft and others (2008), Stockwell and others 

(2008), and Yager and others (2008). Other key references included Carmack 

and Wassmann (2006); Grebmeier and Maslowski, 2014; Moore and Stabeno, 

2015; and Grebmeier and others (2015).
Chapter 1 5

Bering Sea Cold Pool

Mecklenburg and others (2011) described the boundaries 

of an “arctic region” to include the Arctic Ocean and its seas 

and adjacent waters of the North Atlantic and North Pacific 

southward to an ichthyofaunal boundary (Arctic–Boreal 

species in chapter 2). Mecklenburg and others (2011, p. 110) 

reported that, “...position of the boundary is reflected in 

major differences in species composition associated with 

seafloor topography, such as sills and canyons, and water 

characteristics, such as temperature and salinity, that form 

barriers against fish movements.” In the northern Bering 

Sea, the location of the 2 °C isotherm during summer 

months, generally south of St. Lawrence Island, demarks this 

zoogeographic boundary and the northern edge, or ecotone, 

of marine waters associated with the Bering Sea cold pool 

(bottom temperatures ≤ 2 °C) that relates to the seasonal 

presence of sea ice (see chapters 4 and 5).

Sea Ice Dynamics 

The presence of ice in the Arctic is one of the most 

important physical conditions to deal with for developing 

offshore OCS oil and gas resources. The seasonal sea ice 

cycle is a pervasive force in the Arctic, influences human 

activities and many aspects of the region’s natural history, 

and demonstrates great seasonal and inter-annual variability 

off the coast of Alaska. Generally, there are two forms of sea 

ice: fast ice that is anchored along the shore and free-floating 

pack ice that moves with winds and currents. Shore fast and 

pack ice interact to cause an extensive, somewhat predictable, 

system of flaw leads (swathes of open water in between ice) 

and polynyas off the coasts of the Chukchi and Beaufort Seas 

eastward to the Canadian Archipelago. These flaw leads and 

polynyas become more prevalent in spring and are biologically 

significant features with respect to the timing and location of 

seasonal movements and northward migrations of wildlife 

species, such as bowhead whales and marine birds. 

Figure 1.2. Schematic circulation map of the northern Chukchi Sea and western Beaufort Sea. From Weingartner and 

others (2008). 

tac11-5179_fig2-01

0

4,500

1,000

250

100

50

20 Beaufort Gyre

Atlantic Water (subsurface)

Alaska Coastal Water

Bering Sea Shelf Water

Anadyr Water

Siberian Coastal Current

Depth of water, in meters
6 Alaska Arctic Marine Fish Ecology Catalog

Maximum sea-ice cover occurs in March or early 

April, lagging minimum insolation in late December by 

3 months because of the heat capacity of the ocean and the 

cold atmosphere. During these months, essentially all of the 

Beaufort and Chukchi Seas are ice-covered (fig. 1.3). Winter 

ice extent in the Arctic has decreased since the late 1970s 

(fig. 1.3) along the southern margins of sea-ice extent, but 

not severely (fig. 1.4). Maximum retreat of the sea ice occurs 

in September, again lagging maximum insolation by about 

3 months. The extent of sea-ice loss in Septembers since 

the satellite record began has been remarkable (fig. 1.5). 

By September, in normal years, the ice pulls away from the 

Arctic coasts of Canada, Alaska, and Siberia, leaving a nearly 

continuous, relatively ice-free corridor that varies in width 

around the permanent ice pack. In recent years (2010–15), 

the ice-free corridor has expanded to hundreds of kilometers 

in the East Siberian Sea and offshore of the northern Alaska 

coast. The contrasts between 1980, a representative year with 

extensive ice cover, and 1987, when sea-ice extent in the 

Arctic was at a record minimum, and the long-term median ice 

edge are evident (fig. 1.5). 

In addition to decreases in sea-ice extent during late 

summer and autumn, the amount of multi-year compared 

with summer sea ice in the Arctic also is changing; conditions 

are moving toward an abundance of younger and thinner sea 

ice (figs. 1.3–1.6). The longer sea ice remains in the Arctic 

Ocean the thicker it becomes because of additional freezing 

and deformation. The thinning of Arctic Ocean sea ice has 

occurred largely because of the export of older, thicker sea ice 

out of the Arctic through Fram Strait, east of Greenland. This 

is important because young, thin ice is more vulnerable to the 

melting that results from warm air and water temperatures, 

perpetuating a feedback cycle because the open ocean absorbs 

solar insolation.

Figure 1.3. Extent of sea ice for single months and single years, using 1980 as an example of an extensive ice cover year, and 2007 as 

the record minimum year—maximum winter extent. The border plots the long-term median ice edge based on data from 1979 to 2000. 

From National Snow and Ice Data Center, 2011a, monthly sea ice extent for 1980 and 2007: National Snow and Ice Data Center database, 

accessed April 15, 2011, at http://nsidc.org/.
Chapter 1 7

Figure 1.4. Sea-ice extent anomalies for March (maximum sea-ice extent) and September (minimal sea-ice extent) 

expressed as percentage-departure from average (that is, anomalies as compared to the 1979–2000 mean). From National 

Snow and Ice Data Center (2011b).

Figure 1.5. Extent of sea ice for single months and single years, using 1980 as an example of an extensive ice cover year, and 2007 as 

the record minimum year. The border plots the long-term median ice edge based on data from 1979 to 2000. Source: National Snow and 

Ice Data Center (2011a).
8 Alaska Arctic Marine Fish Ecology Catalog

Seafloor Substrates

Soft sediments dominate the sea floors of the continental 

shelves of the Beaufort and Chukchi Seas. These are largely 

combinations of muds, sands, and gravels (fig. 1.7). These 

soft-sediment bottoms support high densities and biomass of 

benthic invertebrates, particularly in the extensive shallow 

shelf areas of the Chukchi Sea where productivity is high 

(for example, Hanna Shoal and Barrow Canyon [fig. 1.1]). 

Only two areas with hard substrates have been identified in 

the entire region (Smith, 2010) (fig. 1.7)—one in Peard Bay, 

southwest of Barrow, and the other in Steffanson Sound near 

Prudhoe Bay that is known as the “boulder patch” (Dunton 

and others, 1982). The boulder patch is characterized as 

sediment with greater than 10 percent boulder cover. It 

provides attachment habitat for the endemic kelp Laminaria 

solidungula and other macroalgae, which are the primary 

carbon source for consumers living there.

Figure 1.6. Old and new ice in the Arctic for February. These maps show the median age of February sea ice from (A) 1981 to 2009 and 

(B) February 2009. As of February 2009, ice older than 2 years accounted for less than 10 percent of the ice cover. Dark blue indicates 

ice greater than 2 years old; medium blue indicates 2-year-old ice; pale blue indicates annual ice. From National Snow and Ice Data 

Center (2011c), accessed April 15, 2011, at http://nsidc.org/sotc/sea_ice.html.
Chapter 1 

9

tac11-5179_fig2-06

A R C T I C O C E A N

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Bay

Camden

Bay

Bering Strait

Kolyuchin

Bay

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Mackenzie

Bay

Smith

Bay Kugmallit

Bay

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Peard

Bay

Wrangel Island Herald Island

St. Lawrence Island

R U S S I A

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Old Crow

Inchoun R.

Mechigmensk

Bay

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Bay

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Cape Lisburne

Cape

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Icy Cape

Kasegaluk

Lagoon

Point

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R

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Tanana R.

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Emmonak

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Arctic

Village

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Point

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Wainwright

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Pass

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128°0'0"W 132°0'0"W 136°0'0"W 140°0'0"W 144°0'0"W 148°0'0"W 152°0'0"W 156°0'0"W 160°0'0"W 164°0'0"W 168°0'0"W 172°0'0"W 176°0'0"W 180°0'0" 176°0'0"E

72°0'0"N70°0'0"N68°0'0"N66°0'0"N64°0'0"N62°0'0"N

Arctic Marine Synthesis:

Atlas of the Chukchi and Beaufort Seas

150 0 150 300 75

Miles

January 2010

Sea Floor Substrate

Sources: (1) NOAA. 1988. Bering, Chukchi, and Beaufort seas coastal and ocean zones strategic assessment

data atlas; (2) Carmack, E.C., and R. MacDonald. 2002. Oceanography of the Canadian shelf of the Beaufort

Sea: a setting for marine life. Arctic 55:29-45; (3) Dunton, K.H., E. Reimnitz, and S. Schonberg. 1982. An

Arctic kelp community in the Alaskan Beaufort Sea. Arctic 35(4):465-484; (4) NOAA. 2002. Environmental

sensitivity index, version 3.0. NOAA, Seattle, Washington; (5) Mohr, J.L., N.J. Wilimovsky, and E.Y. Dawson.

1957. An arctic Alaskan kelp bed. Arctic 19:45-54; (6) Horowitz, W.L. 2002. Evaluation of sub-sea physical

environmental data for the Beaufort Sea OCS and incorporation into a geographic information system (GIS)

database. OCS Study MMS 2002- 017. <http://www.mms.gov/ itd/pubs/2002/2002-017/

User_Manual_contract30985.pdf>. Accessed June 2009.

Boulder PatchNo Data

3 4 5 6

Gravel1 2Gravelly Mud1Gravelly Muddy Sand1Gravelly Sand1Mud1Muddy Gravel1Muddy Sand1Sand1Sandy Mud1

This map is based on the best available data, which varies in quality across the project area. For more information on

data quality and knowledge gaps, see the introduction to the Arctic Marine Synthesis and the description of this map.

Clay2Silt2

Figure 1.7. Seafloor substrates of the Alaska Beaufort and Chukchi Seas. From Smith (2010).
10 Alaska Arctic Marine Fish Ecology Catalog

Benthos

Benthic food supply originates in surface waters and is 

highly seasonal in the Arctic. Densities of sediment particles 

and their nutritional values range vastly from the nutrient 

rich waters of the northern Bering and Chukchi Seas to the 

oligotrophic deep waters of the Arctic Basins. Less is known 

about the benthos from the Beaufort Sea although preliminary 

data from ongoing BOEM surveys suggest there is much less 

biomass and diversity, and that different ecological processes 

occur (appendix A). Generally, however, comparisons of 

energy fluxes show that the benthic systems receive more 

energy in the Arctic than from temperate and tropical systems.

Much of the broad, shallow shelf of the Chukchi Sea 

is strongly influenced by northward flowing nutrient-rich 

Pacific Ocean water through the Bering Strait, resulting in 

high benthic biomass, which is among the highest worldwide 

in soft-sediment macrofaunal communities (for example, 

Grebmeier and Maslowski, 2014; fig. 1.8). Specifically, the 

south-central Chukchi Sea has the highest algal and faunal 

biomass on the combined Bering Sea and Chukchi Sea shelf 

because of the high settlement rates of organic production that 

is not grazed by microbes and zooplankton. These rich benthic 

communities, tied to high pelagic production and advection, 

serve as prey for various diving sea birds and marine 

mammals, a key feature of the productive Chukchi Sea. 

About 1,200 species are known from the Chukchi Sea fauna 

to date with amphipods, clams, and polychaetes dominating 

infaunal community. Important macrofauna prey species for 

higher trophics include bivalves taken by walrus, in particular 

Macoma spp. and Mya truncata, and benthic amphipods 

used by gray whales and bearded seals. Within the epifauna, 

ophiuroids dominate abundance and biomass in much of 

the surveyed Chukchi Sea, and other patchily distributed 

echinoderms (especially asteroids), gastropods, ascidians, 

sponges, cnidarians, and bryozoans are locally abundant.

The comparatively narrow Beaufort Sea Shelf is 

influenced by large freshwater inflow from numerous small 

rivers and streams, the larger Colville and Mackenzie Rivers 

(fig. 1.1), and permafrost resulting in estuarine conditions in 

the nearshore. Because of this freshwater flow, non-marine 

sources of carbon may play an increasingly important role 

for the benthic food web in parts of the nearshore Beaufort 

Sea. The Beaufort Sea floor is dominated by soft sediments 

(fig. 1.7), but high ice cover and associated scouring, along 

with glacial erratics, have left coarser sediments (gravel and 

boulders) in various areas of the Beaufort Sea. The Alaskan 

part of the Beaufort Sea coast is fringed by sandy barrier 

islands forming numerous shallow lagoons with average 

depths less than 5 m and ecological traits different from those 

in the open water. Compared to the Chukchi Sea, productivity 

and benthic biomass in the Alaskan Beaufort Sea are 

dramatically lower. Consequently, benthic-pelagic coupling is 

not as pronounced as in the Chukchi Sea and food chains are 

shorter. Much less is known about the slopes of the Chukchi 

Sea and especially the Beaufort Sea, and the adjacent basins 

(Bluhm and others, 2008). The existing investigations of the 

slopes and abyssal infaunal benthos in the western Arctic 

revealed low abundances and biomass values relative to the 

shelves, especially with increasing water depth and distance 

from the shelves. At taxonomic levels of phylum and orders, 

the soft-bottom deep Arctic macrofauna appear to be similar to 

the shelf communities: polychaetes, bivalves, and crustaceans 

are dominant, but on a family, genus, and species level, 

inventories differ from the shelves.
Chapter 1 11

tac11-5179_fig 02-08

Figure 1.8. Benthic biomass distribution in the Chukchi and northern Bering Seas. From Grebmeier and 

others, 2006a. Feder and Stoker station locations reference historical sampling stations by University of 

Alaska-Fairbanks researchers in the Bering and Chukchi Seas as part of NOAA’s Outer Continental Shelf 

Environmental Assessment Program. (g C m-2, grams of carbon per square meter.)
12 Alaska Arctic Marine Fish Ecology Catalog

Primary and Secondary Production

In the Arctic, the combination of cold temperature, the 

occurrence of sea ice, and the extreme seasonal variations in 

light regimes controls phytoplankton growth and governs its 

spatial and temporal growth patterns. The stabilizing effect 

of sea ice allows production to occur near the surface under 

low light intensities. A large number of planktonic algae 

thrive in Arctic waters, but there seem to be relatively few 

truly Arctic species. Estimates of phytoplankton biomass 

vary widely depending on the region, with the highest 

amounts in the Chukchi Sea. Algal production and biomass 

in the Arctic primarily are controlled by light, stratification, 

and nutrient fields. On the shelves, advection and turbulent 

mixing of nutrients through the Bering Strait and local 

nutrient remineralization sustain extremely high primary 

production values on the Chukchi Sea Shelf (fig. 1.9). Much 

of the production is not grazed, falls to the sea floor, and fuels 

benthic communities. In addition to phytoplankton, ice algae 

contribute to the total primary production of the Arctic Ocean 

with higher production values in first-year ice compared to 

multi-year ice. The contributions of ice algae to total primary 

production range from less than 1 percent in coastal regions to 

as much as 60 percent in the central Arctic Ocean.

Secondary producers include the microbes, protists, 

and zooplankton that consume phytoplankton and algae. 

Compared to phytoplankton and mesozooplankton, much less 

is known about the composition, distribution, and rates of 

activity of microbes and protists in the Arctic Ocean, and this 

confounds the ability to estimate the effect of climate change 

or other disturbances on food webs and basic biogeochemical 

processes. Biomass of heterotrophic microbes in Arctic 

surface waters shows a strong response to seasonal changes 

in phytoplankton stocks. In the Chukchi Sea, concentrations 

of bacteria are low in spring, increase over the course of the 

bloom, and are highest in late summer. Heterotrophic protists 

include nanoflagellates, ciliates, and dinoflagella.

Recent work in the Gulf of Alaska, the Bering Sea, 

and shelf and slope regions of the western Arctic Ocean 

has confirmed the role of these organisms, known as 

microzooplankton, as consumers of phytoplankton in 

sub‐Arctic and Arctic food webs. Although it is likely that 

phytoplankton and sea ice algae still represent a crucial food 

source for the larger zooplankton, use of microzooplankton 

as food is recognized as being of similar import, particularly 

during periods when phytoplankton standing stock is low 

or of poor quality. Because strong local pulses of primary 

production are a frequent characteristic of high-latitude 

oceans, including the Chukchi and Beaufort Seas, the 

response of microbes (including both bacteria and protists) to 

these pulses determines the rate of remineralization and the 

fraction of total production exported to the benthos. Weak 

microbial activity in the Arctic contributes to the high degree 

of bentho-pelagic coupling in many shelf regions of the Arctic 

and the consequent strength of demersal ecosystems.

Zooplanktons are the major grazers of the primary 

production in the Arctic and determine the resources available 

to many higher trophic levels, such as fishes, seabirds, and 

marine mammals. In the Chukchi Sea, large quantities of 

Pacific zooplankton enter the region through the Bering 

Strait, in a complicated mixture of water masses. The influx 

of the rich Pacific water determines the reproductive success 

of both the imported and resident zooplankton communities. 

Both inter-annual and long-term variation in climate will 

affect the relative transport of these various water masses 

and hence the composition, distribution, standing stock, and 

production of zooplankton and their predators in the Chukchi 

Sea. Zooplankton abundance and community structure also 

affect the amount and quality of carbon exported to the benthic 

communities in this region. In contrast, the Beaufort Sea 

primarily is Arctic in character, with cross-shelf exchange 

mechanisms more important in determining the relative 

contribution of “oceanic,” “shelf,” and “estuarine” species. 

In the Eastern Beaufort Sea, the outflow of the McKenzie 

River has significant effect on both the composition of the 

zooplankton and its productivity. Thus, the Beaufort Sea is 

responding to a fundamentally different set of factors than 

the Chukchi Sea, even if they are both driven by similar 

climate-related variations and trends.

Although copepods typically predominate throughout the 

Arctic, there is a broad assemblage of other planktonic groups. 

Euphausiids are less abundant and diverse in Arctic waters 

than elsewhere, but can be important prey for higher trophic 

levels such as bowhead whales, birds, and fishes. Larvaceans 

(Appendicularians) have been shown to be abundant in Arctic 

polynyas, and are transported in high numbers through the 

Bering Strait into the Chukchi Sea. Similarly, important 

and common predatory groups, such as the chaetognaths, 

amphipods, ctenophores, and cnidarians have been reported 

on in detail by only a few surveys. Hyperiid amphipods also 

can be common in Arctic waters and, like chaetognaths, have a 

potential to graze a large proportion of the Calanus population. 

Relatively little is known of the abundance, composition, 

or ecology of the delicate gelatinous zooplankton, such as 

jellyfish. There are indications that climate change has resulted 

in increased numbers of jellyfish in the Bering Sea in recent 

years. Scientists have recorded jellyfish piled up several feet 

deep along shorelines near Barrow, Alaska. The ecological 

effect of these predators is substantial and underestimated in 

polar waters.

The ongoing reduction of the sea-ice cover will have 

major effects on the ecosystems and biogeochemical fluxes on 

the extensive continental shelves of the Arctic Ocean. Many 

processes involved in the regulation of the vertical and trophic 

fluxes of particulate organic carbon, and the production of 

dissolved organic carbon, are controlled by the zooplankton. 

Knowledge of zooplankton community ecology, especially 

the temporal and spatial distribution patterns of the different 

classes of zooplankton, is needed to understand the role of 

sea‐ice variability in dictating fluxes of biogenic carbon on 

and off the shelves.
Chapter 1 13

tac11-5179_fig03-07

Figure 1.9. Contours of integrated chlorophyll a concentrations, in milligrams per square meter [mg m-2]) 

based on discrete measurements at sampling locations in the northern Bering Sea, east Siberian Sea, and 

Chukchi and Beaufort Seas, April–September 1976–2004 (Grebmeier and others, 2006a).
14 Alaska Arctic Marine Fish Ecology Catalog

Environmental Context Within the 

Pacific Arctic Region

The information summarized in the section, 

“Environmental Setting” focused on the United States 

Chukchi and Beaufort Seas, and included relevant physical 

and biological information from the Pacific Arctic Region. The 

description of the Pacific Arctic Region includes the northern 

Bering Sea, Chukchi and Beaufort Seas, East Siberian Sea, 

and western sectors of the Canadian Arctic Archipelago (for 

example, Grebmeier and others, 2015). Regional information, 

from data collections outside the United States, was included 

because large scale understanding of physical and biological 

processes (for example, the delineation of boundaries for 

the Arctic Large Marine Ecosystem [Protection of the 

Arctic Marine Environment, 2013] evaluates interactions 

and connections at scales of the northeast Pacific and 

Arctic Ocean) is needed to address historical and ecological 

biogeography objectives of this study (chapter 4). Additional 

focus and synthesis on the environmental features of the 

Pacific Arctic Region address scale issues by drawing special 

attention to the importance of shelf geography and advection 

on regional ecology (this chapter) as it pertains to marine fish 

distributional patterns described in chapters 3 and 4.

Geographic relations between advection, sea ice, and 

stratification properties were reviewed by Carmack and 

Wassmann (2006) to classify shelf typologies and compare 

regional connections between physical habitats and biota 

in Arctic seas. Inflow, interior, and outflow shelves were 

described for the Pacific Arctic Region and collectively 

these shelf types were estimated to cover 2,145 × 103

 km2

or approximately 35 percent of the entire shelf area of the 

Arctic Ocean (6,048 × 103 km2). Continental shelf area is 

greatest in the East Siberian Sea (16 percent of total Arctic 

Ocean shelf area) followed by the Chukchi Sea (10 percent), 

Beaufort Sea (3 percent), and Canadian Arctic Archipelago 

(about 2 percent). The total volume of seawater over Arctic 

shelves is 829 × 103 km3 of which 7 percent is associated with 

the East Siberian Sea, 6 percent the Chukchi Sea, about 4–5 

percent in the Canadian Arctic Archipelago, and 3 percent the 

Beaufort Sea.

The northern Bering/Chukchi Sea is a shallow inflow 

shelf (mean depth = 58 m) as the transport of Pacific water 

through the Bering Strait connects Pacific and Arctic Oceans 

(S ~ 32 psu, VT ~ 1 Sv) (fig. 1.2; salinity and annual mean 

volume as reported by Woodgate and others, 2015). As these 

waters flow across the Chukchi shelf they are responsive to 

atmospheric forcing and are strongly modified by geophysical 

and physical processes before their subduction along the 

break and upper slope. Freshwater transport from rivers, ice 

(primarily first year ice), and stratified waters of the Bering 

Sea is significant and supports strong seasonal stratification 

and high primary productivity in the Chukchi Sea. The Bering 

Sea inflows, especially from upwelled deep waters, are rich 

in nutrients and biogenic materials such as phytoplankton, 

zooplankton, and meroplankton. Large herbivores are not 

abundant in the zooplankton community overlying shelf 

waters and the grazing efficiency is low. The advection of 

freshwater and suspended organic matter combined with high 

primary production and inefficient cropping over shallow 

depths results in tight pelagic-benthic coupling and high 

standing stocks of benthic communities (Renaud and others, 

2008). As a consequence of the large vertical flux of locally 

and advected carbon, the Chukchi shelf (and southwestern 

Beaufort Sea) are also the site of substantial denitrification and 

high benthic biomass in offshore areas compared to the more 

river-influenced central and eastern Beaufort Sea. Benthic 

ecosystems and short food chains are adaptations to the fluxes 

and cycling of carbon as evidenced in the composition and life 

history traits of the benthos and seasonal migrations of large 

numbers of benthic feeding animals in the Bering Strait and 

Chukchi Sea each year.

The East Siberian Sea (mean depth = 48 m) and Beaufort 

Sea are interior shelves (mean depth = 80 m). According to 

Carmark and Wassman (2006), interior shelves are shallow 

and characterized by strong effect of major Arctic rivers (for 

example, Kolyma and Mackenzie Rivers). Interior shelves 

exhibit a positive estuarine flow (plume spreading) in summer 

and a negative estuarine circulation (brine release) in winter. 

During summer, sediments and organic matter are transported 

to the nearshore in a highly variable mix of riverine, estuarine, 

and marine waters. The horizontal exchange of water masses 

is thus considerable and variation in salinity is significant. 

Carmack and others (2015) defined the brackish water 

mass that forms near the coast (≤30 m) as the River Coastal 

Domain. In deeper waters beyond the shelf break, boundary 

currents flow to the east and contain Arctic water derived from 

Atlantic and Pacific origins. Interior shelf waters are turbid 

(large terrigenous loads) and seafloors featureless except in 

large river areas where small valleys continue across the bed. 

The interior shelves are characterized by a predominance of 

landfast ice that melts during summer and flaw polynyas in 

winter. Primary production and the general biological activity 

are low over offshore areas of the interior shelves due to high 

turbidity, export of surface waters below the ice cover, and 

nutrient limitation due to stratification. Suspended biomass 

of planktonic organisms is thus low, but that of benthic 

organisms is relatively high. This relates to benthic foods from 

marine advection and local production and significant imports 

from littoral and riverine sources. Shoreline erosion delivers 

large amounts of peat to coastal waters and represents a major 

source of carbon in lagoonal food webs (Dunton and others, 

2005; 2006; 2012). During winter, the lagoons freeze to depths 

about 2–5 m, often to the bottom, a disruption so great that 

the epibenthic community at the base of their food webs must 

re-colonize them annually.

The Beaufort Sea is a narrow interior shelf. The primary 

production in some of the coastal lagoon ecosystems of the 

United States Beaufort Shelf is less than 10 grams of carbon 

per meter (g C m2

/yr) as light penetration in spreading and 

meandering river plumes is extremely limited. Land fast 

ice cover is persistently strong and severely limits primary 
Chapter 1 15

production. Primary production on the adjacent Canadian 

Beaufort Shelf with its mixture of land fast and multi-year ice, 

flow polynyas, stamukhi, and higher light penetration outside 

the river plumes, is high, in the range of 30 to 70 g C m2/yr. 

High freshwater inputs from numerous rivers and streams 

produce an environment that is decidedly estuarine in 

character, especially during the late spring and the summer 

months. Coastal erosion and river discharge are largely 

responsible for introducing high concentrations of suspended 

sediment from upland regions into the nearshore zone and 

often trapped in nearshore lagoons. Shelf energetics are 

characterized by a relatively low autochthonous production 

and a rich terrestrial, allochthonous carbon supply, especially 

in more productive regions. Benthic biomass is relatively rich 

in terrestrial carbon. Zooplankton populations, however, are 

poorly developed in the inner coastal region and the brackish 

waters that form along the coast each summer allows access 

to more marine foraging areas outside of the lagoons by 

several species of amphidromous fishes of importance in 

traditional economies.

In contrast the East Siberian Sea is a wide interior shelf. 

The wide interior shelves may extend far offshore (about 

800 km) before reaching the shelf break. Most of the trophic 

dynamics in this sea remain to be resolved. This sea is heavily 

covered by land fast ice, is shallow (<50 m), and is exposed 

to most of the discharge of freshwater to the Arctic Ocean 

when the Lena River is considered. As a consequence, total 

primary production is low and the contribution from ice algae 

is probably significant. Stratification is strong during the 

productive season and the biota is dominated by benthos. High 

chlorophyll a concentrations in the sediments indicate a tight 

coupling between sympagic and pelagic primary production 

and nutrient supply to the benthos throughout the entire 

Laptev Sea. However, pronounced regional differences exist 

in the magnitude of primary production. The shallow nature 

of the ecosystem implies that the effect of zooplankton on 

carbon flux is limited and may increase toward the shelf break. 

Preliminary carbon budgets indicate that a high proportion 

of primary production is channeled through the benthic food 

web. As in the case of the narrow interior shelves input of 

allochthonous organic carbon seems to be required to balance 

the overall carbon demand. Despite its refractory nature, the 

supply of riverine dissolved organic carbon and particulate 

matter of terrestrial origin, mediated by microbial food webs, 

may be significant for this shelf.

The Canadian Arctic Archipelago is a network outflow 

shelf (mean depth = 124 m). Outflow shelves allow Arctic 

Water to drain back into the North Atlantic. The Canadian 

Arctic Archipelago is a complicated network of channels, 

straits, and sounds, and water mass transit times are long 

enough for thermohaline and biogeochemical changes to 

occur en route. There are no large rivers in this system, but 

moderately sized rivers enter from the Canadian mainland; 

there is other, local freshwater run-off. Stratification derives 

primarily from ice melt. Land fast ice in the region is regular, 

but strongly variable with regard to depth, time of melt, and 

snow-cover. The archipelago is ice-covered during most of 

the year with extensive, but variable ice melt and stratification 

observed during summer and early autumn. Primary 

production on outflow shelves is spatially variable. Over the 

southern parts of outflow shelves the primary production can 

be significant and is highly seasonal, quickly nutrient limited, 

and highly variable between years. The comparatively great 

depth implies that much of the carbon flux is channeled 

through plankton, with additional supplies from sympagic 

biota. Existing measurements of total primary production in 

the entire Archipelago suggest that it may support as much 

as 32 percent of the total primary production of pan-Arctic 

shelves. The zooplankton dynamics are even more variable, 

probably because of irregular advection episodes through the 

Archipelago. Ice fauna and flora are rich in the Archipelago 

and close pelagic–benthic coupling results in rich benthic 

communities in the shallow sections. However, primary 

production is generally low and the transport of organic matter 

(mostly of terrestrial origin) and ice biota by multi-year ice 

and the Transpolar Drift is significant along longitudinal 

outflow shelves. Interannual comparisons of ice algae 

production in Barrow Strait (to the east of M’Clure Strait, 

Canada) reveal strong year-to-year variability, ranging from 

2 to 23 g C m2

 y1 (Smith and others,1988). Thus, this outflow 

shelf probably experiences similar significant variability in its 

planktonic/sympagic primary production. 

A conceptual model for the Pacific Arctic Region 

relates observed patterns in ecosystem structure and function 

to advection and related functional attributes (fig. 1.10; 

Grebmeier and others, 2015). The patterns are consistent with 

shelf processes described by Carmack and Wassman, (2006) 

denoting significance to areas of convergence in the Chukchi 

and Beaufort Seas and along the shelf break, large freshwater 

influences from the Mackenzie River and through the Bering 

Strait, and within different water masses of the Chukchi Sea. 

The conceptual model also is consistent with information 

about the physical environment and the marine resources in 

this chapter and portrays the role of advection more clearly 

with respect to seasonality and patterns in species distribution 

and abundance and further description of biological 

connections between Chukchi and Beaufort Sea shelves and 

other Pacific Arctic environments (for example, northern 

Bering Shelf and nearshore waters). 

Two other conceptual models also inform current 

understanding of the present state: (1) a food web model 

linking physical factors to biological distribution and 

abundance and food web relationships (Wiese and others, 

2013); and (2) a marine-pulsed model focusing on the 

seasonality of production cycles and biological activity as they 

relate to connections between riverine exports, benthic-pelagic 

coupling and remineralization, advection, winds and episodic 

upwelling events, and sea ice coverage (Moore and Stabeno, 

2015). The advection, food web, and marine-pulsed models 

are complementary, building on the unifying constructs of 

the contiguous domain, and depict current understanding 

and suggest how energy pathways and efficiencies in benthic 
16 Alaska Arctic Marine Fish Ecology Catalog

and pelagic communities could be affected by the changes 

in meteorology (temperature, winds and cloudiness), sea 

ice (timing and extent of open water) hydrology (freshwater 

discharge), hydrography (stratification and halocline), 

and plankton dynamics (timing, intensity, magnitude, and 

production) and related ecological efficiencies (energy 

transfers between trophic levels). 

Advection is a key forcing function for the Arctic marine 

environment in general and the Pacific Arctic Region in 

particular (Grebmeier and others, 2015). In this conceptual 

model, advection of water, ice, and biological constituents 

through the Bering Strait creates the energetic pathways and 

connections that affect much of the region and influence 

regional productivity and abundance of biota. Sea ice is 

a primary forcing factor in the region and is inherently 

connected to advection. Because of a steep decadal decline 

in seasonal sea ice coverage in the Chukchi Sea, it is thought 

to be among the most vulnerable Arctic continental seas for 

ecosystem change (Wiese and others, 2013; Grebmeier and 

others, 2015). 

Figure 1.10. Advective conceptual model for Pacific Arctic Region (emphasis on Chukchi and Beaufort Seas, Alaska. 

(From Grebmeier and others, 2015). A 30-meter contour separates nearshore and offshore regions. The inflow of Pacific 

water masses (Anadyr Water [AW], Bering Shelf Water [BSW], and Alaska Coastal Water [ACW]) is critical to ecosystem 

dynamics, for example benthic ecosystem functioning, and transport processes in this conceptual framework. The East 

Siberian Coastal Current, not shown, while important, may be of less significant to the United States Chukchi and Beaufort 

Sea marine ecosystem dynamics.
Chapter 1 17

Methods, Sources, and Terminology

Standard methods of literature review for published and 

unpublished data sources were followed. The search process 

for data and information included reviews of: current and 

historical studies; peer-reviewed journal and monographs; 

regional guides; museum specimens; technical reports and 

survey databases; searchable Web-based databases; oral 

communications; limited new data collection; and use of 

defensible unpublished scientific data. 

Unless otherwise noted we conformed with conventions 

of scientific and common nomenclature used by Nelson 

and others (2004), Nelson and Bouchard (2013), Love and 

others (2005), Mecklenburg and others (2011), Maslenikov 

and others (2013), and Mecklenburg and Steinke (2015). 

This publication includes a key list of Arctic species 

referred to as the Online Resource 1 (OR1). The American 

Fisheries Society-American Society of Ichthyologists and 

Herpetologists (AFS-ASIH) recently completed an update 

to its list of common and scientific names (Page and others, 

2013). In this report, Boreogadus saidi refers to Arctic Cod 

instead of Polar Cod (Page and others, 2013). Arctogadus 

glacialis is called Ice Cod. Other changes in nomenclature 

as they pertain to differences in taxonomy since the Fishes 

of Alaska (Mecklenburg and others, 2002) was published 

are noted. The Fishes of Alaska is a critical companion 

document to this report for the physical description of species 

including the key diagnostic features of anatomy for proper 

identifications.

Synthesis Methods

The preparation of species accounts involved several 

interrelated tasks including (1) documenting marine fish 

biodiversity; (2) defining terminology relative to the fauna 

and environment; (3) developing a standard template 

for information delivery (outline of species accounts); 

(4) acquiring and reviewing biological, ecological, and 

economic data and information; (5) producing species 

accounts; (6) synthesizing environmental (physical, biological, 

and ecological) information; and (7) managing the scientific 

review. The second and third tasks were most challenging 

because of limitations in existing data and the related need 

for consistent, reliable information in each species account. 

Pilot efforts involved the development of prototype species 

accounts and their review and improvement in an iterative 

process that introduced an evolutionary quality to product 

formation and information portrayal. The pilots highlighted 

the need for clarity in usage of standard terminologies being 

applied to Arctic baselines.

The fifth synthesis task involved an in-depth analysis 

of the collective body of the biological and ecological 

information presented in the species accounts to give it proper 

context to the biogeography of the fauna. Synthesis efforts 

were framed using historical and ecological components of 

biogeography (Nelson, 2006) to describe patterns of species 

occurrence, habitat and population relationships, and the 

ecosystem processes that affect the distribution and abundance 

of marine fishes in the Chukchi and Beaufort Seas. These 

environmental influences reflect physical, biological, and 

ecological attributes and interactions in the region and include 

human uses, such as fisheries, that can affect the population 

dynamics, community dynamics, and species presence in an 

area. In addition to providing information about environmental 

constraints (limiting factors) to marine fish populations, the 

BOEM requested more generalized information about the 

region’s physical setting, adaptation and acclimation, life 

history strategies, marine fish assemblages, foraging and 

feeding behaviors, seasonality, growth and reproduction, 

predation and disease, population abundance and production, 

and human uses. We were asked to consider potential effects 

of climate change and, in light of all of the above, provide 

our impressions of information needs and research directions. 

Our opinions on potential effects of climate change should be 

viewed as hypotheses and, our thoughts on information needs 

are directed at the Arctic scientific community, in general, and 

no single organization, in particular. In many instances, these 

opinions were directed at deficiencies in understanding of 

species biology and ecology and basic information needs for 

any assessment or management action.

Biodiversity Assessment

The assessment of regional biodiversity included: 

(1) developing and updating an Arctic marine fish checklist; 

(2) reviewing recent (since Fishes of Alaska was published) 

ichthyologic sampling and management documents; 

(3) searching online databases; (4) systematic investigations 

that include documenting current taxonomic understanding 

with genetic information, historical museum collections, and 

voucher specimens; (5) acquiring digital images (photographic 

and line drawings); (6) creating visualization products that 

include distribution maps and vertical species profiles; and 

(7) conducting peer reviews. 

Arctic Marine Fish Checklist

A provisional working list of marine fishes inhabiting 

the Chukchi and Beaufort Seas was initially prepared. 

This important first list incorporated new discoveries and 

taxonomic resolutions from ichthyologic sampling, genetic 

analysis, and museum studies conducted since the regional 

monograph Fishes of Alaska (Mecklenburg and others, 

2002) and checklist of fishes (Love and others, 2005) 

were published. The list included only marine fish species 

confirmed from the region and greatly benefited from recently 

completed and ongoing field, laboratory, and museum research 

establishing new baselines for marine fish in the Pacific Arctic 

Region (Mecklenburg and Steinke, 2015). It was evident to 

BOEM and the U.S. Geological Survey (USGS) that much 
18 Alaska Arctic Marine Fish Ecology Catalog

new information on species distribution and occurrence had 

been collected in the decade following publication of the 

Fishes of Alaska and, as noted by Mecklenburg and Steinke 

(2015), this reference is no longer complete for the region. 

An important objective of this study was to illustrate the 

relationship of the final checklist to Mecklenburg and others 

(2002), which remains an important companion document and 

primary source for taxonomic information and identification 

guides for Alaskan fishes. 

In addition to Mecklenburg and others (2002), other key 

references included reviews and syntheses on the taxonomy 

and zoogeography of Arctic marine fishes by Mecklenburg and 

others (2007), Mecklenburg and others (2011, 2013, 2014), 

Mecklenburg and Steinke (2015), and Maslenikov and others 

(2013). A new atlas and identification guide for the marine 

fishes of the Pacific Arctic Region is an important source of 

information about the diversity, taxonomy, and geographic 

distribution of marine fishes in the Chukchi and Beaufort Seas 

(Mecklenburg and others, 2016). 

Scientific Reviews

Recent publications, technical reports, and unpublished 

data were reviewed for new species occurrences and 

expansions in range. Survey results from fish sampling in 

the Chukchi Sea (Mecklenburg and others, 2007, 2011; 

Norcross and others, 2009, 2013a) and Chukchi Borderlands 

(for example, Mecklenburg and others, 2013; Longshan and 

others, 2014) and industry research (Gallaway and Norcross, 

2011; Norcross and others, 2013b) were reviewed. Surveys 

from the southwestern Beaufort Sea (Johnson and others, 

2010; Parker‐Stetter and others, 2011; Rand and Logerwell, 

2011; and Johnson and others, 2012) were reviewed for new 

information from nearshore and shelf waters. Additionally, 

National Oceanic and Atmospheric Administration (NOAA), 

University of Alaska-Fairbanks (UAF), and RUSALCA 

scientists are involved in ongoing fishery investigations for 

BOEM’s Alaska OCS Region in the Chukchi and Beaufort 

Seas. Taxonomic support provided to these studies and 

other research (for example, Mecklenburg and Steinke, 

2015) and analysis of previously unprocessed samples from 

WEBSEC-71 (1971) (Hufford, 1974) was a source of new 

records from the Beaufort Sea. 

Relevant environmental analyses and Arctic resource 

management plans done by the BOEM (Minerals 

Management Service [MMS], 2008) and North Pacific 

Fishery Management Council (NPFMC, 2009), respectively, 

were reviewed in this assessment of biodiversity, human 

uses, and environmental relationships. The DOI synthesis of 

priority science needs for continued oil and gas development 

in the Arctic was consulted for information about the 

region’s environmental setting, living resource information, 

and USGS-identified research needs (Holland-Bartels and 

Pierce, 2011). A supplemental draft EIS addressing possible 

offshore oil and gas development in the U.S. Chukchi and 

Beaufort Seas was reviewed for species and community 

assemblage information (National Oceanic and Atmospheric 

Administration, 2013). A significant study of subsistence 

use and traditional ecological knowledge of whitefish in 

the southeastern Chukchi Sea (Georgette and Shiedt, 2005) 

provided an important source of information about Iñupiat 

taxonomy and seasonal use of whitefishes in the Noatak, 

Kobuk, and Selawik Rivers of Kotzebue Sound (fig. 1.7).

Searchable Databases

The exploration of the Arctic is at the forefront of 

ecological research to document and understand marine 

ecosystems and population ecologies with respect to climate 

change, possible commercial fisheries, and other stressor 

effects. In recent years, active research in conjunction with 

RUSALCA, Census of Marine Life (CoML) programs [that 

is, Pan-Arctic Inventory/Arctic Ocean Diversity (ArcOD) and 

Marine Barcode of Life] has emphasized fishery objectives for 

resource inventory. In this capacity, data for this report were 

from the Global Marine Life Database, Census of Marine Life 

Community Database, and CoML bibliographic database (for 

example, Mecklenburg and others, 2007 and Mecklenburg 

and Mecklenburg, 2009). Other searchable Web-based 

databases used included: FishBase; BOEM Arctic Fisheries 

Database; United Nations Ocean Biogeographic Information 

System (OBIS Database); Encyclopedia of Life; World 

Register of Marine Species; Catalogue of Life Species; and 

Integrated Taxonomic Information System (ITIS). NOAA’s 

on-line database services provide access to life history and 

distributional information for nearshore atlases; offshore 

fishery resources were accessed at the National Oceanic and 

Atmospheric Administration Web site (http://alaskafisheries.

noaa.gov). The automated database at the University of 

Washington School of Fisheries and Ocean Sciences provided 

key access to type specimens and mapping tools.

The BOEM database, “Alaska Scientific and Technical 

Publications” (http://www.boem.gov/Alaska-ScientificPublications/), was a significant resource for marine fishery, 

subsistence, and socioeconomic data. A summary of ongoing 

and recently completed BOEM fishery studies is provided in 

appendix A. A major synthesis by Braund and Kruse (2009) 

provided an authoritative review of subsistence harvest 

patterns for fish and wildlife patterns over the past 30 years 

(from the mid-1980s). During this period, the OCS subsistence 

studies and this synthesis focused on coastal harvest patterns 

in the Alaska North Slope villages of Wainwright, Barrow, 

Nuiqsut, and Kaktovik. An early study by Craig (1987) 

provided important baseline documentation of subsistence 

patterns and Iñupiat taxonomy from the Chukchi and 

Beaufort Seas. 
Chapter 1 19

Systematic Investigations

Recent information about the taxonomy and 

biogeography of Alaskan Arctic fishes has come from 

numerous analyses using field and museum studies, DNA 

sequencing, and exhaustive literature review, as well as efforts 

to synthesize the accumulated results for the Pacific Arctic 

region; particularly the works of Mecklenburg and others 

(2006, 2007, 2011, 2013, 2014, 2016), Mecklenburg and 

Mecklenburg (2009), and Mecklenburg and Steinke (2015). 

Voucher specimens provide a crucial link to historical data 

sets, allowing determination of existing baselines and accurate 

identification of new field collections. Museum collections 

housing the specimens are globally located and managed 

by many institutions. The following collections are some of 

those studied for the works just cited: California Academy 

of Sciences, San Francisco; Canadian Museum of Nature, 

Gatineau, Quebec; Hokkaido University Museum of Zoology, 

Hakodate, Japan; U.S. National Museum of Natural History, 

Washington, D.C.; NOAA, Alaska Fisheries Science Center, 

Auke Bay Laboratory, Juneau, Alaska; University of Alaska 

Museum, Fairbanks, Alaska; University of British Columbia, 

Vancouver, Canada; University of Washington, Seattle; and 

Zoological Institute, St. Petersburg, Russia. Detailed data on 

voucher specimens examined, including the authors’ verified 

or redetermined identifications, have been published (for 

example, Mecklenburg and others, 2006; Mecklenburg and 

Mecklenburg, 2009; and Mecklenburg and others, 2011, 

Online Resources 1 and 2) and the most recent results are 

available from the museums’ online search databases. In 

many instances, the museum studies have revealed previously 

misidentified or unreported specimens and have led to 

extensions or contractions in known patterns of distribution; 

these results are reported here in abbreviated form from the 

authors’ more detailed works. The origins, classification, 

diversity, human uses, and vulnerabilities of circumpolar 

freshwater and marine Arctic fishes were reviewed as part of 

the most recent Arctic Biodiversity Assessment (Christiansen 

and others, 2013).

Genetic Analyses

Mecklenburg and others (2011, 2014, 2016) and 

Mecklenburg and Steinke (2015) described the DNA analyses 

(barcoding) performed on Arctic fish specimens as part of 

the CoML, RUSALCA, and other fisheries investigations. 

This analytical tool has provided critical information to 

ichthyologists to help resolve taxonomic problems, assist 

difficult identifications, and, in many instances, generate 

hypotheses about potential population structuring. Many of 

the Arctic fishes that have been barcoded were collected in 

the Chukchi and Beaufort Seas and in other parts of Alaska. 

Online Resources 3 and 4 of Mecklenburg and others (2011) 

provide museum catalog numbers and collection data for 

barcoded specimens. That collection has been increased, and 

data on additional specimens may be found on the Barcode 

of Life Data System (BOLD) Web site from links supplied by 

Mecklenburg and Steinke (2015). 

Other sources of genetic identification information 

were found in the literature review. Research on Pacific 

salmon has been extensive in Alaska including north through 

Kotzebue Sound. BOEM and other organizations including 

the North Pacific Research Board (NPRB) are funding new 

research on the genetic diversity and population structure of 

Arctic Gadidae (for example, Arctic, Saffron, and Ice cod) 

and the new results and ongoing studies are, or will, assist in 

addressing existing taxonomic complexities (for example, see 

appendix A and Nelson and Bouchard, 2013).

Digital Images

Many recent high-quality images of marine fishes were 

obtained from active Arctic researchers who photographed 

freshly sampled specimens in the field. For example, most 

of the images in this report are products of research of 

Mecklenburg and others (2016). Each photograph is properly 

attributed with names of providers and date and location of 

sample collection.

Distribution Maps and Vertical Species Profiles 

ArcGIS geographic range maps were developed for 

each confirmed species from the Chukchi and Beaufort 

Seas. Vertical distribution profiles were created in Adobe 

software to portray specific information in a generalized life 

cycle schemata depicting water column distributions of life 

stages over continental shelf and slope habitats. The maps 

and profiles depicting geographic distributions and vertical 

distribution information are composite products from multiple 

sources of data collection and observation. A BOEM study 

with objectives of compiling existing fishery data sets for 

the Chukchi and Beaufort Seas into an electronic format 

was not completed in time for use herein. Although the data 

were acquired through traditional sources, easier access to, 

and manipulation of these data, in concert with the results of 

ongoing fishery research, would improve the efficiency and 

number of Alaskan records in future updates.

Peer Review

USGS standards for science quality (USGS Fundamental 

Science Practices) were followed (http://www.usgs.gov/

fsp/) for this study and report. The USGS process includes 

technical and policy reviews for technical reports and journal 

publications. For this report, we relied on external subject 

matter experts due to the diversity of species and ecological 

topics addressed. The USGS reviews were supported by 

BOEM technical reviews that included additional internal and 

external scientists having appropriate ichthyologic expertise. 
20 Alaska Arctic Marine Fish Ecology Catalog

Operational Definitions

In some cases, especially in the species accounts and 

synthesis discussions, highly technical information relative 

to various aspects of marine biogeography is presented. 

In several disciplinary areas, especially those dealing with 

quantitative population ecology, special attention to defining 

the scientific terminology used was needed. To facilitate 

consistency and understanding, a glossary of key terms was 

developed as an aide for users. The glossary (chapter 7) 

defines many of the more complex terms and quantitative 

relationships used in the species accounts. Each definition 

is appropriately cited so that, if they are not clear, additional 

sources can be quickly accessed and referenced. Other 

terms commonly used to describe spatial relationships in 

biogeography, involve ecological scaling considerations of 

importance to conservation and are described in the section, 

“Geospatial Terminology.” The terms described in this chapter 

are used widely throughout the report and, for that reason, are 

introduced here. Abundance terminology represented a special 

case where the derivation of a unique classification scheme for 

the Arctic region was necessary and developed in chapter 2

with application to other chapters as well. 

Geospatial Terminology

This study focuses on the biological geography of marine 

fishes in Alaska’s Arctic Ocean and describes their distribution 

and abundance using various spatial terms. Some have 

zoogeographic meaning (for example, realm and province) and 

others refer to common descriptions of a species occurrence 

by scale (for example, region, area, and local). The mapping 

classifications for biogeographic realms and faunal provinces 

used herein are as described by Lourie and Vincent (2004) and 

Spalding and others (2007), respectively. The Arctic region, as 

described by Briggs and Bowan (2012), adds additional clarity 

to the boundaries of marine provinces by considering the 

zoogeography of fishes of fishes. For Alaska, Sigler and others 

(2011) described three provinces that relate to conventional 

management areas in the United States segments of the 

northern Bering Sea and the Chukchi and Beaufort Seas. 

In each case, the biogeographic concepts relate to 

ecological patterns and processes, and thus to conservation. 

Lourie and Vincent (2004, p. 1,005) noted that, “...biodiversity 

exists on multiple scales biologically, spatially and temporally, 

and these scales are, to some extent, independent of one 

another.” They related spatial scales of conservation planning 

to physical, temporal, and biological patterns and processes, 

to mapping scale as follows (Lourie and Vincent (2004, p. 

1,007): 

(1) global approximate map scale 1:100 million 

– ocean basin divisions, major currents, global 

climate, historical biogeography, highly migratory 

species, large marine ecosystems (LMEs); 

(2) regional, 1:10 million – regional currents, 

historical biogeography, genetic connectivity, 

widespread species, individual LMEs or ecoregion; 

(3) provincial, 1.1 million – small scale currents, 

upwelling, genetic connectivity, major habitats 

with LME or ecoregion restricted-range species, 

bioregion; (4) local, 1:500,000 – local gyres and 

eddies, watershed runoff, coastal geomorphology, 

ecological connectivity, planning unit; (5) site, 

1:10,000 – tides, watershed runoff, ecological 

connections, habitat specialists, habitats within 

planning unit, zoning for marine protected areas. 

The scales are germane because they give additional texture 

to how (unless otherwise specified) the concepts of realm and 

province should be understood and, generally, the geographic 

distinctions between “region” and “area” as used in this report. 

Typically, “region” is used to refer to large-scale patterns and 

process (for example, scale 3 in Lourie and Vincent [2004]) 

and “area” refers to local or site specific ecological conditions 

(for example, scales 4 and 5).

The use of other geographical or habitat terms, including 

inshore, coastal, nearshore, and offshore in this report are 

somewhat more ambiguous with respect to meaning or area 

involved. For example, in some instances “coastal” may have 

a legal meaning with respect to a state’s territorial waters, or 

meteorological and oceanographic meaning with respect to 

wave influence. Similarly, “nearshore” also may have multiple 

oceanographic meanings including reference to littoral zone 

components, photic zones, or waters extending to the shelf 

break. There can be considerable overlap in area with respect 

to how these terms are applied with respect to the notion of 

“close to shore.” In an attempt to reduce such ambiguity, 

some practical distinctions are made for these terms based 

on scientific knowledge of Arctic fish habitat characteristics 

(depth, temperature and salinity influences, and distance from 

shore) and realities of their sampling by proven fisheries 

techniques:

• Inshore reflects the area from shoreline to depths of 

2 m. This reflects the shallow water areas where much 

of the historical fish baseline data were collected in the 

Chukchi and Beaufort Seas surveys with fyke net and 

gill net sampling techniques. Freshwater influences 

typically are important seasonally, especially in the 

Beaufort Sea.

• Coastal habitats can include lower reaches of rivers 

and streams, deltas, inshore, and deep waters adjacent 

to the coast. Lagoons and shallow waters adjacent 

to outer coasts of barrier islands are included in 

the category. Freshwater influences are seasonally 

important in coastal habitats.

• Nearshore, or nearshore marine, as used herein, 

generally refers to those waters located inside the 

10-m isobath. This depth is often where large (class 1) 

oceanographic vessels cease to operate for safety 

reasons. Additionally, the presence of brackish water 
Chapter 1 21

can be a factor in describing the distance that nearshore 

waters occur from the coast. For example, during 

summer, brackish waters can extend more than 10 km 

offshore and often overlay cold more saline water 

masses deep in the water column. The nearshore 

includes inshore and coastal habitats, but tend to 

extend farther offshore.

• Offshore as used here describes shelf, slope, and 

deep‐water habitats located beyond the nearshore. 

• Nearshore-Marine refers to a descriptive adaptive 

model in this report as evidenced by those species 

having distribution patterns that are widespread over 

nearshore, shelf, and slope environments. An example 

would be the distribution of Arctic Cod in the United 

States Chukchi and Beaufort Seas. 

A fish species can be distributed in one or more habitats 

in time and space. This report provides a comprehensive 

examination of how fish in Arctic environments have adapted, 

which depending on life strategy adaptations, represents a 

continuum of habitats on a gradient from freshwater to marine. 

For each species, patterns of habitat use are multi-dimensional 

with respect to areas and volumes occupied, space and time, 

and life cycle requirements for reproduction, growth and 

survival, dispersal, and other life functions. The distribution 

and abundance of life stages for each species throughout 

the water column are presented in chapter 3 to capture these 

relationships in the context of life zones (Allen and Smith, 

1988) that portray key aspects of reproductive ecology by 

depth and location. Depictions in this report are unique for this 

region because the shelf break is depicted at 75 m on the basis 

of the bathymetry of the Chukchi and western Beaufort Seas.

Organization of the Catalog

The BOEM requested that the USGS conduct a synthesis 

of available environmental information addressing Arctic 

marine fish biology and ecology in the following areas: 

environmental and biological constraints; oceanographic 

overview; adaptation and acclimation; life history 

strategies; fish assemblages; foraging and feeding behavior; 

bioenergetics; use of time and space; growth and reproduction; 

migration; predation, parasitism, competition, and mutualism; 

dynamics of population abundance and production; 

conservation; subsistence; climate change; and information 

needs. The resultant Arctic Marine Fish Ecology Catalog is 

organized in eight chapters, a glossary, and three appendixes. 

Chapters 1–3 provide detailed characterizations of the 

United States Arctic region’s environmental setting and 

marine fish biodiversity and establish much of the physical 

and biological background information supporting ecological 

synthesis and management objectives of this project. Chapter 1

provides a broad overview of the Arctic marine environment 

and scientific goals and methods of this synthesis as it pertains 

to BOEM information needs as well as other users. Additional 

purposes are to describe the meaning of commonly used 

oceanographic and zoogeographic terminologies to avoid 

possible ambiguity associated with their use herein. Chapter 2

presents an inventory of the marine fish known from the 

United States Chukchi and Beaufort Seas including an analysis 

of changes resulting from new data and information acquired 

from the region since the early 2000s. Chapter 3 includes more 

than 100 species accounts and describes current knowledge 

about the biology and ecology for each marine fish known 

from the United States Arctic region using information from 

studies in Alaska and elsewhere within the species ranges. 

The species accounts are quick references to some topics (for 

example, climate change) and are described in more detail in 

chapters 5–6.

Chapter 4 provides a synthesis of information that, in 

its entirety, addresses the ecological objectives requested by 

BOEM. The USGS approach to this synthesis was to evaluate 

data and information about the descriptive and interpretive 

components of biogeography as described by Nelson (2006). 

By necessity, much information is incorporated by reference 

in this description of historical and environmental influences 

on marine fish presence and patterns of distribution and 

abundance in the United States Arctic marine environment.

Chapter 5 focuses on the modeling of climate change 

effects on the distribution of Arctic and Saffron cod in the 

Bering Sea. The Bering Sea was selected because of the 

extensive fishery data that are available from this region. 

Arctic and Saffron cod were selected because they are notably 

important in Arctic food webs and subsistence fisheries north 

of the Bering Strait. Our emphasis on modeling approach 

and distributional effects represent an important potential 

application to the Chukchi and Beaufort Seas as Arctic fishery 

data become more widely accessible in electronic formats. The 

results of modeling in the Bering Sea are considered in light of 

potential warming effects in the Chukchi and Beaufort Seas.

Chapter 6 focuses on how scientific information about 

the Arctic marine fish can inform policy and the conservation 

and management of Arctic species. Information needs relate to 

environmental assessment for OCS oil and gas development, 

fisheries management, and understanding potential effects of 

climate change. A traits-based approach can be used to assist 

resource management until population-level information and 

dynamics are better known. The information needs for highpriority species identified in chapter 3 also are evaluated with 

respect to future research and ecosystem-based management.

This report includes many descriptive and quantitative 

terms and complex concepts common in scientific literature. 

A “Glossary of Ecological Terms” is provided in chapter 7 to 

reduce confusion about these terms and concepts as used in 

this report. Chapter 8 provides a list of scientific references 

cited in narrative sections of the report. Each species account 

presented in chapter 3 includes a listing of the scientific 

reports and articles used. 
22 Alaska Arctic Marine Fish Ecology Catalog

Appendix A describes age, weight, and size relationships 

for marine fish species where data are available. The 

information supports life history and population dynamic 

sections of the report in chapters 4 and 6. Appendix B provides 

additional information about the availability of recent fishery 

investigations supported by BOEM as well ongoing studies, 

and topics where new information is anticipated. Appendix C

provides summaries of the models developed to examine 

temperature effects on Arctic and Saffron cods in chapter 5.

Acknowledgments

The authors wish to thank the U.S. Department of the 

Interior, Bureau of Ocean Energy Management (BOEM), 

Environmental Studies Program for their support of this 

study. The support of the U.S. Geological Survey (USGS) 

Outer Continental Shelf (OCS) Studies Program, the USGS 

Western Fisheries Research Center, and the USGS Alaska 

Region also is appreciated. This report has been technically 

reviewed by expert specialists in BOEM and by the USGS 

managed peer review processes. The authors wish to 

acknowledge all the fishery scientists and oceanographers who 

have contributed to the regional knowledge of Arctic marine 

fishes and marine ecosystem processes. We have relied on a 

vast amount of research by others in preparing this catalog. 

More specifically, we are especially grateful for the insights, 

support, and reviews our team received from Katherine 

Wedemeyer from BOEM’s Alaska OCS Region. Kate’s 

original vision, guidance, and coordination with other BOEM 

researchers greatly enhanced the scientific quality and form of 

information delivery herein. Similarly, we are grateful for the 

administrative support of Colleen Charles, USGS Ecosystem 

Mission Area, and members of the USGS Western Fisheries 

Research Center who contributed to this study. In particular, 

we thank Debra Becker (USGS) for her assistance in all 

production phases of the final report. Finally, special thanks 

to Carl J. Markon and Leslie Holland-Bartels from the USGS 

Alaska Science Center for allowing me to continue in this 

study following my return to Alaska in 2011. Carl Markon’s 

assistance with final publication will be forever appreciated. 

Tony DeGange’s (formerly USGS Alaska Science Center) 

literature review in support of USGS Circular 1370 provided 

important background on the regional setting of the ChukchiBeaufort Sea study area. The authors are especially grateful 

for the efforts of the USGS Science Publishing Network, 

Tacoma Publishing Service Center for producing this report 

and specifically to acknowledge Debra Grillo for editorial 

assistance and William Gibbs for the cover design.

Summary

The need for reliable fishery information for National 

Environmental Policy Act analysis related to offshore oil 

and gas development in Chukchi and Beaufort Sea Planning 

Areas is described. The conduct of systematic surveys in these 

areas has been hampered by environmental and budgetary 

constraints and thus, in many respects, the existing data remain 

scattered and fragmented and are difficult to access. Lack 

of access to quantitative catch and observational data from 

ongoing research precluded our full use of modern geospatial 

technologies. Despite, this limitation, new information and 

visualization products were developed for each species of 

confirmed presence in the study area. The biological and 

ecological information presented in this report is regional in 

scale and, unless otherwise noted, local variations in seasonal 

fish habitat and abundance can be found in the literature cited. 

Given the spatial resolution of existing data, background 

descriptions of the physical and biological environments of 

the Chukchi and Beaufort Seas are presented at the landscape 

scale. All methods, terminologies, and operational definitions 

for this study are described. The organization of the report and 

linkages between chapters are described as they relate to the 

Bureau of Ocean Energy Management’s information need for 

this synthesis.
Abstract

Several other marine fishery investigations, including 

efforts for Arctic data recovery and regional analyses of range 

extensions, were ongoing concurrent to this study. These 

included Bureau of Ocean Energy Management-sponsored 

research in the Chukchi Sea and nearshore and transboundary 

surveys in the Beaufort Sea. New collections in the Chukchi 

Sea and Chukchi Borderland also were being obtained 

through National Oceanic and Atmospheric Administration’s 

multidisciplinary Russian-American Long-term Census of the 

Arctic (RUSALCA) program. These were major exploratory 

efforts and represented potential sources of new marine 

species information. Building on the “Fishes of Alaska,” an 

updated checklist was constructed of known and probable 

marine fishes with catch data obtained in these studies. There 

are 109 known species from 24 families of marine fish from 

the United States High Arctic, with 97 species reported 

from the Chukchi Sea and 83 from the Beaufort Sea. As this 

synthesis was being done, changes in the confirmation process, 

based on morphological and genetic analysis, were captured 

as information became available in scientific publications. 

The known species are reviewed with respect to taxonomic, 

geographic, and large-scale abundance considerations. 

Each species’ zoogeographic pattern with respect to our 

understanding of broadly defined evolutionary origins is noted. 

Changes in species nomenclature are described as they relate 

to newly described standards. 

Introduction

A large number of Arctic fisheries studies were 

started following the publication of the Fishes of Alaska

(Mecklenburg and others, 2002). Although the results of 

many of these efforts are not yet available, those involving 

field sampling are important new sources of biodiversity 

information including voucher specimens and genetic 

confirmation of a species existence and phylogenetic 

relationships. The addition of new morphological data and 

genetic analyses is important because despite many efforts, 

systematic sampling in the Chukchi and Beaufort Seas has 

been logistically difficult due to ice, storms, and platform 

limitations. Further variations in research missions and 

sampling methods have hampered regional assessments of 

diversity, relative abundance, and habitat importance because 

the resulting database is sparse and fragmented in time and 

space. Quantitative evaluations of abundance based on catch 

data are difficult, although local variability and persistent hot 

spots are known to exist. This chapter builds on the species 

inventory established by the Fishes of Alaska by incorporating 

new species information obtained primarily in Bureau of 

Ocean Energy Management (BOEM)-sponsored (appendix A) 

and National Oceanic and Atmospheric Administration 

(NOAA) Russian-American Long-term Census of the Arctic 

(RUSALCA) surveys (Mecklenburg and Steinke, 2015). 

This information is complemented with a uniformly applied 

assessment of relative abundance and qualitative analysis 

of population viability based on present day understanding 

of endemism. 

Chapter 2. Alaska Arctic Marine Fish Inventory

By Lyman K. Thorsteinson1

1

U.S. Geological Survey.
24 Alaska Arctic Marine Fish Ecology Catalog

Checklist of Marine Fish

The updated checklist of confirmed occurrences is 

significant because it incorporates the new knowledge gained 

since the publication of the Fishes of Alaska (Mecklenburg and 

others, 2002). It includes the most recent information from the 

Pacific Arctic Region including the Chukchi and Beaufort Seas 

from Mecklenburg and Steinke (2015). Unlike Mecklenburg 

and Steinke (2015), this checklist includes diadromous species 

from the family Salmonidae as these species spend significant 

portions of their lives in the marine environment and have 

great importance in subsistence lifestyles. Even though this list 

represents a significant advance, many uncertainties remain 

about the true composition of the marine fish assemblage. 

For this reason, information about probable occurrences of 

marine fishes is included in this chapter. The uncertainties 

result in part because the taxonomy of Arctic fishes was 

still in its infancy at the turn of the 21st century, with many 

species misidentified in historical literature and others not 

recognized from the region due to sampling inadequacies 

(Mecklenburg and others, 2011). The scientific improvements 

in knowledge about the Arctic region’s biodiversity reflect 

new data collection and, importantly, advances in taxonomic 

and zoogeographic understanding that have resulted from 

genetic resolution of issues that morphological studies failed to 

notice or resolve (for example, Mecklenburg and others, 2011; 

Lynghammar and others, 2012; Mecklenburg and Steinke, 

2015). The Arctic marine fish inventory, presented in table 2.1, 

is a major biodiversity achievement, which documents 

known (verified) marine fish species from the Chukchi and 

Beaufort Seas. 

Although advances have been significant in recent years, 

the taxonomy, including naming of Arctic marine fishes 

remains a source of inconsistency and confusion between 

leading authorities such as the American Fisheries Society 

(AFS), California Academy of Sciences Catalog of Fishes, 

Integrated Taxonomic Information Systems, and World 

Register of Marine Fishes, for members of several families. 

This is especially true for common names associated with 

different languages (Mecklenburg and others, 2013). Changes 

in nomenclature are not uncommon in the classification 

process, especially for lesser-studied species, and can 

affect the regional use of scientific and common names in 

literature (Eschmeyer and others, 2010). Thus, in creating 

checklists, it is important to cite the authority used and 

rationale for inclusion or naming of a particular species where 

disagreement occurs. In this report, the marine fish inventory 

presented in table 2.1 reflects the editors’ understanding of 

the classification and nomenclature of marine Arctic fishes. 

In most instances, the scientific and common names are those 

listed by Page and others (2013) and Eschmeyer and others 

(2015). However, in several instances where differences exist, 

we relied on the work of contemporary Arctic specialists and 

note areas of transitional change with respect to common 

names for species.

The nomenclature of a small number of fish species from 

four families (Squalidae, Gadidae, Liparidae, and Zoarcidae) 

are in transition and reflect areas where we disagree with the 

AFS nomenclature (Page and others, 2013). For purposes of 

this report, we decided to agree with the AFS nomenclature 

for Boreogadus saida due to the species’ significance to the 

region’s marine ecology. Although, we acknowledge that 

Polar Cod is emerging as the widely accepted common name 

for B. saida, we decided to use Arctic Cod (as recommended 

by Page and others [2013]) to avoid any confusion with 

new and historical Alaska literature. In contrast, we decided 

to use Ice Cod, not Polar Cod, as the common name for 

Arctogadus glacialis. This decision was made to avoid any 

potential ambiguity and confusion with B. saida. The use of 

Ice Cod is emerging nomenclature in scientific literature (for 

example, Arctic Council’s Conservation of Arctic Flora and 

Fauna) and clarification of the vernacular name is supported 

by genetic analysis (Nelson and Bouchard, 2013). Regarding 

other families and species, we recognize (1) Spotted Spiny 

Dogfish instead of Pacific Spiny Dogfish as the common 

name for Squalus suckleyi (Squalidae, see Lynghammar and 

others [2012]); (2) Estuarine Eelpout, not Polar Eelpout as 

the common name for L. turneri (Mecklenburg and others, 

2002); and (3) Arctic Sand Lance for Ammodytes hexapterus

following Orr and others (2015) and Mecklenburg and others 

(2016). Other more recent changes (for example, Mecklenburg 

and others, 2016) include changes in the scientific names for 

Capelin (now Mallotus catervarius) and Arctic Flounder (now 

Liopsetta glacialis). In every case, the scientific name of a fish 

is conclusive for the species being described. 

One hundred nine (109) species from 24 families are 

confirmed with 97 verified occurrences from the Chukchi 

Sea and 83 from the Beaufort Sea (table 2.1). Mecklenburg 

and others (2002) suggested other species (<20) also might 

be present and it remains likely that the total number will 

change with additional field sampling, examination of existing 

voucher specimens, and with shifts in range due to climate 

warming. Mecklenburg and others (2007, 2011, 2014, 2016) 
Chapter 2 25

and Mecklenburg and Steinke (2015) have recently confirmed 

the presence of at least 18 species in the Alaskan Chukchi and 

Beaufort Seas that were not previously verified to be present: 

Somniosus pacificus, Amblyraja hyperborea, Bathyraja 

parmifera, Myoxocephalus polyacanthocephalus, Triglops 

nybelini, Hypsagonus quadricornis, Careproctus reinhardti, 

Liparis bathyarcticus, Paraliparis bathybius, Lycenchelys 

kolthoffi, Lycodes adolfi, L. frigidus, L. jugoricus, L. marisalbi, 

L. pallidus, L. reticulatus, Anarhichas denticulatus, and 

Zaprora silenus. The status of three of these species was 

previously described as being of “probable” occurrence in the 

provisional checklist developed for this study:

• Pacific Sleeper Shark (Somniosus pacificus): Benz 

and others (2004) reported a new occurrence of 

the species from an animal found washed onshore 

at Point Hope, Alaska. The animal remains were 

highly decomposed; this occurrence was reviewed 

by Lynghammar and others (2012), who determined 

that the physical condition of the shark suggested 

its death from outside the Chukchi Sea. In 2014, 

a Pacific Sleeper Shark was taken alive by a seal 

hunter in a southeastern Chukchi Sea lagoon. Parts 

and photographs were archived in the University of 

Alaska Museum. This is a mesobenthopelagic shark 

species recorded as deep as 2,205 m. It is abundant in 

the southern Bering Sea and the Gulf of Alaska, and 

reaches 4.4 m TL or more (Mecklenburg and others 

(2016).

• Alaska Skate (Bathyraja parmifera): Mecklenburg 

and others (2011 Online Resource 1) reported that two 

specimens were found washed ashore in the Chukchi 

Sea. Examination of one of the specimens indicated 

advanced deterioration and possible death elsewhere 

in the northern Bering Sea. Later, Mecklenburg 

and others (2016) reported that in 2012 an adult 

specimen was taken alive in the southern Chukchi 

Sea. Mecklenburg and others (2016) also reported the 

collection locations of other beachcast specimens and 

spent egg cases from the Chukchi and Beaufort Sea. 

The authors noted the species can reach a size of 135 

cm TL. They also reported that the occurrences of 

the Alaska Skate in the Chukchi Sea could indicate a 

northerly shift in its distribution.

• Pale Eelpout (Lycodes pallidus): Mecklenburg and 

others (2016) identified this species in a collection 

made in 1971 in the Beaufort Sea and confirmed 

presence in the Beaufort Sea again in voucher 

collections from 2012 and 2013. Common at least 

in the Greenland, Barents, and Kara Seas, otherwise 

found nearly circumpolar. It is epi-meso-bathybenthic 

and lives on muddy substrate at depths of 11–1,750 m 

and temperatures of -1.8–3.7 °C. Maximum size is 

28.9 cm SL or more (Mecklenburg and others, 2016).

The updated checklist was examined with respect to 

new species occurrences and differences in the geographic 

distributions reported in the Fishes of Alaska (Mecklenburg 

and others, 2002) for the Chukchi and Beaufort Seas 

(table 2.2). Our decision to include two species, Artediellus 

ochotensis and Lumpenus sagitta, on the checklist was made 

recognizing that uncertainty exists about their occurrence 

in the Chukchi Sea (Mecklenburg and others, 2016). As 

an example, Mecklenburg and others (2016) state “Many 

of the specimens that had been previously identified as 

“A. ochotensis” on further investigation have been found 

to be A. scaber.” However, not all of the identifications 

were refuted. In the case of L. sagitta, Mecklenburg and 

others (2011) described a range extension for this species 

into the Chukchi Sea based on a single specimen. Presently, 

Mecklenburg and others (2016) raise doubts “The provenance 

of a specimen reported to be a possible record of L. sagitta

from the Chukchi Sea...is uncertain.” The uncertainty stems 

from conflicting issues in the 1960s field collection data and 

location of sampling (possibly not Chukchi Sea), and not 

with species identification. Until these issues can be resolved 

with species confirmations with new specimens, we chose 

to retain them on the checklist. As a final note, we recognize 

Icelus spiniger as a valid species from the Chukchi Sea noting 

that genetic similarities with I. spatula specimens from the 

region raise questions about relationships. Other technical 

and unresolved issues for the marine fishes in the Chukchi 

and Beaufort Seas are described in Mecklenburg and Steinke 

(2015) and Mecklenburg and others (2016).
26 Alaska Arctic Marine Fish Ecology Catalog

Table 2.1. Inventory of confirmed species of Arctic marine fish in the United States Chukchi and Beaufort Seas, Alaska.

[Blank cells indicate that the species occurrence has not been confirmed in that sea. Marine waters out to the U.S. Exclusive Economic Zone (200-mile limit) are 

included]

Family Scientific name Common name

Confirmed occurrence

Chukchi Sea Beaufort Sea

Petromyzontidae Entosphenus tridentatus Pacific Lamprey X

Petromyzontidae Lethenteron camtschaticum Arctic Lamprey X X

Squalidae Squalus suckleyi Spotted Spiny Dogfish X

Somniosidae Somniosus pacificus Pacific Sleeper Shark X

Rajidae Amblyraja hyperborea Arctic Skate X

Rajidae Bathyraja parmifera Alaska Skate X

Clupeidae Clupea pallasii Pacific Herring X X

Osmeridae Hypomesus olidus Pond Smelt X

Osmeridae Mallotus catervarius Pacific Capelin X X

Osmeridae Osmerus dentex Arctic Smelt X X

Salmonidae Coregonus autumnalis Arctic Cisco X X

Salmonidae Coregonus laurettae Bering Cisco X X

Salmonidae Coregonus nasus Broad Whitefish X X

Salmonidae Coregonus pidschian Humpback Whitefish X X

Salmonidae Coregonus sardinella Least Cisco X X

Salmonidae Oncorhynchus gorbuscha Pink Salmon X X

Salmonidae Oncorhynchus keta Chum Salmon X X

Salmonidae Oncorhynchus kisutch Coho Salmon X X

Salmonidae Oncorhynchus nerka Sockeye Salmon X X

Salmonidae Oncorhynchus tshawytscha Chinook Salmon X X

Salmonidae Salvelinus malma Dolly Varden X X

Salmonidae Stenodus leucichthys Inconnu X

Myctophidae Benthosema glaciale Glacier Lanternfish X

Gadidae Arctogadus glacialis Ice Cod X X

Gadidae Boreogadus saida Arctic Cod X X

Gadidae Eleginus gracilis Saffron Cod X X

Gadidae Gadus chalcogrammus Walleye Pollock X X

Gadidae Gadus macrocephalus Pacific Cod X X

Gasterosteidae Gasterosteus aculeatus Threespine Stickleback X X

Gasterosteidae Pungitius pungitius Ninespine Stickleback X X

Hexagrammidae Hexagrammos stelleri Whitespotted Greenling X X

Cottidae Artediellus ochotensis Okhotsk Hookear Sculpin X

Cottidae Artediellus scaber Hamecon X X

Cottidae Enophrys diceraus Antlered Sculpin X X

Cottidae Gymnocanthus tricuspis Arctic Staghorn Sculpin X X

Cottidae Hemilepidotus papilio Butterfly Sculpin X

Cottidae Icelus bicornis Twohorn Sculpin X X

Cottidae Icelus spatula Spatulate Sculpin X X

Cottidae Icelus spiniger Thorny Sculpin X

Cottidae Megalocottus platycephalus Belligerent Sculpin X X

Cottidae Microcottus sellaris Brightbelly Sculpin X

Cottidae Myoxocephalus jaok Plain Sculpin X X

Cottidae Myoxocephalus polyacanthocephalus Great Sculpin X

Cottidae Myoxocephalus quadricornis Fourhorn Sculpin X X

Cottidae Myoxocephalus scorpioides Arctic Sculpin X X

Cottidae Myoxocephalus scorpius Shorthorn Sculpin X X

Cottidae Trichocottus brashnikovi Hairhead Sculpin X X

Cottidae Triglops nybelini Bigeye Sculpin X X

Cottidae Triglops pingelii Ribbed Sculpin X X

Hemitripteridae Blepsias bilobus Crested Sculpin X

Hemitripteridae Nautichthys pribilovius Eyeshade Sculpin X X

Psychrolutidae Cottunculus microps Polar Sculpin X

Psychrolutidae Eurymen gyrinus Smoothcheek Sculpin X

Agonidae Aspidophoroides monopterygius Alligatorfish X X

Agonidae Aspidophoroides olrikii Arctic Alligatorfish X X
Chapter 2 27

Family Scientific name Common name

Confirmed occurrence

Chukchi Sea Beaufort Sea

Agonidae Hypsagonus quadricornis Fourhorn Poacher X

Agonidae Leptagonus decagonus Atlantic Poacher X X

Agonidae Occella dodecaedron Bering Poacher X

Agonidae Pallasina barbata Tubenose Poacher X

Agonidae Podothecus veternus Veteran Poacher X X

Cyclopteridae Eumicrotremus andriashevi Pimpled Lumpsucker X

Cyclopteridae Eumicrotremus derjugini Leatherfin Lumpsucker X X

Liparidae Careproctus reinhardti Sea Tadpole X X

Liparidae Liparis bathyarcticus Nebulous Snailfish X X

Liparidae Liparis fabricii Gelatinous Seasnail X X

Liparidae Liparis gibbus Variegated Snailfish X X

Liparidae Liparis tunicatus Kelp Snailfish X X

Liparidae Paraliparis bathybius Black Seasnail X X

Zoarcidae Gymnelus hemifasciatus Halfbarred Pout X X

Zoarcidae Gymnelus viridis Fish Doctor X X

Zoarcidae Lycenchelys kolthoffi Checkered Wolf Eel X

Zoarcidae Lycodes adolfi Adolf’s Eelpout X

Zoarcidae Lycodes eudipleurostictus Doubleline Eelpout X

Zoarcidae Lycodes frigidus Glacial Eelpout X X

Zoarcidae Lycodes jugoricus Shulupaoluk X

Zoarcidae Lycodes marisalbi White Sea Eelpout X

Zoarcidae Lycodes mucosus Saddled Eelpout X X

Zoarcidae Lycodes palearis Wattled Eelpout X X

Zoarcidae Lycodes pallidus Pale Eelpout X

Zoarcidae Lycodes polaris Polar Eelpout X X

Zoarcidae Lycodes raridens Marbled Eelpout X X

Zoarcidae Lycodes reticulatus Arctic Eelpout X

Zoarcidae Lycodes rossi Threespot Eelpout X

Zoarcidae Lycodes sagittarius Archer Eelpout X

Zoarcidae Lycodes seminudus Longear Eelpout X X

Zoarcidae Lycodes squamiventer Scalebelly Eelpout X

Zoarcidae Lycodes turneri Estuarine Eelpout X X

Stichaeidae Acantholumpenus mackayi Blackline Prickleback X

Stichaeidae Anisarchus medius Stout Eelblenny X X

Stichaeidae Chirolophis snyderi Bearded Warbonnet X

Stichaeidae Eumesogrammus praecisus Fourline Snakeblenny X X

Stichaeidae Leptoclinus maculatus Daubed Shanny X X

Stichaeidae Lumpenus fabricii Slender Eelblenny X X

Stichaeidae Lumpenus sagitta Snake Prickleback X

Stichaeidae Stichaeus punctatus Arctic Shanny X X

Pholidae Pholis fasciata Banded Gunnel X

Anarhichadidae Anarhichas denticulatus Northern Wolffish X X

Anarhichadidae Anarhichas orientalis Bering Wolffish X X

Zaproridae Zaprora silenus Prowfish X

Ammodytidae Ammodytes hexapterus Arctic Sand Lance X X

Pleuronectidae Hippoglossoides robustus Bering Flounder X X

Pleuronectidae Hippoglossus stenolepis Pacific Halibut X

Pleuronectidae Limanda aspera Yellowfin Sole X X

Pleuronectidae Limanda proboscidea Longhead Dab X X

Pleuronectidae Limanda sakhalinensis Sakhalin Sole X

Pleuronectidae Liopsetta glacialis Arctic Flounder X X

Pleuronectidae Platichthys stellatus Starry Flounder X X

Pleuronectidae Pleuronectes quadrituberculatus Alaska Plaice X

Pleuronectidae Reinhardtius hippoglossoides Greenland Halibut X X

 Totals: 109 97 83

Table 2.1. Inventory of confirmed species of Arctic marine fish in the United States Chukchi and Beaufort Seas, Alaska.—Continued
28 Alaska Arctic Marine Fish Ecology Catalog

Table 2.2. Changes in occurrence and geographic distribution of Arctic marine fishes in the Chukchi and Beaufort Seas since 2002.

[Changes were made since publication of Mecklenburg and others (2002)]

Scientific name Common name New occurrence Geographic distribution

Range change No change

Entosphenus tridentatus Pacific Lamprey X

Lethenteron camtschaticum Arctic Lamprey X

Squalus suckleyi Spotted Spiny Dogfish X

Somniosus pacificus Pacific Sleeper Shark X X

Amblyraja hyperborea Arctic Skate X X

Bathyraja parmifera Alaska Skate X X

Clupea pallasii Pacific Herring X

Hypomesus olidus Pond Smelt X

Mallotus catervarius Pacific Capelin X

Osmerus dentex Arctic Smelt X

Coregonus autumnalis Arctic Cisco X

Coregonus laurettae Bering Cisco X

Coregonus nasus Broad Whitefish X

Coregonus pidschian Humpback Whitefish X

Coregonus sardinella Least Cisco X

Oncorhynchus gorbuscha Pink Salmon X

Oncorhynchus keta Chum Salmon X

Oncorhynchus kisutch Coho Salmon X

Oncorhynchus nerka Sockeye Salmon X

Oncorhynchus tshawytscha Chinook Salmon X

Salvelinus malma Dolly Varden X

Stenodus leucichthys Inconnu X

Benthosema glaciale Glacier Lanternfish X

Arctogadus glacialis Ice Cod X

Boreogadus saida Arctic Cod X

Eleginus gracilis Saffron Cod X

Gadus chalcogrammus Walleye Pollock X

Gadus macrocephalus Pacific Cod X

Gasterosteus aculeatus Threespine Stickleback X

Pungitius pungitius Ninespine Stickleback X

Hexagrammos stelleri Whitespotted Greenling X

Artediellus ochotensis Okhotsk Hookear Sculpin X

Artediellus scaber Hamecon X

Enophrys diceraus Antlered Sculpin X

Gymnocanthus tricuspis Arctic Staghorn Sculpin X

Hemilepidotus papilio Butterfly Sculpin X

Icelus bicornis Twohorn Sculpin X

Icelus spatula Spatulate Sculpin X

Icelus spiniger Thorny Sculpin X X

Megalocottus platycephalus Belligerent Sculpin X

Microcottus sellaris Brightbelly Sculpin X

Myoxocephalus jaok Plain Sculpin X

Myoxocephalus polyacanthocephalus Great Sculpin X X

Myoxocephalus quadricornis Fourhorn Sculpin X

Myoxocephalus scorpioides Arctic Sculpin X

Myoxocephalus scorpius Shorthorn Sculpin X

Trichocottus brashnikovi Hairhead Sculpin X

Triglops nybelini Bigeye Sculpin X X

Triglops pingelii Ribbed Sculpin X

Blepsias bilobus Crested Sculpin X

Nautichthys pribilovius Eyeshade Sculpin X

Cottunculus microps Polar Sculpin X

Eurymen gyrinus Smoothcheek Sculpin X

Aspidophoroides monopterygius Alligatorfish X

Aspidophoroides olrikii Arctic Alligatorfish X
Chapter 2 29

Scientific name Common name New occurrence Geographic distribution

Range change No change

Hypsagonus quadricornis Fourhorn Poacher X X

Leptagonus decagonus Atlantic Poacher X

Occella dodecaedron Bering Poacher X

Pallasina barbata Tubenose Poacher X

Podothecus veternus Veteran Poacher X

Eumicrotremus andriashevi Pimpled Lumpsucker X

Eumicrotremus derjugini Leatherfin Lumpsucker X

Careproctus reinhardti Sea Tadpole X X

Liparis bathyarcticus Nebulous Snailfish X X

Liparis fabricii Gelatinous Seasnail X

Liparis gibbus Variegated Snailfish X

Liparis tunicatus Kelp Snailfish X

Paraliparis bathybius Black Seasnail X X

Gymnelus hemifasciatus Halfbarred Pout X

Gymnelus viridis Fish Doctor X

Lycenchelys kolthoffi Checkered Wolf Eel X X

Lycodes adolfi Adolf’s Eelpout X X

Lycodes eudipleurostictus Doubleline Eelpout X

Lycodes frigidus Glacial Eelpout X X

Lycodes jugoricus Shulupaoluk X X

Lycodes marisalbi White Sea Eelpout X X

Lycodes mucosus Saddled Eelpout X

Lycodes palearis Wattled Eelpout X

Lycodes pallidus Pale Eelpout X X

Lycodes polaris Polar Eelpout X

Lycodes raridens Marbled Eelpout X

Lycodes reticulatus Arctic Eelpout X X

Lycodes rossi Threespot Eelpout X

Lycodes sagittarius Archer Eelpout X

Lycodes seminudus Longear Eelpout X

Lycodes squamiventer Scalebelly Eelpout X

Lycodes turneri Estuarine Eelpout X

Acantholumpenus mackayi Blackline Prickleback X

Anisarchus medius Stout Eelblenny X

Chirolophis snyderi Bearded Warbonnet X

Eumesogrammus praecisus Fourline Snakeblenny X

Leptoclinus maculatus Daubed Shanny X

Lumpenus fabricii Slender Eelblenny X

Lumpenus sagitta Snake Prickleback X X

Stichaeus punctatus Arctic Shanny X

Pholis fasciata Banded Gunnel X

Anarhichas denticulatus Northern Wolffish X X

Anarhichas orientalis Bering Wolffish X

Zaprora silenus Prowfish X X

Ammodytes hexapterus Arctic Sand Lance X

Hippoglossoides robustus Bering Flounder X

Hippoglossus stenolepis Pacific Halibut X

Limanda aspera Yellowfin Sole X

Limanda proboscidea Longhead Dab X

Limanda sakhalinensis Sakhalin Sole X

Liopsetta glacialis Arctic Flounder X

Platichthys stellatus Starry Flounder X

Pleuronectes quadrituberculatus Alaska Plaice X

Reinhardtius hippoglossoides Greenland Halibut X

 Totals: 109 20 59 50

Table 2.2. Changes in occurrence and geographic distribution of Arctic marine fishes in the Chukchi and Beaufort Seas since 2002.—

Continued
30 Alaska Arctic Marine Fish Ecology Catalog

Species Range Extensions

Comparing Arctic marine fish distributions as described 

by recent studies to similar historical information reveals some 

significant changes. Some of the changes in distributional 

patterns relate to increased sampling and new discovery and 

others to changes in species classifications and nomenclature. 

Since 2002, 20 new species were confirmed from the Chukchi 

and Beaufort Seas. By family, the new Alaska records were 

reported from Somniosidae (1), Rajidae (2), Cottidae (3), 

Agonidae (1), Liparidae (3), Zoarcidae (7), Stichaeidae (1), 

Anarhichadidae (1), and Zaproridae (1). The new Alaskan 

records are valid range extensions resulting from BOEMsponsored surveys by NOAA (Logerwell and others, 2009; 

Rand and Logerwell, 2011) and University of AlaskaFairbanks (UAF) and NOAA’s RUSALCA (Mecklenburg and 

others, 2014; Mecklenburg and Steinke, 2015) or changes 

in taxonomy and distribution reported by Mecklenburg and 

others (2011, 2015, 2016). 

Extensions to the known ranges for five marine species 

were reported from trawl collections in the Beaufort Sea 

by Rand and Logerwell (2011): Walleye Pollock (Gadus 

chalcogrammus), Pacific Cod (G. macrocephalus), Festive 

Snailfish (Liparis marmoratus), Bigeye Sculpin (Triglops 

nybelini), and Eyeshade Sculpin (Nautichthys pribilovius). 

Apparently, Rand and Logerwell (2011) did not notice that the 

Antlered Sculpin (Enophrys diceraus) listed in their tables also 

was an extension of this species’ known range in the Beaufort 

Sea. This extension was confirmed by Maslenikov and others 

(2013). 

Additional information about the Rand and Logerwell 

(2011) collection includes:

• Walleye Pollock (Gadus chalcogrammus): This 

probably was the first published record from the 

Beaufort Sea for G. chalcogrammus, since the previous 

record from Elson Lagoon (near Barrow, Alaska) was 

shown to pertain to Boreogadus saida (Mecklenburg 

and others, 2011, Online Resource 1, p. 15). Rand and 

Logerwell (2011, p. 484) reported this occurrence from 

Elson Lagoon incorrectly. 

• Pacific Cod (Gadus macrocephalus): “Major changes 

in the composition of gadine genera in recent years 

mean patterns of species distribution are different” 

(Mecklenburg and others, 2011, Online Resource 

1, p. 120). Historically, G. macrocephalus from the 

Beaufort Sea were identified as G. ogac (Mecklenburg 

and others, 2002). Owing to recent DNA studies 

and similarity in morphology, and the consequent 

synonymization of G. ogac in G. macrocephalus, this 

species is now recognized to have a broad distribution 

from the Chukchi Sea and continuing eastward in the 

Arctic across Canada to Greenland and the White Sea 

(Mecklenburg and others, 2011, Online Resource 1, 

p. 120). Given this information, the range extension 

reported by Rand and Logerwell (2011) was not truly 

an extension of known range.

• Bigeye Sculpin (Triglops nybelini); From Maslenikov 

and others, 2013, p. 12): “An Arctic species reported 

from northwest of Alaska in the international waters 

of the Chukchi Borderland (Mecklenburg et al., 2011), 

the Beaufort Sea off Alaska (eastward to 71.21836N, 

149.90316W by benthic trawl in 2011; C.W. 

Mecklenburg, personnel [sic.] communication), and 

from western Canada in Mackenzie Bay and Amundsen 

Gulf eastward around the Arctic to the Laptev Sea 

(Pietsch, 1993). Captured in a midwater tow, the 

specimens reported here from the Beaufort Sea east of 

Point Barrow are vouchers for the 1st Alaskan records 

cited by Rand and Logerwell (2011).”

• Eyeshade Sculpin (Nautichthys pribilovius); From 

Maslenikov and others, 2013, p. 12) “Known from 

the Chukchi Sea off Wainwright south through the 

western and eastern Bering Sea, Sea of Okhotsk, 

and Sea of Japan, and the Aleutian and Commander 

Islands to southeastern Alaska, the record cited here 

provides a voucher for the northern range extension of 

Nautichthys pribilovius into the Beaufort Sea cited by 

Rand and Logerwell (2011).”

As an additional note, the single specimen reported by 

Rand and Logerwell (2011) from their 2008 cruise is the first 

record from the Beaufort Sea, as described by Mecklenburg 

and others (2011, Online Resource 1, p. 27). Others have 

been collected in the same area since then (Mecklenburg and 

others, 2016).

• Festive Snailfish: Reports of L. marmoratus in 

the Bering Sea and the Alaskan Arctic have been 

questioned because the voucher specimens are similar 

morphologically and genetically to L. tunicatus

(Mecklenburg and others, 2011, 2016; Mecklenburg 

and Steinke, 2015). The DNA sequences from all 

specimens collected in the Bering and Chukchi Seas 

in recent years with the external appearance described 

for L. marmoratus (Mecklenburg and others, 2002), are 

identical to sequences from L. tunicatus. Mecklenburg 

and others (2011) did not count this species among 

those considered valid in their review and list of Arctic 

marine fishes. The specimens reported by Rand and 

Logerwell (2011), which would be the only records 

from the U.S. Arctic, look morphologically the same as 

the barcoded specimens2

. 2

Mecklenburg and others (2011) questioned the validity of collection 

records for L. marmoratus from the Beaufort Sea. DNA barcoding indicate the 

species is not valid from the Chukchi or Beaufort Seas but may be found in 

the Sea of Okhotsk (Mecklenburg and Steinke, 2015). 
Chapter 2 31

• Longear Eelpout (Lycodes seminudus); from 

Maslenikov and others (2013): L. seminudus is an 

Arctic species known from the Beaufort Sea and the 

Chukchi slope north of Alaska (Mecklenburg and 

others, 2011), as well as the Norwegian and Kara Seas. 

The record cited here extends the range 200 km west 

in the Beaufort Sea and represents only the second 

Alaskan record.

Mecklenburg and others (2016, p. 197) reported a 

significant extension in the northern distribution of the 

Bearded Warbonnet in sampling conducted off Pt. Barrow:

• Bearded Warbonnet (Chirolophis snyderi): “The 

northernmost documented locality is Point Barrow, 

Alaska, at 71°23′N, 156°29′W, where juvenile 

specimens were taken in a beach seine in 2012 

(e.g., CAS 237946, 2 specimens, each 31 mm). The 

previously reported northernmost record was from 

northwest of Point Franklin in the eastern Chukchi Sea 

(Mecklenburg et al., 2002, 2011; UAM 4582).”

These descriptions highlight the dynamic nature and 

taxonomic issues associated with documenting or confirming 

range extensions from the Arctic. In many instances, Alaska 

records seem to be new because there has been so little 

previous sampling. Researchers often are not able to review 

historical collections in light of new taxonomic knowledge. 

In fairness, the historical data are often difficult to access and 

interpret because of documented changes in taxonomy and 

geographical distribution. To illustrate, Rand and Logerwell 

(2011) noted that Bering Flounder (Hippoglossoides robustus) 

was not reported in Arctic fish sampling by Frost and Lowry 

(1983) in 1977, nor was it reported north of Point Barrow by 

Mecklenburg and others (2002). However, additional research 

by Mecklenburg and others (2011) discovered that this species 

was distributed eastward through the Beaufort Sea to Bathurst 

Inlet (Canadian Arctic) as early as 1965. 

The Walleye Pollock catches reported by Logerwell and 

others (2011) were to the north of all larval, age-1, and adult 

pollock catches depicted by Wyllie Echeverria (1996) for the 

northeastern Chukchi Sea and the geographic distributions 

described by Mecklenburg and others (2002). The new 

Alaskan records, and size of fish captured, raise the possibility 

that this species, particularly juvenile pollock, may have been 

misidentified in earlier Beaufort Sea surveys as Polar Cod3

. It 

also raises questions about their potential origins and dispersal 

into this part of the Arctic Ocean. Walleye Pollock are 

captured off Norway and thus their dispersal into the Beaufort 

Sea with possible genetic interchange along the upper slope 

is plausible. Cold temperatures are constraints to northerly 

movements and abundance of pollock; therefore, large 

expansions and swarms, such as those reported for Arctic Cod 

(Boreogadus saida) by Crawford and others (2012), would not 

be expected under present environmental conditions. 

Other range extensions are reported in Love and others 

(2005), Mecklenburg and others (2011), Maslenikov and 

others (2013), Mecklenburg and Steinke (2015), or evaluated 

more conventionally in the Pacific Arctic Marine Fishes guide 

(Mecklenburg and others, 2016). In addition to the species 

discussed from the Rand and Logerwell (2011) collection, an 

annotated list of range extensions provides new distributional 

information with supporting documentation as available, or 

appropriate, for citation at this time. The new information 

results from an increased Arctic research sampling emphasis 

in offshore marine habitats in the Chukchi and Beaufort 

Seas. The annotated list of extensions is represented by 

lesser-known marine fishes having no commercial value that 

researchers are aware of and are presented as they relate to 

the (1) Beaufort Sea, (2) Chukchi Sea, and (3) Chukchi and 

Beaufort Seas:

1. From the Beaufort Sea:

• Arctic Skate (Amblyraja hyperborea): Mecklenburg 

and others (2016) report this species to be circumpolar; 

found in polar basins southward to Beaufort Sea 

of Alaska and Western Canada at depths ranging 

from 92 to 2,925 m. Size information is reported in 

Wienerroither and others (2011).

• Plain Sculpin (Myoxocephalus jaok): New record 

identified from the Beaufort Sea in 2011 (Mecklenburg 

and others, 2016).

• Alligatorfish (Aspidophoroides monopterygius):

Identified by Catherine W. Mecklenburg from a 

specimen voucher in the NOAA RACE groundfish 

surveys 2012 survey collections (Mecklenburg and 

others, 2016).

• Black Seasnail (Paraliparis bathybius): Previously 

reported from the western Arctic Canada Basin north 

of Alaska in 2005 (Mecklenburg and others, 2016). 

Alaskan collections were from slope and deep waters 

of the Beaufort Sea in 2011.

• Checkered Wolf Eel (Lycenchelys kolthoffi): One 

specimen was taken in the Alaskan Beaufort Sea near 

the U.S.–Canada border at 70°28′N, 141°09′W at 

a depth of 500 m (Mecklenburg and others, 2016). 

Otherwise, the species lives eastward from that point 

to Greenland and on to the Kara Sea. It is common 

at least around Greenland. It is mesobenthic, lives 

on sandy or muddy or mixed soft-stony sea floors, at 

depths of 202–930 m and temperatures of -0.9–4.1 °C. 

Maximum size is 29 cm TL.

3

This is a common misidentification in museum collections (Mecklenburg 

and others, 2011; Mecklenburg and Steinke, 2015). 
32 Alaska Arctic Marine Fish Ecology Catalog

At least five new occurrences of eelpout species (table 

2.2) from the Beaufort Sea sampling were confirmed in 

the processing of voucher specimens from University of 

Alaska Fairbanks fishery collections between 2010 and 2011 

(Mecklenburg and Steinke, 2015; Mecklenburg and others, 

2016). Specimens collected from the WEBSEC-71 survey in 

the Beaufort Sea (U.S. Coast Guard Cutter [USCGC] Glacier; 

Hufford, 1974) were evaluated in 2007 and new information 

gleaned from the specimens was reported by Mecklenburg 

and others (2016). Additional records and information on the 

following species may also be found in Mecklenburg and 

others (2016):

• Adolf’s Eelpout (Lycodes adolfi): A voucher specimen 

collected in the Beaufort Sea in University of Alaska 

Fairbanks surveys conducted between 2010 and 2012 

(Mecklenburg and others, 2016). This species was also 

sampled on Chukchi slope, outside the 200-mi limit, 

by 2009 RUSALCA (Russian-American Long-term 

Census of the Arctic, 2009); no other records near or in 

the Chukchi Sea); see Mecklenburg and others (2011, 

Online Reference 1).

• Glacial Eelpout (Lycodes frigidus); From 

Mecklenburg and others (2011, Online Resource 1, 

p. 39): “Arctic; Makarov Basin northwest of Ellesmere 

Island, deep slopes and basins of Greenland and 

Norwegian Seas, Nansen to Canada basins off Laptev, 

East Siberian, Chukchi, and Beaufort Seas. Benthic, 

at depths of 475 m to 3,000 m, rare in depths less than 

1,000 m. The first vouchered collection from north of 

Alaska (California Academy of Sciences, Department 

of Ichthyology, San Francisco, USA CAS 230372; 

2 specimens, 53–76 mm) was made by remotely 

operated vehicle (ROV) and suction sampler in 2005 

north-northwest of Point Barrow at a depth of about 

2,500 m.”

• Shulupaoluk (Lycodes jugoricus): The University of 

Alaska Fairbanks Beaufort Sea 2012 voucher specimen 

collection includes several L. jugoricus. (Mecklenburg 

and others, 2016).

• White Sea Eelpout (Lycodes marisalbi); From 

Mecklenburg and others (2011, Online Resource 1), 

p. 40: “Arctic; White Sea and Beaufort Sea. Benthic, 

at depths of 6–335 m. Records from the Beaufort Sea 

off Alaska identified as L. pallidus (e.g., Mecklenburg 

et al. 2002) are L. marisalbi (Canadian Museum of 

Nature, Ottawa, Canada CMNFI 1974-279, 1974-285, 

1974-287, 1978-302; CWM, 15 May 2008).”

• Arctic Eelpout (Lycodes reticulatus): Mecklenburg 

and Steinke (2015) investigated the validity of species 

in the family Lycodes from the Pacific Arctic Region 

using morphological observations and DNA barcoding 

sequences (http://dx.doi.org/10.5883/DS-LYCODES). 

Their genetic analysis revealed that specimens 

identified as L. rossi always fell into the Beaufort Sea 

clade for L. reticulatus.

• Threespot Eelpout (Lycodes rossi): The inclusion 

of this species is based on information presented in 

Mecklenburg and others (2016). 

• Longear Eelpout (Lycodes seminudus): Based on 

UAF’s fishery surveys in the Beaufort Sea (2011–12 

records), an offshore pattern reflects a geographic 

distribution along the shelf deeper than 200–300 m to 

about 1,400 m. The species occurs only in the offshoremost tracts in the Beaufort Sea lease area. 

• Scalebelly Eelpout (Lycodes squamiventer): Catherine 

W. Mecklenburg tentatively identified this species from 

specimens collected from the Beaufort Sea in 2012 

(Mecklenburg and others, 2016).

2. From the Chukchi Sea:

• Ice Cod (Arctogadus glacialis): Identified from 

samples collected from the Chukchi Sea (Mecklenburg 

and others, 2016). 

• Sea Tadpole (Careproctus reinhardti): Geographic 

distribution from Baffin Bay and Davis Strait, off 

eastern Greenland, off northern Russia (Kara, and 

Laptev Seas), and Chukchi Sea slope and northern 

Bering Sea (Mecklenburg and others, 2016). Benthic, 

at depth of 100–1,840 m. Preliminary genetic data 

suggest the sea tadpole could be a species complex 

(Mecklenburg and others, 2016).

• Great Sculpin (Myoxocephalus polyacanthocephalus):

Okhotsk Sea and eastern Japan Sea to Commander–

Aleutian Island chain to Chukchi Sea, 70°20′N, 

163°06′W (Mecklenburg and others, 2016) to southern 

Puget Sound, Washington. Benthic; intertidal and to 

825 m. 

• Prowfish (Zaprora silenus): Hokkaido, Japan, and 

Sea of Okhotsk to Bering Sea (Mecklenburg and 

others, 2002) and to southeastern Chukchi Sea (west 

of Kivalina, 67°32′N, 165°54′W) (Mecklenburg and 

others, 2011) and Aleutian Islands (Mecklenburg and 

others, 2002) to San Miguel Island, southern California 

(Allen and Smith, 1988). At depths of 10–801 m. 

Adults found near bottom, young fish often collected 

near surface (Mecklenburg and others, 2002). 
Chapter 2 33

3. From the Chukchi and Beaufort Seas:

• Bigeye Sculpin (Triglops nybelini): Arctic Ocean, 

practically circumpolar, Chukchi Sea north of Alaska, 

western Beaufort Sea slope between 152°W and 

155°W (Mecklenburg and others, 2011). Benthic, at 

depths of 30–1,354 m, usually deeper than 200 m 

(Mecklenburg and others, 2016).

• Nebulous Snailfish (Liparis bathyarcticus):

Circumpolar, Beaufort and Chukchi Seas to 

southeastern Bering Sea (Mecklenburg and others, 

2016). Benthic at depths of about 12–510 m 

(Mecklenburg and others, 2016).

Probable Occurrences in Alaska Arctic 

More species are known to occur in the study area than 

are described in the species accounts in this catalog because 

the taxonomic examinations of these fish are in an early 

stage, and valid identifications are not possible without all 

results from the multi-faceted research (that is, meristic, 

morphological, and molecular genetic). The ongoing research 

approach will include formal descriptions of new species 

and documentation of significant distributional changes. 

For instance, Mecklenburg and others (2011) reported an 

unidentified sculpin, Icelus sp., from the Chukchi Sea (editor’s 

note: recently confirmed by Mecklenburg and Steinke, 2015), 

and Rand and Logerwell (2011) reported an unidentified 

snailfish, Careproctus sp. cf. rastrinus, from the Beaufort Sea. 

For this study, we determined that the biological information 

available for these two species to be too scant and preliminary 

to permit construction of dependable species accounts. Species 

accounts were not developed for Sominiosus pacificus, 

Bathyraja parmifera, or Lycodes pallidus due to their recent 

confirmation from the Chukchi Sea (December 2015); 

information about these species is available in the Fishes of 

Alaska and species accounts are presented in the Pacific Arctic 

Marine Fishes guide (Mecklenburg and others, 2016).

The additional species identified here are likely to be 

confirmed from Alaska waters. The information presented was 

primarily from an ongoing effort by several authors of this 

report to update Love and others (2005):

• Atlantic Hookear Sculpin (Artediellus atlanticus); 

From: Mecklenburg and others, (2011, Online 

Resource 1, p. 20): Arctic–Boreal; to southern 

Baffin Island, northwest and northeast Greenland, 

Iceland, Jan Mayen, Faroe Islands, Norway, and 

Barents Sea, northern Kara Sea, and northwestern 

Laptev Sea; upper slope off the Chukchi Sea; south 

to Cape Cod, the Skagerrak, Irish Sea, and coasts of 

Ireland (Fedorov, 1986, p. 1,245; Mecklenburg and 

others, 2011, fig. 2; this study). Benthic, at depths of 

11–1,366 m. Found on the Chukchi slope (Chukchi 

Sea Borderlands, outside U.S. EEZ) in 2009: two at 

74°29′N, 165°58′W, depth 365–370 m (California 

Academy of Sciences, Department of Ichthyology, 

San Francisco, USA CAS 228530, fig. 3; Zoological 

Institute of the Russian Academy of Sciences, St. 

Petersburg, Russia ZIN 54840); and five at 74°07′N, 

166°00′W, depth 227–236 m (CAS 228533, University 

of Alaska Museum, University of Alaska Fairbanks 

5534). This is the first record from the East SiberianChukchi-Beaufort region. The specimens fit the 

description of A. a. corniger (Andriashev, 1954) from 

the Kara and Laptev Seas, Russia, with large parietal 

spines, but the most recent review of the species in 

the western Atlantic (Van Guelpen, 1986) determined 

this character not to discriminate between taxonomic 

groups and did not recognize any subspecies of 

A. atlanticus. Parietal spine size resembled a cline, 

with the most strongly developed found in northern 

parts of the range. The large size fish length (Total 

Length) of specimens from the Chukchi Sea (as long 

as 153 mm) agrees with Van Guelpen’s determination 

that the largest specimens are found in the north. The 

four Co-1 (Cytochrome C Oxidase 1) gene sequences 

from the Chukchi Sea slope specimens match those of 

three specimens from southern Baffin Bay (Canadian 

Museum of Nature, Ottawa, Canada CMNFI 2002-

0028.3, 2002-0031.1, 2002-0033.1; 68°26′–69°37′N). 

The species was confirmed from the Chukchi 

Borderland by Mecklenburg and others (2014).

• Spinyhook Sculpin (Artediellus gomojunovi): From 

Mecklenburg and others (2011, Online Resource 1)—

Boreal Pacific north to Gulf of Anadyr, Russia, 

and Bering Strait. Benthic, at depths of 37–380 m 

(Mecklenburg and others, 2002). Bering Sea records 

are no deeper than 90 m. A specimen collected in 

1950 near Point Barrow (Smithsonian National 

Museum of Natural History, Washington, D.C., USNM 

152901) and tentatively identified as A. gomojunovi

(Andriashev, 1961 as cited by Mecklenburg and others, 

2011 Online Resource 1) was misplaced (Mecklenburg 

and others, 2002), but was found in 2005, and 

subsequently identified as A. scaber. The current 

northernmost record of A. gomojunovi is a specimen 

caught in 1933 at Bering Strait (ZIN 33093). 

• Hookhorn Sculpin (Artediellus pacificus): From 

Mecklenburg and others, (2011, Online Resource 1)—

Boreal Pacific north to southern Gulf of Anadyr and 

northeast of St. Lawrence Island, Bering Sea. Benthic, 

at depths of 15–250 m (Mecklenburg and others, 

2002). The northern records are not recent: Zoological 
34 Alaska Arctic Marine Fish Ecology Catalog

Institute of the Russian Academy of Sciences, 

St. Petersburg, Russia ZIN 33657, Gulf of Anadyr at 

63°53′N, 178°48′W, depth 87 m, collected in 1952; 

and University of Washington Fish Collection at the 

Burke Museum, Seattle, WA UW 11740, at 63°40′N, 

167°28′W, depth 22–40 m, in 1949. 

• Leister Sculpin (Enophrys lucasi): Mecklenburg 

and others (2016, p. 55) described the distribution 

as “Southeastern Kamchatka, Commander Islands, 

Aleutian Islands, Gulf British Columbia, Bering Sea 

north to Bering Strait. The distribution is difficult to 

assess due to misidentifications in the literature and 

museum collections, and the distribution depicted on 

our map may not be accurate north of the Aleutian 

Islands and Gulf of Alaska.”

• Threaded Sculpin (Gymnocanthus pistilliger): 

Mecklenburg and others (2016) suggested this species 

to be common from Bering Strait. The northernmost 

record is CAS 230370, taken in 2006 west of Port 

Clarence at 65°21′N, 167°41′W. Records from Port 

Clarence include CAS 230369, CAS 230370, CMNFI 

1958-0093, UAM 1201, and UW 41676). The several 

records from Port Clarence and nearby suggest it also 

occurs in the southeastern Chukchi Sea and there 

are reliable identifications northward to Kotzebue 

Sound, but distribution in the Chukchi Sea has not 

been documented with voucher specimens. Historical 

records from the Chukchi Sea remain unverified or 

have been reidentified (Mecklenburg and others, 2016).

• Yellow Irish Lord (Hemilepidotus jordani): From 

Mecklenburg and others, (2011, Online Resource 1)—

Boreal Pacific north to Bering Strait, perhaps to 

southeastern Chukchi Sea. Benthic, at depths of 

15–400 m or more, typically shallower than 150 m. 

Reported from Chukchi Sea (Allen and Smith, 1988), 

but voucher specimens are lacking. A specimen from 

the Chukchi Sea southwest of Kivalina identified as 

H. jordani (University of British Columbia, Vancouver, 

B.C., UBC 61-0064) is H. papilio (Mecklenburg and 

others, 2016). The northernmost verified record is 

CAS 230368, two juveniles (77–82 mm) caught in 

2006 near eastern Bering Strait at 64°59’N, 168°30’W, 

by bottom trawl in 25 m of water.

• Frog Sculpin (Myoxocephalus stelleri): This species, 

once believed to possibly be present rarely in Alaskan 

waters (Mecklenburg and others, 2002), has been 

determined from evaluation of all pertinent literature 

and voucher records identified as this species to be 

absent from the eastern Pacific and Alaskan waters 

(Mecklenburg and others, 2011, 2016). See also 

Mecklenburg and Steinke (2015, p. 176).

• Polar Sculpin (Cottunculus microps): One record 

is from the Arctic Ocean on the slope north of the 

Chukchi Sea, outside the 200-mi limit. Taxonomic 

revision is in progress; confirmed from the Chukchi 

Borderland (Mecklenburg and others, 2014).

• Threaded Sculpin (Gymnocanthus pistilliger): 

Mecklenburg and others (2016) extended the range for 

this species into the southeastern Chukchi Sea based on 

“reliable” identifications but not confirmed specimen 

vouchers. Although this species is likely to occur in the 

region, our list includes only species whose presence is 

known from voucher specimens.

• Northern Sand Lance (Ammodytes dubius): 

Predominantly in boreal western Atlantic; Canadian 

Arctic from Beaufort Sea to Baffin Bay, and Davis 

Strait. Benthic, shallows to 108 m (Coad and Reist, 

2004). These authors also listed Alaska but provided 

no specific locality.

Other Changes in Species Distributions 

In addition to the new occurrences and range extensions 

in table 2.2, mapping for this study (chapter 3) documents 

expansions in the known geographic distributions of another 

45 species in this updating of information presented in the 

Fishes of Alaska (Mecklenburg and others, 2002). These shifts 

in range result from the new information and knowledge 

gained from recent field surveys and discoveries, improved 

identification techniques, evaluation of museum collections, 

and the wide-ranging search of data and literature for this 

study. In light of the recent field sampling, some changes in 

geographic, depth, or both, were realistic expectations. Marine 

ecosystems are dynamic and fish distributions may contract 

and expand due to various factors including, but not limited to, 

temperature changes, current patterns, changes in population 

size, and changes in predator and prey distribution.
Chapter 2 35

Table 2.3. Changes in scientific and common names of Arctic marine fishes in the Chukchi and Beaufort Seas since 2002.

[Changes were made since publication of Mecklenburg and others (2002). Reference: Information is critical for users. In some instances, specialists may 

disagree (“not clearly distinguishable” as supporting genetic information may not be available; for example, Liparis herchelinus not clearly distinguished from 

L. tunicatus), or species may be part of an evolving complex (for example, Lycodes rossi and L. reticulatus)]

Family Common or

scientific name

Change in common or scientific name Reference

From To

Squalidae Spotted Spiny Dogfish Squalus acanthias Squalus suckleyi Lynghammar and others, 2012

Osmeridae Capelin Mallotus villosus Mallotus catervarius Mecklenburg and Steinke, 2015

Osmeridae Mallotus catervarius Capelin Pacific Capelin Mecklenburg and Steinke, 2015

Osmeridae Arctic Smelt Osmerus mordax Osmerus dentex Mecklenburg and others, 2011

Gadidae Arctogadus glacialis Polar Cod Ice Cod Jørgen Schou Christiansen, oral 

commun, 20131

Gadidae Walleye Pollock Theragra chalcogramma Gadus chalcogrammus Mecklenburg and others, 2011; 

Page and others, 2013

Gadidae Pacific Cod Gadus ogac Gadus macrocephalus Mecklenburg and others, 2011; 

Page and others, 2013

Psychrolutidae Polar Sculpin Cottunculus sadko Cottunculus microps Byrkjedal and others, 2014

Agonidae Arctic Alligatorfish Ulcina olrikii Aspidophoroides olrikii Mecklenburg and others, 2011; 

Page and others, 2013

Ammodytidae Ammodytes hexapterus Pacific Sand Lance Arctic Sand Lance Orr and others, 2015

Pleuronectidae Arctic Flounder Pleuronectes glacialis Liopsetta glacialis Eschmeyer and others, 2015; 

Mecklenburg and others, 2016

1

From Jørgen Schou Christiansen, University of Tromsø, Norway, written commun., May 12, 2013, regarding common names for A. glacialis and B. saida: 

“The scientific name is conclusive and should follow the vernacular name at first mention for these species. Whenever vernacular names are used, we suggest 

‘ice cod’ for Arctogadus glacialis (Latin: glacialis = ice; Russian: ледовая треска) and ‘polar cod’ for Boreogadus saida. Fish names lists representing the 

official stand of national and international organizations such as the American Fisheries Society and American Society of Ichthyologists and Herpetologists 

(Nelson et al. 2004) and the Fisheries Society of the British Isles (Wheeler 1992) differ and, thus, preclude establishment of universal common names. 

Fortunately, use of scientific names at first mention allows preferential use of vernacular names.” We acknowledge that our of use Ice Cod is different from other 

North American nomenclature (Page and others, 2013; Hoff and others, 2015; Mecklenburg and others, 2016). Mecklenburg and others (2013) list Ice Cod as 

one of the “English” names for A. glacialis.

Nomenclature and Previous 

Misidentifications

Genetic analysis, in concert with other species 

identification techniques, has greatly assisted ichthyological 

efforts to clarify and resolve existing taxonomic uncertainty 

surrounding many Arctic species. This work has resulted in 

several important changes in scientific and common names, 

which must be continually tracked for natural resource 

inventories to remain current. The changes in nomenclature 

in table 2.3 relate to the greater resolution in phylogenetic 

understanding that presently exists than was available at the 

time when the Fishes of Alaska (Mecklenburg and others, 

2002) was published.

Many species relationships require further study and 

additional changes in synonymy are expected. As an example, 

the relationship between Hippoglossiodes robutus (Bering 

Flounder) and H. elassodon (Flathead Sole) are currently 

being investigated (Kartavtsev and others, 2008; Mecklenburg 

and Steinke, 2015; Mecklenburg and others, 2016); 

preliminary results suggest the two species are the same.
36 Alaska Arctic Marine Fish Ecology Catalog

Zoogeographic Patterns

Definitions of zoogeographic pattern are from 

Mecklenburg and others (2011, Online Resource 1) and 

relate to species endemism and ranges of their population 

viability:

• Arctic—Species that live and reproduce in Arctic 

waters and are not found, or only infrequently found, 

in adjacent boreal waters.

• Predominantly Arctic—Species that are usually 

found in Arctic waters.

• Predominantly Boreal—Species that are 

characteristic of boreal waters and common in the 

border regions of the Arctic (for example, eastern 

Barents Sea, Gulf of Anadyr [Russia], Norton 

Sound [Alaska]). Can be more specific, such as 

predominantly Boreal Pacific or predominantly 

Boreal eastern Atlantic.

• Boreal—Species that are characteristic of boreal 

waters and only rarely or temporarily found in the 

border regions of the Arctic. Can be more specific, 

such as Boreal Pacific or Boreal eastern Atlantic. 

Includes subtropical, southern boreal category of 

Andriashev and Chernova (1994). 

• Arctic-Boreal—Species are distributed and spawn in 

both Arctic and Boreal waters.

• Widely Distributed—Species common in the 

temperate, boreal, and subtropical zones and in the 

warm waters of at least two oceans or known from 

the southern hemisphere, and occurring only rarely 

in the Arctic.

4

The decision to use three abundance categories used by Miller and Lea 

(1976) relates to the lack of consistent, long-term data sets needed for more 

precise estimates. 

5

Marine fishes are not uniformly distributed in time or space. This analysis 

was unable to capture seasonal and locality differences but rather portrays 

regional patterns in abundance.

6

“Acceptable,” as used here, refers to the precision of existing data with 

respect to the seasonal, geographic, and taxonomic qualities of data collection 

and how accurately they describe baseline conditions.

Abundance Terminology

Species distribution and abundance was determined from 

the reports of a large number of reconnaissance surveys and 

research expeditions. The wide variation in sampling times, 

areas, and collection methods reported raised issues of data 

comparability, which required reviewing existing abundance 

measures, defining appropriate relative abundance terms, and 

consistently applying operational definitions. Many methods 

and descriptors have been used to express fish abundance in 

Arctic studies. Relative abundance terms such as abundant, 

common, fairly common, present, occasional, uncommon, or 

rare are commonplace in the reported literature (for example, 

Coad and Reist, 2004). Existing baselines for Arctic marine 

fishes are not consistent for sampling in time, space, or 

methods to support a quantitative, multi-scaled classification 

that might be suggested by the definitions from Mecklenburg 

and others (2011) and Mecklenburg and Steinke, 2015). The 

challenge was to define a simple, but robust classification 

system that would have broad regional application. 

Four ecological principles guided the definition process: 

(1) abundance is a continuum between rare and abundant, 

(2) abundance is scale dependent, (3) sampling bias affects 

estimates of abundance, and (4) environmental influences on 

species abundance are taxon-specific. 

 “Rare,” “uncommon,” and “common” terms are defined 

in this report to classify relative abundance in the species 

accounts. These terms also were used by Miller and Lea 

(1976) to describe the coastal marine fishes of California4. 

For the Arctic ichthyofauna, abundance was classified using 

the same terms but with different operational definitions 

determined by species compositions and capture rates of 

marine fishes. These descriptive statistics are commonly 

used to characterize catch and, with respect to a regional 

classification scheme, meet pragmatic criteria related to 

acceptability (are ecologically sound and reliable), practicality 

(are applicable to existing records), and effectiveness (are 

realistic with broad-scale5 patterns). Species composition 

(percentage of occurrence of a species with respect to total 

number of individuals in the catch) provides a simple and 

scale dependent quantification of an assemblage on an 

acceptable6

 continuum. Capture rates are a lower resolution 

measure of abundance based on the presence or absence 

of a species detected per sampling session and frequency 

of capture (or rate) over sampling period and survey area. 
Chapter 2 37

Table 2.4. Relative abundance of Arctic marine fishes in the 

Chukchi and Beaufort Seas.

[Symbols: ≥, greater than or equal to; >, greater than; <, less than; ≤, less than or 

equal to]

Relative 

abundance term

(ecological science)

Species composition

(by geographic locale)

Frequency of capture

(encounter rates in 

catch records)

Common Regularly collected 

(comprise ≥10 to 

50 percent of catch in 

terms of total numbers 

of individuals)

Constant rate of capture 

(>10 to 25 percent of 

records)

Uncommon Infrequently observed or 

collected (<10 percent 

of 

catch in terms of total 

numbers of individuals)

Low rate of capture 

(≤10 percent of 

records)

Rare Seldom reported 20 or fewer records 

in Arctic (or 1 to 2 

captures per cruise)

Detection rates allow a higher order, less precise comparison 

of abundance across gear-types and habitats and are a more 

subjective index. With respect to gear bias and comparability 

issues, the resultant abundance categories represent semiquantitative composite metrics from descriptive analyses of 

catch and expert opinion about relative abundance across at 

the large-scales of distributional patterns (table 2.4).

Endemism and Relative Abundance 

Mecklenburg and others, (2011) describe the 

zoogeographic pattern of 242 marine fish species from the 

Arctic zoogeographic province. This province includes the 

northern Bering Sea and North Atlantic Ocean at latitudes 

far south of this study area. These authors reported 41 

percent of the zoogeographic patterns of the marine fauna 

were Arctic (that is, Arctic, Predominantly Arctic, and 

Arctic-Boreal zoogeographic patterns) and 59 percent were 

Boreal (that is, Predominantly Boreal, Boreal, and Widely 

Distributed zoogeographic patterns). Applying this approach 

to just the United States area of the Arctic Province, we 

determined that the Arctic pattern in 58 percent (n = 92) of 

the marine fishes was from the Chukchi Sea and 73 percent 

(n = 80) was from the Beaufort Sea. (Editors note: Analysis 

does not include newly confirmed species to the region—

Somniosus pacificus, Bathyraja parmifera, Icelus spiniger, 

and Lycodes pallidus.) The Boreal pattern comprised 42 and 

26 percent of the marine faunas, respectively. The ratio of 

Arctic to Boreal patterns for marine fish found in the U.S. 

Chukchi and Beaufort Seas is about 60:40 (table 2.5). 
38 Alaska Arctic Marine Fish Ecology Catalog

Scientific name Common name

Confirmed occurrence

Zoogeographic pattern Chukchi Sea Beaufort Sea

Entosphenus tridentatus Pacific Lamprey Rare Boreal Pacific

Lethenteron camtschaticum Arctic Lamprey Common Common Arctic-Boreal

Squalus suckleyi Spotted Spiny Dogfish Rare Boreal Pacific

Somniosus pacificus Pacific Sleeper Shark Rare Boreal Pacific

Amblyraja hyperborea Arctic Skate Rare Arctic

Bathyraja parmifera Alaska Skate Rare Boreal Pacific

Clupea pallasii Pacific Herring Common Uncommon Arctic-Boreal Pacific

Hypomesus olidus Pond Smelt Rare Predominantly Boreal Pacific

Mallotus catervarius Pacific Capelin Common Common Arctic-Boreal

Osmerus dentex Arctic Smelt Common Common Arctic Boreal Pacific

Coregonus autumnalis Arctic Cisco Rare Common Arctic

Coregonus laurettae Bering Cisco Common Common Predominantly Arctic

Coregonus nasus Broad Whitefish Common Common Predominantly Arctic

Coregonus pidschian Humpback Whitefish Common Common Predominantly Arctic

Coregonus sardinella Least Cisco Common Common Predominantly Arctic

Oncorhynchus gorbuscha Pink Salmon Common Uncommon Predominantly Boreal Pacific

Oncorhynchus keta Chum Salmon Common Uncommon Predominantly Boreal Pacific

Oncorhynchus kisutch Coho Salmon Uncommon Uncommon Predominantly Boreal Pacific

Oncorhynchus nerka Sockeye Salmon Uncommon Uncommon Predominantly Boreal Pacific

Oncorhynchus tshawytscha Chinook Salmon Uncommon Uncommon Predominantly Boreal Pacific

Salvelinus malma Dolly Varden Common Common Arctic-Boreal Pacific

Stenodus leucichthys Inconnu Common Arctic-Boreal1

Benthosema glaciale Glacier Lanternfish Rare Arctic-Boreal Atlantic

Arctogadus glacialis Ice Cod Rare Arctic

Boreogadus saida Arctic Cod Common Common Arctic

Eleginus gracilis Saffron Cod Common Common Arctic-Boreal Pacific

Gadus chalcogrammus Walleye Pollock Uncommon Uncommon Predominantly Boreal Pacific and Atlantic

Gadus macrocephalus Pacific Cod Uncommon Uncommon Arctic-Boreal Pacific and western Atlantic

Gasterosteus aculeatus Threespine Stickleback Common Rare Predominantly Boreal

Pungitius pungitius Ninespine Stickleback Common Common Predominantly Boreal

Hexagrammos stelleri Whitespotted Greenling Uncommon Rare Predominantly Boreal Pacific

Artediellus ochotensis Okhotsk Hookear Sculpin Rare Boreal Pacific

Artediellus scaber Hamecon Common Common Arctic

Enophrys diceraus Antlered Sculpin Common Uncommon Predominantly Boreal Pacific

Gymnocanthus tricuspis Arctic Staghorn Sculpin Common Common Arctic

Hemilepidotus papilio Butterfly Sculpin Common Predominantly Boreal Pacific

Icelus bicornis Twohorn Sculpin Rare Uncommon Predominantly Arctic

Icelus spatula Spatulate Sculpin Common Common Arctic-Boreal

Icelus spiniger Thorny Sculpin Rare Predominantly Boreal

Megalocottus platycephalus Belligerent Sculpin Common Rare Predominantly Boreal Pacific

Microcottus sellaris Brightbelly Sculpin Rare Predominantly Boreal Pacific

Myoxocephalus jaok Plain Sculpin Rare Rare Predominantly Boreal Pacific

Myoxocephalus 

polyacanthocephalus

Great Sculpin Rare Boreal Pacific

Myoxocephalus quadricornis Fourhorn Sculpin Common Common Predominantly Arctic

Myoxocephalus scorpioides Arctic Sculpin Uncommon Uncommon Arctic

Myoxocephalus scorpius Shorthorn Sculpin Common Common Arctic-Boreal

Trichocottus brashnikovi Hairhead Sculpin Common Uncommon Predominantly Boreal Pacific

Triglops nybelini Bigeye Sculpin Uncommon Uncommon Arctic

Triglops pingelii Ribbed Sculpin Common Uncommon Arctic-Boreal

Blepsias bilobus Crested Sculpin Uncommon Boreal Pacific

Nautichthys pribilovius Eyeshade Sculpin Common Rare Predominantly Boreal Pacific

Cottunculus microps Polar Sculpin Rare Arctic-Boreal Atlantic

Eurymen gyrinus Smoothcheek Sculpin Rare Predominantly Boreal Pacific

Table 2.5. Relative abundance and zoogeography of Arctic marine fishes in the Chukchi and Beaufort Seas.

[Zoogeographic pattern: Main references, Mecklenburg and others (2011) Online Resource 1, Mecklenburg and Steinke (2015), and Mecklenburg and others 

(2016). Blank cell indicates the species is unreported or unknown from the area]
Chapter 2 39

Scientific name Common name

Confirmed occurrence

Zoogeographic pattern Chukchi Sea Beaufort Sea

Aspidophoroides monopterygius Alligatorfish Common Common Predominantly Arctic

Aspidophoroides olrikii Arctic Alligatorfish Common Common Predominantly Arctic

Hypsagonus quadricornis Fourhorn Poacher Rare Boreal Pacific

Leptagonus decagonus Atlantic Poacher Uncommon Uncommon Arctic-Boreal

Occella dodecaedron Bering Poacher Rare Boreal Pacific

Pallasina barbata Tubenose Poacher Rare Predominantly Boreal Pacific

Podothecus veternus Veteran Poacher Common Common Arctic-Boreal Pacific

Eumicrotremus andriashevi Pimpled Lumpsucker Rare Arctic-Boreal Pacific

Eumicrotremus derjugini Leatherfin Lumpsucker Rare Uncommon Arctic

Careproctus reinhardti Sea Tadpole Rare Rare Arctic

Liparis bathyarcticus Nebulous Snailfish Common Common Predominantly Arctic

Liparis fabricii Gelatinous Seasnail Common Common Arctic

Liparis gibbus Variegated Snailfish Common Uncommon Arctic-Boreal

Liparis tunicatus Kelp Snailfish Common Common Arctic

Paraliparis bathybius Black Seasnail Rare Rare Arctic

Gymnelus hemifasciatus Halfbarred Pout Common Common Arctic-Boreal

Gymnelus viridis Fish Doctor Common Common Predominantly Arctic

Lycenchelys kolthoffi Checkered Wolf Eel Rare Arctic

Lycodes adolfi Adolf’s Eelpout Rare Arctic

Lycodes eudipleurostictus Doubleline Eelpout Uncommon Arctic

Lycodes frigidus Glacial Eelpout Rare Rare Arctic

Lycodes jugoricus Shulupaoluk Rare Arctic

Lycodes marisalbi White Sea Eelpout Rare Arctic

Lycodes mucosus Saddled Eelpout Common Uncommon Arctic

Lycodes palearis Wattled Eelpout Common Uncommon Predominantly Boreal Pacific

Lycodes pallidus Pale Eelpout Rare Arctic

Lycodes polaris Polar Eelpout Common Common Arctic

Lycodes raridens Marbled Eelpout Common Common Arctic-Boreal Pacific

Lycodes reticulatus Arctic Eelpout Common Arctic

Lycodes rossi Threespot Eelpout Rare Arctic

Lycodes sagittarius Archer Eelpout Common Arctic

Lycodes seminudus Longear Eelpout Rare Rare Arctic

Lycodes squamiventer Scalebelly Eelpout Rare Arctic

Lycodes turneri Estuarine Eelpout Uncommon Uncommon Predominantly Arctic Pacific

Acantholumpenus mackayi Blackline Prickleback Uncommon Predominantly Boreal Pacific

Anisarchus medius Stout Eelblenny Common Uncommon Arctic-Boreal

Chirolophis snyderi Bearded Warbonnet Uncommon Predominantly Boreal Pacific

Eumesogrammus praecisus Fourline Snakeblenny Uncommon Uncommon Arctic-Boreal Pacific and western Atlantic

Leptoclinus maculatus Daubed Shanny Common Common Arctic-Boreal Pacific and Atlantic

Lumpenus fabricii Slender Eelblenny Common Common Arctic-Boreal 

Lumpenus sagitta Snake Prickleback Rare Boreal Pacific

Stichaeus punctatus Arctic Shanny Common Common Arctic-Boreal Pacific and western Atlantic 

Pholis fasciata Banded Gunnel Uncommon Rare Arctic-Boreal

Anarhichas denticulatus Northern Wolffish Rare Rare Arctic-Boreal

Anarhichas orientalis Bering Wolffish Uncommon Uncommon Predominantly Boreal Pacific

Zaprora silenus Prowfish Rare Boreal Pacific

Ammodytes hexapterus Arctic Sand Lance Common Common Arctic-Boreal Pacific

Hippoglossoides robustus Bering Flounder Common Common Arctic-Boreal Pacific

Hippoglossus stenolepis Pacific Halibut Rare Boreal Pacific

Limanda aspera Yellowfin Sole Common Rare Predominantly Boreal Pacific

Limanda proboscidea Longhead Dab Uncommon Rare Predominantly Boreal Pacific

Limanda sakhalinensis Sakhalin Sole Uncommon Predominantly Boreal Pacific

Liopsetta glacialis Arctic Flounder Common Common Predominantly Arctic

Platichthys stellatus Starry Flounder Uncommon Uncommon Arctic-Boreal Pacific

Pleuronectes quadrituberculatus Alaska Plaice Uncommon Predominantly Boreal Pacific

Reinhardtius hippoglossoides Greenland Halibut Uncommon Uncommon Arctic-Boreal Pacific and Atlantic

1

Author determination.

Table 2.5. Relative abundance and zoogeography of Arctic marine fishes in the Chukchi and Beaufort Seas.—Continued
40 Alaska Arctic Marine Fish Ecology Catalog

Acknowledgments

The personal observations and ichthyologic data and 

insights provided by Catherine W. Mecklenburg (California 

Academy of Sciences, San Francisco and Point Stephens 

Research, Auke Bay, Alaska) were critical during early 

planning and review phases of this study. Her participation in 

RUSALCA, BOEM, Conservation of Arctic Flora and Fauna, 

U.S. Fish and Wildlife Service, National Park Service, NOAA 

Auke Bay Laboratories, and other Arctic research provided 

biogeographic expertise, taxonomic and diversity insights, and 

critical scientific reviews of developing project information. 

Her scientific contributions to and review of chapter 2 were 

especially significant with respect to development of a 

provisional checklist of valid marine fishes from the Chukchi 

and Beaufort Seas as well the identification of probable and 

questionable species occurrences. Special appreciation is 

extended to Jørgen Schou Christiansen (University of Tromsø, 

Norway) and Robert Lea (formerly of State of California) 

for their thoughtful insights on the consistent nomenclature 

(vernacular names) for Boreogadus saida and Arctogadus 

glacialis. 

Summary

One hundred nine (109) marine fishes from 24 families 

are described and compared regarding their occurrence in 

the United States Chukchi and Beaufort seas. Ninety-seven 

(97) species are confirmed from the Chukchi Sea and 83 

from the Beaufort Sea. Twenty species are newly confirmed 

to the U.S. Arctic since the publication of Fishes of Alaska

in 2002. The taxonomy of Arctic marine fishes has improved 

since 2002, but issues in the naming and acceptance and 

validity of several species in the Liparidae and Zoarcidae 

require additional resolution. Many of the common species 

are benthic or demersal in their habitat orientation. Smallsized marine species such as those representing Cottidae, 

Liparidae, Stichaeidae, and Zoarcidae families, were common 

to both seas. The diversity of Salmonidae species reflects 

the plasticity of this group of fishes and its adaptive linkage 

to freshwaters. Our analysis supports high rates of Arctic 

endemism in both seas, especially in the Beaufort Sea. 

Numerous range extensions and their sources are noted. The 

estimates of species diversity can be expected to increase 

with new sampling, with greater reliance on genetic and 

molecular identification aids, and in response to large-scale 

effects of changing environmental conditions on present 

patterns of regional fish distribution and abundance. Fishes of 

Alaska remains an important reference for species taxonomy 

and identification of more than 600 species from the Gulf of 

Alaska and Bering Sea to the Arctic but is not complete with 

respect to species in the Chukchi and Beaufort Seas.
Abstract

Species accounts provide brief, but thorough descriptions 

about what is known, and not known, about the natural life 

histories and functional roles of marine fishes in the Arctic 

marine ecosystem. Information about human influences on 

traditional names and resource use and availability is limited, 

but what information is available provides important insights 

about marine ecosystem status and condition, seasonal patterns 

of fish habitat use, and community resilience. This linkage has 

received limited scientific attention and information is best 

for marine species occupying inshore and freshwater habitats. 

Some species, especially the salmonids and coregonids, are 

important in subsistence fisheries and have traditional values 

related to sustenance, kinship, and barter. Each account is an 

autonomous document providing concise information about a 

species zoogeography, western and Alaska Native taxonomy, 

life history, niches, and life requirements. Each account is 

fully referenced with the identification of the most critical 

literature for Alaska and a more comprehensive listing of 

referencing from which biological and ecological information 

was drawn. New-to-science narratives, distributional maps, 

and vertical profiles, provide quick, reliable sources of 

information about fish life history and habitat requirements for 

this segment of the Arctic fauna.

Purpose and Design of Species 

Accounts

Individual species accounts were prepared for 104 of the 

109 confirmed marine fishes for which adequate biological 

information was available from the U.S. Chukchi and 

Beaufort Seas. These descriptions are an important source 

of documentation about Arctic Alaska’s marine fish fauna. 

Although tailored to address the specific needs of BOEM 

Alaska OCS Region NEPA analysts, the information presented 

in each species account also is meant to be useful to other 

users including state and Federal fisheries managers and 

scientists, commercial and subsistence resource communities, 

and Arctic residents. Readers interested in obtaining additional 

information about the taxonomy and identification of marine 

Arctic fishes are encouraged to consult the Fishes of Alaska

(Mecklenburg and others, 2002) and Pacific Arctic Marine 

Fishes (Mecklenburg and others, 2016). By design, the species 

accounts enhance and complement information presented in 

the Fishes of Alaska with more detailed attention to biological 

and ecological aspects of each species’ natural history 

and, as necessary, updated information on taxonomy and 

geographic distribution. 

Each species account includes a concise summary of 

the natural history, population dynamics, functional roles, 

and traditional and economic values of the marine fish found 

off Alaska. An initial organizational task was to create a 

standard format for effective information delivery. The species 

descriptions by Ehrlich and others (1988) were provided to 

the USGS by BOEM as an example of a creative template for 

information transfer. Four pilot species accounts, representing 

well known to poorly known species, were developed, 

reviewed, and repeatedly revised for improvements, 

interagency approval, and selection of the final layout and 

design. Final decisions about content represented the priority 

needs of BOEM. 

More than 1,200 individual scientific publications 

relevant to Arctic marine fishes were reviewed in preparation 

of the species accounts. In each species account, the most 

relevant literature for each species is cited. A shorter list 

(about 5–10 articles) identifies key Alaskan information 

sources that, in our opinion, have had the greatest scientific 

effect on understanding the species of the Arctic area of the 

United States. 

Chapter 3. Alaska Arctic Marine Fish Species Accounts

By Milton S. Love1, Nancy Elder2, Catherine W. Mecklenburg3, Lyman K. Thorsteinson2, and 

T. Anthony Mecklenburg4

1

University of California, Santa Barbara.

2

U.S. Geological Survey.

3

California Academy of Sciences, San Francisco, and Point Stephens 

Research, Auke Bay, Alaska.

4

Point Stephens Research, Auke Bay, Alaska.
42 Alaska Arctic Marine Fish Ecology Catalog

Limitations of Data

The species accounts reveal many gaps in the biological 

information needed to conduct vulnerability assessments 

of the marine fishes of the Beaufort and Chukchi Seas to 

human interventions. Part of this problem relates to the 

geographic coverage of existing research and surveys in 

Alaska as, in many instances, we were required to incorporate 

the results of investigations conducted outside the region. 

This raises an important caution because, even though the 

best available information was used in preparing the species 

accounts, our reliance on data and information from outside 

Alaska will introduce uncertainty to EIS expectations. 

Ideally, and with respect to oil and gas activities, baseline 

information for fishery resources should be collected from 

the potentially affected environment to appropriately evaluate 

the potential effects of oil spills or other possible industrialrelated disturbances. However, as has been widely noted 

(for example, Bluhm and others, 2011), systematic and 

methodologically comparable data typically are not available 

from Arctic Alaska marine ecosystems. Evaluating change in 

populations and communities from natural and anthropogenic 

stressors is limited by the variable quality and lack of 

quantitative reports on abundance, distribution, community 

structure, and demographics for Arctic marine fishes. 

In each species account, an attempt was made to 

incorporate the most reliable baseline information available 

and offer impressions of information needs. Important ongoing 

studies sponsored by BOEM, and others, may be addressing 

some of these needs. The needs assessments for this study 

considered these efforts to the extent that oral and (or) written 

communications and preliminary results allowed. The focus 

of this study was on impressions of the population parameters 

(Williams and others, 2002) and environmental measurements 

needed to detect changes in marine fish populations (Reist 

and others, 2006; Wassmann and others, 2011) and their 

resilience to a variable and rapidly changing environment 

(Holland-Bartels and Pierce, 2011). For key marine fish 

species, examples might include changes in range, community 

structure, abundance, phenology, behavior, and population 

growth and survival.

Each species account is designed as a self-contained 

article; therefore, no references to other accounts are included. 

Additionally, to reduce complexity in the presentations, only 

common names were used to identify the major predator 

and prey species for the marine fish described. Because this 

document was meant to be a companion document to the 

Fishes of Alaska (Mecklenburg and others, 2002), interested 

readers are encouraged to consult this book or Page and others 

(2013) and Mecklenburg and others (2016) for more complete 

information about the scientific authorities and literature 

citations associated with the original descriptions of each 

species. Readers are directed to the references cited in each 

species account for additional information on the species. 

Operational Definitions

In chapter 1, several concepts about the temporal and 

spatial habitat requirements for Arctic marine fish were 

introduced. More information is presented in this chapter to 

explain the vertical distribution and the location of shelf break, 

as used in this report.

Vertical Distribution

The conceptual design of the species depth profiles 

(vertical structure by life history stage) was patterned after 

the “coastal marine life zones” of Allen and Smith (1988). 

The goal of the profiles is to visualize what is known about 

a species occurrence and reproductive ecology by depth and 

location. An idealized characterization of Arctic shelves was 

designed to visualize these relationships. Additional detail 

about origins of data was included in the depth profiles to 

reflect Alaskan records or collections from other Arctic 

regions. This is important because actual field collections and 

observations are limited from this region. In many instances, 

the actual presence of a life stage remains unverified by field 

sampling. Thus, for many of species, the depth of a fish’s life 

cycle should be considered untested hypotheses in need of 

additional testing. 

Location of Shelf Break

Early versions of the depth profiles were modified at 

the request of BOEM with respect to the depiction of the 

continental shelf break. As a special effect for the Arctic, 

the species depth profiles were redrawn to depict the change 

in bathymetry that typically occurs at depths of about 75 m 

throughout the Chukchi and western Beaufort Seas. This 

depiction is not an attempt to redefine the oceanographic 

definition of shelf break. Instead, it highlights the relatively 

sharp gradient in depths that often occurs near 70- to 80-m 

contours over much of the region. Although species depth 

profiles in this report depict an apparent “break” at 75-m, three 

factors were considered: (1) this is a generalization and the 

actual shelf break may be geographically close but at a slightly 

greater depth; (2) shelf edge effects on fish distribution at 

depths occurring between 75-, 150-, or 200-m are likely 

negligible due to the gradient and area involved; and (3) the 

conceptual depictions of depth distributions by life history 

stage are consistent with accepted oceanographic conventions 

for continental shelf and slope (despite the magnified view at 

75-m) and thus are compatible to the import of biological data 

obtained elsewhere.
Chapter 3 43

Keystone Species

The concept of keystone species describes the critical 

role certain organisms are perceived to have in maintaining 

the structure of biological communities and resilience of 

ecosystem dynamics (Paine, 1966). Arctic Cod (Boreogadus 

saida) are widely distributed in the Arctic Ocean and by virtue 

of their abundance and intermediate trophic position between 

invertebrates and higher-level predators are integral to the 

movement of nutrients in marine food webs. For this reason, 

Arctic Cod are considered a keystone species in the Arctic 

marine (Bradstreet and others, 1986; Walkusz and others, 

2011). Arctic Cod are common in United States waters of 

the Beaufort and Chukchi Seas being considered for energy 

exploration and development and are an ecological focus of 

BOEM fishery studies to understand potential effects on the 

species (Maule and Thorsteinson, 2012).

Outline of Species Accounts

The species accounts are scientifically accurate 

descriptions of the life histories, populations, habitats, and 

community values of individual species in the Arctic marine 

ecosystem. The mix of quantitative and qualitative information 

presented reflects state-of-the-art knowledge, a faunal 

assessment of information gaps, and prioritization of priority 

needs for population and process understanding. Limited 

information for many Alaskan species required that relevant 

observations from other geographic locales be included. Each 

species account attempts to be clear about the geographic 

origins of data and information, through scientific referencing 

or special notations in graphics. As an example, italics are 

used in the species accounts to highlight data collections from 

the Alaska study area. In several instances, species information 

was so lacking that inferences from a closely related species 

were required. 

The generic species account includes a comprehensive 

accounting of scientific and cultural information in a standard 

format. The scientific information addresses multiple 

disciplinary areas including taxonomy, life history and 

habitats, ecological relationships including predator-prey 

interactions and environmental preferences, and population 

ecology. The population information is critical to evaluations 

of population status and health, resilience, and vulnerability to 

natural and anthropogenic changes in the marine environment. 

Each species account includes a photograph of an adult 

specimen (or line drawing if an image was not available); 

distribution maps (horizontal and vertical); and concise 

descriptions of abundance, life history, and ecology (11 life 

history categories); major stressors; research needs; and 

key references. To assist users, a suite of easily recognized 

icons was developed to provide quick access to specific life 

history information. In addition, some species attributes 

regarding life history, population dynamics, and biological 

interactions are defined in the Glossary (chapter 7).

Information presented in each species account is outlined 

and described as:

Taxonomic—Scientific and Common Names 

The format of the species accounts was, by design, 

intended to link the biologic and ecologic information 

presented in this document directly to the species identification 

guides contained in the “Fishes of Alaska.” This connection 

was established by adherence to naming conventions as 

described by Mecklenburg and others, 2002 (p. 25 and 26). 

The common names of each marine fish are presented first, 

followed by scientific and family names. Each scientific name 

includes a reference to the name of the person (author) who 

formally described and named the species in the ichthyological 

literature. The bibliographic data for the authors and dates of 

publication of scientific names can be found in Eschmeyer’s 

Catalog of Fishes online (http://researcharchive.calacademy.

org/research/ichthyology/catalog/fishcatmain.asp) and are 

not reported here. In some instances, a Note (italicized) has 

been included to describe exceptional details about existing 

biological data, morphology, nomenclature, taxonomic status, 

life history strategy, or occurrence of a species in the United 

States Chukchi and Beaufort Seas.

Iñupiat Name

The existence of colloquial Iñupiat (Iñupiaq) names for 

the Arctic’s marine fish fauna by indigenous peoples is an 

important component of traditional ecological knowledge. 

Relatively few marine fish species are abundant or 

susceptible enough to subsistence fisheries to have received 

special names. For those species having Iñupiat names, this 

information is reported to assure that a common vocabulary 

can facilitate future exchanges of ideas and knowledge across 

disciplinary boundaries. In this manner, colloquial names 

can provide a cultural link between local marine resources 

and science supporting sustainability of Arctic communities

and ecosystems.

Ecological Role

Fishes play a pivotal role in marine ecosystems as 

secondary and higher-level consumers in many marine food 

webs. In many instances, information about predator-prey 

relationships is so limited that only preliminary, qualitative 

assessments of the relative role of each species are possible. 

The ecological niche describes how an organism or population 

responds to resources and competitors. Importance or 

significance descriptors do not diminish the fact that all 

organisms contribute in ways large or small to the provision 
44 Alaska Arctic Marine Fish Ecology Catalog

of ecosystem goods and services. These descriptors however, 

may provide useful information about the relative importance 

of a particular species as an indicator of ecosystem condition 

and trajectories of change associated with climate change, 

habitat fragmentation, ecosystem stress, effect of pollutants, or 

other anthropogenic effects.

Physical Description/Attributes

A brief physical description of the species is summarized 

from information presented by Mecklenburg and others, 

(2002) in the Fishes of Alaska; the relevant page number 

is included for quick referral to more comprehensive 

morphological information. An image of the adult form of 

each fish is presented with appropriate attribution. Highquality images were selected to highlight the key identifying 

features of a particular species. 

Information about the presence of a swim bladder and 

antifreeze glycoproteins is included because of its relevance 

to geo-seismic oil and gas exploration, climate change issues, 

and evolutionary life history. 

Range

The geographic occupancy of the species in United States 

sectors of Chukchi and Beaufort Seas and adjacent waters is 

presented in brief narratives and depicted on maps. Known 

occurrence in the Arctic OCS Planning Areas is highlighted by 

symbols indicating locations of valid species identifications 

from properly archived voucher specimens on each map. 

Although the symbols on the maps may suggest that some 

of the species are rare in the region, the study of historical 

collections from the United States and Canadian sectors of 

the Beaufort Sea, as well as the collections from BOEM 

surveys in the Beaufort in 2011 and 2012, is still in progress 

and may reveal that these species are more abundant in deep 

sectors of the study area than the maps suggest. Definitions 

of zoogeographic pattern are from the Online Resource 1 

(electronic supplemental to Mecklenburg and others, 2011), 

Mecklenburg and Steinke (2015), and Mecklenburg and others 

(2016) and relate to ranges of population viability (see chapter 

2).

Depth profiles in each species account graphically 

summarize existing information about the benthic and 

reproductive distributions of each marine fish. In both 

depth profiles, the width of areas depicted confers species 

information about horizontal (onshore-offshore) patterns 

of distribution. The italicized captions in the depth profiles 

highlight species information germane to the study area. 

Areas in the graphs denoted by the orange coloration represent 

understanding from data collection within the United States 

Chukchi and Beaufort Seas; olive colors represent data 

collection outside the study area. For benthic distributions, 

solid lines in the depth profiles represent species for which 

no specific information is available about its preferred depth 

range. Solid lines represent a synthesis of understanding that 

includes information not necessarily specific to the study area. 

In some instances, only one record of a species occurrence by 

depth was available and coding in orange was not meaningful. 

In these cases, an explanatory comment, in italicized font, with 

a line pointing to the appropriate depth was included in the 

graph (for example, see the species account for Megalocottus 

platycephalus). Highlighted depths as indicated through 

“bolded” (dark black) and dashed segments, represent most 

common depths where the species has been detected, and 

depth distribution as has been reported throughout the species 

range, respectively. Areas denoted with diagonal crosshatching represents depth distribution of juveniles (immature); 

adult distributions are not cross-hatched and age-related 

habitat overlaps, are informed by captioning in the figures.

For reproductive distribution, eggs and larvae 

(pre‐juvenile life stages) of marine fishes are represented 

with respect to depth and distance from the coast. Orange 

areas in the reproductive distribution profiles represent data 

collection in the study area. In many instances, information 

about spawning habitats and egg and larval distributions is 

summarized from information reported from throughout a 

species range. In these cases, dark blue represents species 

distributions in spawning habitats; light blue represents 

the geographic distributions of eggs and larvae; and light 

green is used to highlight areas of substantial habitat overlap 

(for example, see the species account for Hippoglossus 

stenolepsis). Distribution patterns of eggs and larvae are 

symbolized by “dots” and “horizontal dashes,” respectively, 

in the graphs. As for benthic distribution, solid lines represent 

species-specific information from data collections from 

throughout the species entire range. Highlighted (dark black 

lines) segments of solid lines indicate the most common 

depths where egg and larvae samples have been collected. 

Dashed lines represent areas of hypothesized distributions 

for species for which no information is available about egg 

or larval occurrence. In these instances the hypothesized 

distributions are based on known patterns for closely related 

species; the lack of data is stated in captions above the graph.

Relative Abundance

Relative abundance refers to the contribution a species 

makes to the total abundance of the fishery community. It is a 

measure that provides an index of the number of individuals 

present, but not the actual numbers. Relative abundance terms, 

such as “common,” “uncommon,” or “rare” often are used 

to express the general population status of a given species, 

but are most useful when they are defined by something 

that is measured or estimated in a manner that makes 

comparison meaningful.
Chapter 3 45

Depth Range

Benthic distribution refers to the spatial arrangement 

of a particular species at different depths over continental 

shelf and slope waters. The life cycle of fishes occurs in 

multiple dimensions in time and space and generally reflects 

genetically determined life history or behavior that has 

evolved to maximize fitness (life time reproductive success, 

see Gross [1987]). Benthic distribution profiles for each 

species represent the location of important habitats as they are 

presently known for juvenile and marine fishes. Reproductive 

distributions depict important habitats for spawning and early 

life history development.

Life History, Population Dynamics, and 

Biological Interactions

Life history theory holds that the schedule and duration 

of key events in a species’ lifetime are shaped by natural 

selection to produce the largest possible number of surviving 

offspring. These events, notably juvenile development, age 

of sexual maturity, first reproduction, number of offspring 

and level of parental investment, senescence, and death, 

depend on the abiotic and biotic environment of the organism. 

Specific information about these traits informs understanding 

of a species’ adaptive capacity including major influences 

on population abundance. A number of fisheries models use 

basic length-weight and age-at-size relationships to describe 

the growth and dynamics of fishery populations (for example, 

von Bertalanffy and Gompertz, growth models and derivatives 

[Ricker, 1975]). Ecological models estimate transfer of 

energy or matter along the trophic chain (Gamito, 1998). The 

parameters that are estimated in these models are individually 

important indicators of population condition and may be 

used with other indicators to derive quantitative information 

about compensatory responses and resilience. Much of this 

information, including population parameters, has been 

compiled in FishBase for the Arctic marine fish (Froese and 

Pauly, 2012).

Habitats and Life History—Basic 

information about the life history (for example, 

body size, reproductive ecology, growth) and 

ecology (for example, mobility, growth, 

habitat) of a species and the environmental area 

inhabited by that species is foundational to 

effective resource management. Habitat is the natural 

environment that influences and is used by a species 

population. Information about abiotic (that is, temperature, 

salinity, other physiochemical factors, depth, and substrate 

types) and biotic (that is, type and abundance of food, 

presence of other biota) often are used to describe fish habitats 

and provide insights about a species environmental 

preferences and habitat associations (for example, water 

masses). Maximum body size often is reported and can be an 

important surrogate of different life history traits (for example, 

age at maturity, growth, and reproductive output). In population 

dynamics studies, the relationships between length and weight 

and size and age form the basis for population growth and 

production models and quantitative analysis of environmental 

effects. Length measurements are reported as standard length 

(SL), total length (TL), and fork length (FL) in fisheries studies. 

Behavior (see also Glossary [chapter 7]).—

Behavior is the manner in which a fish operates 

or functions within its environment (that is, 

home range, territoriality, and many others) to 

procure food, orient to specific locations, or 

relate to other organisms. Knowing how 

individuals respond to the environment (physical, chemical, and 

biological cues) is critical to understanding population 

processes such as distribution, survival, and reproduction and 

recruitment and for managing fisheries. Many behaviors are 

evolutionary adaptations to the physiological and reproductive 

requirements for a species’ survival. For example, migration 

involves the regular movement of animals between different 

geographic locations. Migrations can be extensive in terms of 

time and distance involved (anadromous model) or seasonal 

(amphidromous and marine models). Each of these models 

reflects a life strategy adapted for age and growth at sea. Diel 

relates to daily changes in water column position due to changes 

in light, temperature, and food supply. 

Migratory behaviors are rooted in physiological 

requirements for food, growth, reproductive, and survival 

(“scope for growth”). Movement behaviors are more tactical 

responses to local environmental conditions (for example, 

variable hydrographic conditions in the nearshore Beaufort 

Sea). Fish movement can be active or passive and involve large 

distances in search of suitable habitats and foods. The seasonal 

nature of migration and movement behaviors are typically 

related to life history stage, predator-prey distributions, or 

energetic requirements for growth.

Schooling (that is, social structure of fish of the same 

species moving in more or less harmonious patterns in the sea) 

often is related to survival and reproduction. Schooling confers 

physical benefits to fish movement, safety against predators, 

search behaviors (for example, foods), population immunology, 

and reproduction. 

The functional feeding morphology of a fish relates to its 

anatomical adaptations (for example, body size, gape sizes, 

shape, and body form) to environmental conditions especially 

food preferences. The adage “function determines morphology 

and morphology determines way of life” is an important 

evolutionary concept as it applies to fish feeding behavior, 

dietary preferences, habitat selection, and trophic stature. 

Trophic position (within categories of trophic levels) expresses 

the “tendency of larger (less abundant) fishes feeding on smaller 

(more abundant) fishes, which themselves feed on zooplankton 

and all these animals resting upon primary producers” (from 

Pauly and Watson, 2005). Categories of trophic levels are:
46 Alaska Arctic Marine Fish Ecology Catalog

• Trophic level 1 (T1), plants and animals make their 

own food and are called primary producers; 

• Trophic level 2 (T2), herbivores eat plants and are 

called primary consumers;

• Trophic level 3 (T3), carnivores eat herbivores and are 

called secondary consumers; 

• Trophic level 4 (T4), carnivores eat other carnivores 

and are called tertiary consumers; and 

• Trophic level 5 (T5), apex consumers, which have no 

predators, are at the top of the food chain.

Populations or Stocks—A population often is 

defined as a group of organisms of the same 

species occupying a particular space at a 

particular time with the potential to breed with 

each other (Williams and others, 2002). Stocks 

are subpopulations of a particular species of 

fish that result from reproductive isolation and subdivisions 

within the biological range. The current state of knowledge 

about local stocks and their genetic population structure is 

reported. Grossberg and Cunningham (2001) described the 

combined effects of demographic, behavioral, genetic, 

oceanographic, climate, and tectonic processes as major 

determinants of population structure. These mechanisms act 

across a range of temporal and spatial scales to determine the 

rates and patterns of dispersal of different life stages of marine 

fishes. Dispersal, combined with the successful reproduction 

and survival of immigrants, control the scale and rate of 

processes that build or erode structure within and among 

groups of individuals.

Reproduction Mode—Little information is 

available about the spawning times and 

locations, mating behaviors (breeders or 

nonbreeders), and genetic diversity of Arctic 

marine fishes. What is known is drawn largely 

from observations from populations studied 

outside the United States. For most Arctic marine fish species, 

there is no information about population or stock structure (for 

example, age structure, reproductive behavior, sex ratios, 

age-at-maturity, fecundity, and genetic). These are key 

population parameters needed for understanding reproductive 

ecology, population dynamics (for example, growth, survival, 

and mortality), and assessments of resiliency (response 

to disturbance).

Food and Feeding—Dietary information is 

summarized from literature and, unless in 

italics, is reported from other regions. Fish 

communities can affect the ecological 

characteristics of marine ecosystems in 

response to productivity and abundance patterns, the mobility 

and migratory behavior of species, and through food 

influences in different habitats (for example, Grebmeier and 

others, 2006b). Trophic Index (T) values are reported from 

FishBase (Froese and Pauly, 2012). The T values for Arctic 

marine fishes are largely derived from stomach contents 

analyses, which have correlated well with stable isotopes of 

nitrogen in tissues. The fractional values (between 1 and 5) 

realistically address complexities of consumer feeding 

behaviors (omnivory and feeding across multiple trophic 

levels) and predator-prey relationships. For example, the mean 

T value for Blackline Prickleback (Acantholumpenus mackayi) 

is 3.1 (±0.31). This mid food web value is indicative of a 

primary carnivore that feeds across trophic levels, in this case 

on lower level herbivores.

Biological Interactions.—The effects 

organisms in a community have on one 

another. Competition and consumption 

(predation, herbivory, or cannibalism) are the 

best known of the major ecological processes 

affecting resource abundance, community 

composition, and ecosystem function. Competition involves 

interactions between individuals of the same species 

(intraspecific) or different species (interspecific) in which the 

fitness of one is lowered by the presence of another. 

Competition often is related to food and habitat requirements 

and reproductive behavior. Interspecific competition for foods 

is greatest for species occupying similar trophic positions in 

relatively short food chains and for animals living in regions 

of low biological productivity. 

1980 1990 2000 2010

Resilience—In ecology, resilience traditionally 

refers to the ability of a population or biotic 

community to sustain or return to its former 

state after a disturbance. The rate of recovery is 

a measure of resilience determined by the 

population processes involved in restoring 

abundance to healthy, sustainable, or pre-disturbance levels. 

Four categories of productivity (high, medium, low, and very 

low) are used to classify reliance in marine fish populations 

(Musick, 1999). These categories are based on a combination 

of population parameters for intrinsic rate of growth, growth 

coefficient, fecundity, age at maturity, and maximum age. 

Because population parameters were unavailable, resiliency is 

defined here based on estimated population doubling time 

where high = <15 months, medium = 1.4–4.4 years, and 

low = 4.5–14 years. 

Traditional, Cultural, and Economic Values

In August 2009, the U.S. Secretary of Commerce 

approved a Fishery Management Plan for the Arctic 

Management Area. The plan covers U.S. Arctic waters in the 
Chapter 3 47

Chukchi and Beaufort Seas, and acknowledges that changing 

climate may potentially favor the development of commercial 

fisheries. However, until adequate fisheries resource 

assessments are completed, the region remains closed to 

commercial fishing in federal waters. A small salmon fishery 

exists in Kotzebue Sound; in 2010, a small commercial fishery 

for Arctic Ciscoes in the Colville River was terminated. 

Traditional and Cultural Importance.—

Several species of nearshore marine fishes are 

important in subsistence fisheries. The 

protection of traditional lifestyles and 

economies, including these subsistence 

fisheries, is a responsibility of the Federal 

government. Subsistence relates to resource use patterns (for 

example, seasonal round) and values (that is, sustenance, 

kinship, and barter) in coastal communities of northern Alaska.

Commercial Fisheries.—Currently (2016) 

there are no offshore marine fisheries in the 

U.S. Chukchi and Beaufort seas. Changing 

Arctic environmental conditions and shifting 

distributions of species in response to warming 

suggest that there may be fisheries in the 

future. A precautionary approach by fishery managers has been 

adopted that requires the collection of reliable baseline 

information for decision-making and ecosystem management 

(North Pacific Fishery Management Council [North Pacific 

Fishery Management Council, 2009; Wilson and 

Ormseth, 2009]). 

Climate Change

Alaska’s climate is changing at more than twice the rate 

of the rest of the United States (Mellilo and others, 2014). 

Year-to-year and regional variability in air temperatures are 

evident and the warming trend currently is being moderated 

by large-scale cooling associated with the Pacific Decadal 

Oscillation. Even so, climate effects are pronounced and 

are being seen in changes in sea ice, timing of snowmelt, 

widespread glacier retreat, and changes in hydrology (runoff) 

and coastal processes, such as erosion (Markon and others, 

2012). The effects of rising ocean temperatures and ocean 

acidification on marine food webs are of growing regional 

concern with respect to the condition and trends in marine 

ecosystems and human community resilience are of concern. 

Climate changes potentially can affect marine fish in 

numerous ways, leading to distributional changes, increased 

or decreased mortality rates, changes in growth rates, and by 

altering the timing in reproduction (Clow and others, 2011).

Potential Effects of Climate Change.—A 

pole-ward shift of many fish distributions is 

possible as is a reduction or extinction of 

species that are narrowly adapted to Arctic 

environments. Generally, the species are expected to increase 

in abundance if they are currently present in the Bering Sea 

and decrease if they have very low tolerance for temperatures 

greater than 1.5–2.0 °C. However, it is hypothesized in current 

climate projections that temperatures near the ocean floor in 

the northern Bering Sea will remain cold (<2 °C) due to 

persistence of winter sea ice (Sigler and others, 2011). 

Cold-water conditions and other marine ecosystem effects 

related to seasonal sea ice extent and timing of retreat may 

effectively block northward migrations and production of 

exploitable quantities of species, such as pollock and cod, for 

several decades. Shifts in range and other possible climaterelated effects, such as increased predation or competition for 

food, are identified in the species accounts. Only “loose 

qualitative generalizations” are presently possible (Reist and 

others, 2006).

Research Needs

The compilation and review of species information 

for species in U.S. Arctic waters revealed many gaps in life 

history understanding and environmental relations. These 

are evaluated on the basis of a species current fishery and 

community values and ecological significance in marine 

ecosystem structure and function. The needs reflect the 

researcher’s perceptions and their understanding that new 

fishery information is becoming available for the Arctic region 

and that, although Arctic research is currently a national 

priority, some aspects of population ecology will take many 

years of data collection to accurately assess. 

Areas for Future Research.—The preparation 

of individual accounts led to the identification 

of many information gaps in knowledge about 

the biology and ecology of marine species 

including life history, population dynamics, 

and community associations. Generally, 

species life history and ecology gaps are most pronounced 

with respect to: (1) depth and location of pelagic larvae; 

(2) depth, location, and timing of young-of-the-year habitats; 

(3) preferred depth ranges for juveniles and adults; 

(4) spawning seasons; (5) seasonal and ontogenetic 

movements; (6) population genetics and dynamics; (7) prey–

predator relationships and food web relationships; and 

(8) environmental health (multiple stressor effects on fitness). 

Behavioral studies for all life stages are virtually non-existent. 

New information is being developed and, for the lesser-known 

species, gaps may be slowly addressed over time. Priority 

needs, for species having special significance in subsistence 

fisheries and marine food webs or that may be indicator 

species are emphasized in the species accounts. One of two 

categories of identified research need is identified for each 

species. The meaning of the categories [A] and [B] is 

as follows:
48 Alaska Arctic Marine Fish Ecology Catalog

• [A] Many gaps in our understanding of the species life 

history and ecology remain in Alaska (for example, 

research areas 1 through 8). These are high profile 

species in terms of ecological, subsistence, or potential 

fisheries values. Specific research priorities are 

briefly discussed. 

• [B] Most aspects of the species life history and ecology 

are unknown for Alaska (for example, research areas 1 

through 8). Species information will likely accumulate 

over time and focused studies are not warranted at 

this time.

References Cited and Bibliography

A thorough review of scientific literature was done in 

the preparation of the species account. A list of references 

(References Cited [chapter 8]) is provided for each species for 

readers seeking additional information. This list identifies key 

sources of information that make the greatest contributions 

to current knowledge (2014) and understanding. The 

Bibliography section provides a full accounting of all scientific 

literature cited in each species account. For a small number 

of species from the family Cottidae, only a Bibliography 

was possible to provide and this is indicative of the lack of 

information available. Citations are not always in numerical 

order in species accounts because new information became 

available during the production phase of this publication and 

were incorporated into the species accounts as appropriate. 
Pacific Lamprey 49

Pacific and Arctic Lampreys

Pacific Lamprey (Entosphenus tridentatus)

(Gairdner, 1836)

Family Petromyzontidae 

Note: Except for physical description and geographic range data, 

all information is from areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Its rarity in the U.S. Chukchi Sea and absence from the U.S. Beaufort Sea implies an insignificant role in 

regional ecosystem dynamics.

Physical Description/Attributes: Elongate, eel-like body, blue-black to dark brown dorsally, pale or silver ventrally. For 

specific diagnostic characteristics, see Fishes of Alaska, (Mecklenburg and others, 2002, p. 61, as Lampetra tridentata) [1]. 

Swim bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: Eastern U.S. Chukchi Sea [1, 3]. Elsewhere, from Bering Sea south to Punta Canoas, northern Baja California, 

Commander Islands, and Pacific coast of Kamchatka Peninsula, Russia, and Honshu, Japan [1].

Relative Abundance: Rare in U.S. Chukchi Sea, with one record near Cape Lisburne, Alaska [1, 3]. Common in southeastern 

Bering Sea [6]. Widespread at least as far southward as Honshu, Japan [7]. Rare to occasional in marine waters off Commander 

Islands and Pacific coasts of Kamchatka Peninsula, Russia, and Hokkaido, Japan [1].

Pacific Lamprey (Entosphenus tridentatus). Photograph by 

René Reyes, Bureau of Reclamation.

200

200

50

200

50

50

50

50

YUKON

BRITISH

COLUMBIA

NORTHWEST

TERRITORIES

NUNAVUT

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon Kotzebue

Sound

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Pacific Lamprey

Entosphenus tridentatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

Occurrence records

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Pacific Lamprey (Entosphenus tridentatus), within Arctic Outer Continental 

Shelf Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [3, 5]. 
50 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Over continental shelf and slope, near surface to 1,508 m. Most abundant at depths less than 500 m and 

pelagically most abundant above 100 m [7].

Habitats and Life History

Eggs—Size: Small, 1.1–1.2 mm [8]. Time to hatching: 10–24 days, depending on temperature [9, 10]. Habitat: 

Freshwater, benthic [3]; attached to rocks among gravel nests near riffles in rivers [9, 11]. 

Larvae (ammocoetes)—Size at hatching: About 4–5 mm TL [12]. Size at juvenile transformation: From 

4.7–17.0 cm [9, 13]. Days to juvenile transformation: 3–7 years [11]. Actual transformation process can take 

85–126 days [14]. Habitat: Freshwater, benthic [3]; in gravel redds approximately 2–3 weeks after hatching, then 

drifts downstream and burrows into soft sediments of slow, shallow depositional areas along stream banks and in 

pools and eddies [15, 16]. 

Juveniles (macropthalmia)—Age and size: From 3 to 4.5–8 years [13, 15, 16]. Habitat: Pelagic and benthic in 

marine water [3]. Marine (parasitic) phase not well understood. Over continental shelf and slope sometimes far 

offshore [11]. Resides in ocean for 20 months up to 3.5 years, depending on area [7, 13, 17]. 

Adults—Age and size at first maturity: Unknown. Likely from 4.5–8 years [10–12]. Size at First Maturity: Size 

varies from 13–72 cm TL [13]. Maximum age: 9 years [18]. Maximum size: 85 cm TL and at least 0.5 kg [7, 11]. 

Habitat: Freshwater streams and rivers for a few months up to several years before spawning [3, 19]. Substrate: 

Unknown in ocean. Sandy gravel for spawning. Soft sediment for larval rearing [15, 16]. 

Physical/chemical—Temperature: Unknown at sea. Spawns between 13 and 18 °C [12]. Salinity: Fresh to 

marine waters [1].

Behavior

Diel—In the ocean, makes daily vertical migrations into pelagic zone, higher at night perhaps to feed [7]. 

Ammocoete downstream migrations and adult upstream migrations are primarily at night [9, 19]. 

Seasonal—Ammocoete generally transform into juveniles during July through late November [15, 16]. 

Ammocoetes migrate downstream year-round but mainly from autumn through spring. Migration times differ 

among populations. In British Columbia, Canada, after leaving their mud-silt habitat they reside in gravel and 

boulder fields in moderate to strong current streams and then enter seawater from December–June (occasionally 

earlier than December or later than June) [13]. Adults generally return to freshwater rivers and streams in late 

spring and early summer (April–June in British Columbia) [13] and reside there from a few months to several 

years before spawning [19]. Generally, spawning begins the following spring and summer (about May–July) 

depending on river system [19]. 

Reproductive—Semelparous, most die within a month after spawning [16], though some seem to spawn at 

least twice [20]. Spawning occurs in low-gradient streams in sandy gravel usually at riffle heads and in pool 

tailouts [15, 19]. Males initiate nest building and then joined by females. Nests are constructed by fish attaching 

to rocks to lift them out of the nest and by digging down within the nest to line the bottom with loose sand for 

egg attachment [19]. Adults attach themselves side by side to a rock or to each other and release sperm and eggs. 

3-3-1gPacificLamprey

Open ocean Shore

Depth, in meters

1,500

1,200

900

600

300

0

1,500

1,200

900

600

300

0

Juveniles and younger adults

Some fish to 1,508 m

Entosphenous tridentatus

Pacific Lamprey

Benthic distribution

Open ocean No reproductive distribution in seawater Shore

Depth, in meters

Reproductive distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Anadromous.

Spawning, eggs and 

amnocoetes (larvae)

occur in freshwater

Data from outside U.S. Beaufort-Chukchi Seas

Benthic and reproductive distribution of Pacific Lamprey (Entosphenus tridentatus). 
Pacific Lamprey 51

Fertilized eggs drift into nests and attach to rocks. Some adults then cover eggs with rocks or debris [11]. 

Schooling—Unknown at sea. At times, tends to congregate in certain areas in freshwater rivers [9]. 

Feeding—Freshwater ammocoetes are burrowing filter feeders [9]. Macropthalmia begin parasitic feeding on 

fish during seaward migrations [19]. They are parasitic feeders, attaching themselves to fishes using their toothed 

tongues to penetrate scales and skin to suck out body fluid and blood. While feeding, anticoagulants are produced 

which prevents host’s blood from coagulating [19, 21]. In general, host fish are not killed as various surveys 

show high incidences of fish with scars. For instance, off the Fraser River, British Columbia, Canada, 66 percent 

of Sockeye Salmon and 20 percent Coho Salmon had Pacific Lamprey wounds. Lampreys generally attack 

ventrally and anteriorly, leaving one to three holes, with younger fish creating more holes. They have been shown 

in the laboratory to hang on to a host for several days [13]. Feeding ceases during upstream migrations [22].

Populations or Stocks

There have been no studies within the study area. Elsewhere, recent studies show low levels of genetic 

differentiation between populations separated by large geographic distances [19].

Reproduction

Mode—Oviparous, external fertilization [1]. 

Spawning season—Differs with regions, spawning earlier farther north. April–July in British Columbia 

[12]; in southern California occurs as early as January and may continue into at least May [23]. 

Fecundity—10,000–238,000 eggs [12, 16].

Food and Feeding

Food items–Ammocoetes: Detritus, diatoms and algae [19]. Parasitic macropthalmia and adults: Fishes and 

mammals including Greenland and Pacific Halibut, Arrowtooth and Kamchatka Flounders; Sablefish, Pacific 

Hake, Walleye Pollock, Pacific Cod, and Lingcod; Pink, Sockeye, Coho, and Chinook Salmon; Steelhead; 

Yellowmouth and Rougheye Rockfish; and cetaceans [13, 22]. Off Russia, Greenland Halibut were the most 

common prey [22]. 

Trophic level: 4.5 (standard error ±0.80) [18].

Biological Interactions

Predators—Fishes including Sablefish, rockfishes, various sharks, and White Sturgeon [11, 16, 24–26]. 

Ammocoetes are eaten by Coho Salmon [16]. Larger fish eaten by harbor seals, California sea lion, Steller sea 

lion, northern elephant seal, northern fur seal, sperm whales, Pacific White-sided Dolphin, minks, California 

Gulls, Ring-billed Gulls, Western Gulls, Foster’s Terns, Great Blue Herons, and Common Murres [13, 16, 27–31, 

32, 33]. 

Competitors: Pacific Lamprey in seawater [21].

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (t

m 6–8; Fecundity=10,000–106,000) [18]. 

Traditional and Cultural Importance

None reported.
52 Alaska Arctic Marine Fish Ecology Catalog

Commercial Fisheries

Currently, Pacific Lamprey are not commercially harvested.

Potential Effects of Climate Change

Unknown.

Areas for Future Research [B]

Little is known about the ecology and life history of this species in the U.S. Arctic. Research needs include: 

(1) locations of spawning areas, (2) spawning season, (3) size and age at maturity, (4) seasonal and ontogenetic 

movements, (5) population studies, (6) prey, and (7) predators.

References Cited

Beamish, R.J., 1980, Adult biology of the river lamprey (Lampetra ayresi) and the Pacific lamprey (Lampetra tridentata) from 

the Pacific Coast of Canada: Canadian Journal of Fisheries and Aquatic Sciences, v. 37, no. 11, p. 1,906–1,923. [13]

Close, D.A., Fitzpatrick, M.S., and Li, H.W., 2002, The ecological and cultural importance of a species at risk of extinction—

Pacific lamprey: Fisheries, v. 27, no. 7, p. 19–25. [16]

Kostow, K., 2002, Oregon lampreys—Natural history, status, and analysis of management issues—Information reports: 

Corvallis, Oregon Department of Fish and Wildlife, Oregon State University, Klamath Waters Digital Library, accessed at 

http://digitallib.oit.edu/cdm/ref/collection/kwl/id/3483. [9]

Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p. [11]

Luzier, C.W., Schaller, H.A., Brostrom, J.K., Cook-Tabor, C., Goodman, D.H., Nelle, R.D., and others, 2011, Pacific Lamprey 

(Entosphenus tridentatus)—Assessment and template for conservation measures: U.S. Fish and Wildlife Service, 282 p. [19]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Lagler, K.F., Bardach, J.E., and Miller, R.R., 1962, Ichthyology (1st ed.): New York, John Wiley and Sons, Inc., 545 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 
Pacific Lamprey 53

5. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

6. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

7. Orlov, A.M., Savinykh, V.F., and Pelenev, D.V., 2008, Features of the spatial distribution and size structure of the Pacific 

lamprey Lampetra tridentata in the North Pacific: Russian Journal of Marine Biology, v. 34, no. 5, p. 276–287.

8. Yamazaki, Y., Fukutomi, N., Takeda, K., and Iwata, A., 2003, Embryonic development of the Pacific lamprey, Entosphenus 

tridentatus: Zoological Science, v. 20, no. 9, p. 1,095–1,098.

9. Kostow, K., 2002, Oregon lampreys—Natural history, status, and analysis of management issues—Information reports: 

Corvallis, Oregon Department of Fish and Wildlife, Oregon State University, Klamath Waters Digital Library, accessed at 

http://digitallib.oit.edu/cdm/ref/collection/kwl/id/3483.

10. Hart, J.L., 1973, Pacific fishes of Canada: Ottawa, Fisheries Research Board of Canada, 740 p.

11. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

12. Wang, J.C.S., 1981, Taxonomy of the early life stages of fishes—Fishes of the Sacramento-San Joaquin Estuary and Moss 

Landing Harbor-Elkhorn Slough, California: Concord, California, Ecological Analysts, Inc., 168 p.

13. Beamish, R.J., 1980, Adult biology of the river lamprey (Lampetra ayresi) and the Pacific lamprey (Lampetra tridentata) 

from the Pacific Coast of Canada: Canadian Journal of Fisheries and Aquatic Sciences, v. 37, no. 11, p. 1,906–1,923.

14. McGree, M., Whitesel, T.A., and Stone, J., 2008, Larval metamorphosis of individual Pacific lampreys reared in captivity: 

Transactions of the American Fisheries Society, v. 137, no. 6, p. 1,866–1,878.

15. Pletcher, T.F., 1963, The life history and distribution of lampreys in the salmon and certain other rivers in British Columbia, 

Canada: Vancouver, University of British Columbia, Master’s thesis, 195 p.

16. Close, D.A., Fitzpatrick, M.S., and Li, H.W., 2002, The ecological and cultural importance of a species at risk of 

extinction—Pacific lamprey: Fisheries, v. 27, no. 7, p. 19–25.

17. Kan, T.T., 1975, Systematics, variation, distribution, and biology of lampreys of the genus Lampetra in Oregon: Corvallis, 

Oregon State University, Ph.D. dissertation, 194 p.

18. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

19. Luzier, C.W., Schaller, H.A., Brostrom, J.K., Cook-Tabor, C., Goodman, D.H., Nelle, R.D., and others, 2011, Pacific 

Lamprey (Entosphenus tridentatus)—Assessment and template for conservation measures: U.S. Fish and Wildlife Service, 

282 p.

20. Michael, J.H., Jr., 1984, Additional notes on the repeat spawning by Pacific lamprey: California Fish and Game, v. 70, no. 3, 

p. 186–188.

21. Scott, W.B., and Crossman, E.J., 1973, Freshwater fishes of Canada: Fisheries Research Board of Canada, Bulletin 184, 

966 p.

22. Abakumov, A., 1964, The oceanic period in the life cycle of the lamprey Entosphenus tridentatus (Richardson, 1836), in

Moiseev, P.A., ed., Soviet fisheries investigations in the Northeast Pacific, Part II, Moiseev: Jerusalem, Israel Program for 

Scientific Translations, p. 268–270.

23. Chase, S.D., 2001, Contributions to the life history of adult Pacific lamprey (Lampetra tridentata) in the Santa Clara River 

of southern California: Bulletin of the Southern California Academy of Sciences, v. 100, no. 2, p. 74–85.
54 Alaska Arctic Marine Fish Ecology Catalog

24. Razum, A., 1952, Food of the dogfish, Squalus acanthias and the brown smoothhound, Rhinotriacis henlei, in San Francisco 

Bay, California: Berkeley, University of California, Master’s thesis, 114 p.

25. Ebert, D.A., 1986, Biological aspects of the sixgill shark, Hexanchus griseus: Copeia, no. 1, p. 131–135.

26. Brodeur, R.D., Lorz, H.V., and Pearcy, W.G., 1987, Food habits and dietary variability of pelagic nekton off Oregon and 

Washington, 1979–1984: U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical 

Report NMFS 57.

27. Antonelis, G.A., and Fiscus, C.H., 1980, The pinnipeds of the California current: California Cooperative Oceanic Fisheries 

Investigations Reports, v. 21, p. 68–78.

28. Kawakami, T., 1980, A review of sperm whale food: Scientific reports of the Whales Research Institute, no. 32, p. 199–218.

29. Stroud, R.K., Fiscus, C.H., and Kajimura, H., 1980, Food of the Pacific white-sided dolphin, Lagenorhynchus obliquidens, 

Dall’s porpoise, Phocoenoides dalli, and northern fur seal, Callorhinus ursinus, off California and Washington: Fishery 

Bulletin, v. 78, no. 4, p. 951–959.

30. Harry, G.Y., and Hartley, J.R., 1981, Northern fur seals in the Bering Sea, in Hood, D.W., and Calder, J.A., eds., The Eastern 

Bering Sea shelf—Oceanography and resources: Seattle, University of Washington Press, p. 847–867.

31. Orr, A.J., Banks, A.S., Mellman, S., Huber, H.R., and DeLong, R.L., 2004, Examination of the foraging habits of Pacific 

harbor seal (Phoca vitulina richardsoni) to describe their use of the Umpqua River, Oregon, and their predation on 

salmonids: Fishery Bulletin, v. 102, no. 1, p. 108–117.

32. Merrell, T.R., 1959, Gull food habits on the Columbia River: Fish Commission of Oregon Research Briefs, v. 7, no. 1, p. 82.

33. Matthews, D.R., 1983, Feeding ecology of the common murre, Uria aalge, off the Oregon coast: Eugene, University of 

Oregon, Master’s thesis, 108 p.
Arctic Lamprey 55

Arctic Lamprey (Lethenteron camtschaticum)

(Tilesius, 1811)

Family Petromyzontidae 

Colloquial Name: Iñupiat: Nimiqiaq [24]

Ecological Role: The extent of this lamprey’s parasitisms is 

unknown in U.S. Chukchi and Beaufort Seas.

Physical Description/Attributes: Elongate, eel-like body, blueblack to dark brown dorsally, silvery when fresh on sides and 

ventral surface, with blackish blotch on second dorsal fin and on tail. For specific diagnostic characteristics see Fishes of Alaska

(Mecklenburg and others, 2002, p. 62) [1]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [3]. Elsewhere in Alaska, south through Bering Sea to Kenai Peninsula, Gulf of Alaska. 

Worldwide, White Sea and coasts of southern Barents Sea eastward off Siberia to Beaufort Sea off Anderson River, Canada; in 

western Pacific Ocean, south to Honshu, Japan, and Korean Peninsula, and East-Finnmark, Norway, in eastern Atlantic Ocean. 

Not in western Atlantic [3].

Relative Abundance: Apparently common in some drainages of the U.S. Chukchi and Beaufort Seas. However, abundance in 

these drainages and in marine waters is poorly described. Presence at sea is typically indicated by wounds on pelagic fishes. 

The most common lamprey in Alaska and, although abundance patterns are unknown, thought to occur in high numbers in 

localized areas [1, 6]. Common in Sea of Japan and around Sakhalin Island, Russia [7, 8].

Arctic Lamprey (Lethenteron camtschaticum), 176 mm TL, 

Norton Sound, northeastern Bering Sea, 2002. Photograph by 

C.W. Mecklenburg, Point Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Lamprey

Lethenteron camtschaticum

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Arctic Lamprey (Lethenteron camtschaticum) within Arctic Outer Continental 

Shelf Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [3, 5]. 
56 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Anadromous. Pelagic at sea over continental shelf to bottom depth of 50 m [1].

Habitats and Life History

There are two life-history types, anadromous-parasitic and fluvial-nonparasitic. Eggs and ammocoete larvae of 

both types are demersal in freshwater lakes and streams. There are three juvenile/adult forms: typically 

anadromous, anadromous early maturing forma praecox (predominantly males), and resident freshwater. Both 

forms of anadromous fish are pelagic and migrate to sea. When mature they return to freshwater to spawn. The 

non-parasitic resident fish remain exclusively in fresh water until spawning [1, 9, 10]. 

Eggs—Size: As large as 1.25 mm, average of 0.8 mm [9]. Time to hatching: About 1 month after spawning [9].

Habitat: Pebble-sandy bottoms in rivers. 

Ammocoetes (larvae)—Size at hatching: 6.8 mm long. Size at juvenile transformation: 13.1–16.8 cm [9]. Time

to juvenile transformation: 4 years and longer [9]. Habitat: Sedentary burrowers in river and lake bottoms [11]. 

Juveniles/smolts (anadromous forms)—Age and size: 4–5 years. Transformation to smolt stage takes

approximately 6 months and ends in downstream migration to the sea [10]. Size ranges from 13.0–16.8 to as long

as 16.5–21.7 cm TL and from 2.8–4.4 to about 3.1–9.1 g. [10]. Habitat: Pelagic, in downstream migrations to the

sea [10]. 

Adults—Age and size at first maturity: 7 years for anadromous form, 6 years for forma praecox and 5 years for

freshwater residents. Typically, anadromous form is 25.0–35.0 cm TL and 30–88 g for males and 17.4–33.0 cm

TL and 30–75 g for females [9, 10]. Anadromous forma praecox (predominantly males) is 14.5–22.0 cm TL and

3.2–15 g [9]. Freshwater residents are 11.3–13.9 cm and 1.6–5.0 g for males, and 110–141 cm TL and 2.1–4.5 g 

for females. Maximum age: Same as age at first maturity. Maximum size: 62.5 cm TL (anadromous form) [1]. 

Habitat: Anadromous form migrates downstream and becomes pelagic in shallow marine waters over continental 

shelf. Forma praecox remains in seawater from several months to 1 year, whereas typically anadromous lamprey 

remain as much as 1 year longer [9]. 

Substrate—In freshwater, gravel-sand for spawning and muddy sediments for ammocoete rearing [1]. 

Physical/chemical—Temperature: Spawning occurs between 12 and 15 °C in southwestern Alaska [12]. Salinity: 

Marine and fresh waters.

Behavior

Diel—Ammocoetes are primarily active at night and burrow into sediments during day [6]. 

Seasonal—Metamorphosed ammocoetes migrate downstream to sea during August–November in Alaska [10] 

and May–July in Russia [9]. 

Reproductive—Adults migrate upstream to spawn in spring. Redds are constructed in riffles with pebbly-gravel 

bottom where sand prevails [10]. Redds are made by lampreys sucking on to rocks and swimming them away 

[13]. There is group spawning behavior in fast currents and paired behavior in slow, nearshore zones. In group 

3-1-2g_ArcticLamprey

No reproductive distribution in seawater Open ocean Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Juveniles only

Lethenteron camtschaticum

Arctic Lamprey

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning, eggs and larvae 

all occur in freshwater

Data from outside U.S. Beaufort-Chukchi Seas

Maturing adults return to

freshwater to spawn

Benthic and reproductive distribution of Arctic Lamprey (Lethenteron camtschaticum).
Arctic Lamprey 57

behavior, 6–44 individuals attach themselves by sucking on to each other and drifting downstream. Numerous 

males may attach to one female. Afterwards, individuals return to spawning redds. Females lay several batches 

of eggs in redd. One batch of eggs may be fertilized by several males. After spawning, fish stir up silt and small 

stones to cover the eggs [13]. Adults die after spawning [1]. 

Schooling—Migrating adults frequently congregate in large numbers, particularly around obstructions [14]. 

Feeding—–Ammocoetes are filter feeders, whereas anadromous juveniles and adults are parasitic, feeding on 

other fish tissues and blood. Freshwater residents cease feeding upon sexual maturity [9, 10].

Populations or Stocks

There have been no studies. 

Reproduction

Mode—Separate sexes, oviparous [9, 13]. 

Spawning season—Spring in southwestern Alaska, generally late May to early July [12]. 

Fecundity: 12,272–34,586 eggs [9].

Food and Feeding

Food items—Ammocoetes filter-feed on small aquatic invertebrates, algae and fine organic debris [9]. Adults 

parasitize fish, including Pacific salmon, Starry Flounder, Saffron Cod, Least Cisco, Arctic Cisco, Broad 

Whitefish, Pacific Herring and smelt [11, 15–17]. 

Trophic level—4.5 (standard error ±0.81) [18].

Biological Interactions

Predators—All life stages are preyed on by various fishes including Burbot, Northern Pike, Dolly Varden, and 

Inconnu; also taken by gulls, especially when lamprey are concentrated in shallow streams during migration [19]. 

Competitors—Pacific Lamprey in seawater [14]. In Alaska, often found co-occurring with Alaskan Brook 

Lamprey (L. alaskensis) [1].

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (t

m 4–5) [18].

Traditional and Cultural Importance

None reported. Alaskan Natives on the Yukon and Kuskokwim Rivers have taken them in quantity for food using 

dip nets and sharped sticks [20, 21]. A small commercial fishery on the Yukon River was started in 2003 [6]. 

Commercially harvested in Sea of Okhotsk [22]. 

Commercial Fisheries

Currently, in Alaska, Arctic Lamprey are not commercially fished.
58 Alaska Arctic Marine Fish Ecology Catalog

Potential Effects of Climate Change

Unknown. 

Areas for Future Research [B]

Little is known about the ecology of this species from this region. Research needs include: (1) depth and location 

of pelagic larvae’ (2) depth, location, and timing of young-of-the-year benthic recruitment; (3) preferred depth 

ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic movements; (6) population 

studies; (7) prey; and (8) predators.

Remarks

This is the most abundant and widely distributed lamprey in Alaska [23].

References Cited

Kucheryavyi, A.V., Savvaitova, K.A., Gruzdeva, M.A., and Pavlov, D.S., 2007, Sexual dimorphism and some special traits of 

spawning behavior of the Arctic lamprey Lethenteron camtschaticum: Journal of Ichthyology, v. 47, no. 7, p. 481–485. [13]

Kucheryavyi, A.V., Savvaitova, K.A., Pavlov, D.S., Gruzdeva, M.A., Kuzishchin, K.V., and Stanford, J.A., 2007, Variations of 

life history strategy of the Arctic lamprey Lethenteron camtschaticum from the Utkholok River (Western Kamchatka): Journal 

of Ichthyology, v. 47, no. 1, p. 37–52. [9]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Reference 1. [3]

Nazarov, D.Y., Kucheryavyi, A.V., Savvaitova, K.A., Gruzdeva, M.A., Kuzishchin, K.V., and Pavlov, D.S., 2011, Population 

structure of Arctic lamprey Lenthenteron camtschaticum from the Kol River (western Kamchatka): Journal of Ichthyology, v. 

51, no. 4, p. 277–290. [10]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Lagler, K.F., Bardach, J.E., and Miller, R.R., 1962, Ichthyology (1st ed.): New York, John Wiley and Sons, Inc., 545 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Reference 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

6. Alaska Department of Fish and Game, 2011, Arctic lamprey (Lampetra camtschatica) Tilesius, 1811: Alaska Department of 

Fish and Game, accessed November 2011 at http://www.adfg.alaska.gov/static/species/speciesinfo/_aknhp/Arctic_lamprey.

pdf.
Arctic Lamprey 59

7. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

8. Safronov, S.N., and Nikiforov, S.N., 2003, The list of pisciformes and fishes of the fresh and brackish waters of Sakhalin: 

Journal of Ichthyology, v. 43, no. 1, p. 38–49.

9. Kucheryavyi, A.V., Savvaitova, K.A., Pavlov, D.S., Gruzdeva, M.A., Kuzishchin, K.V., and Stanford, J.A., 2007, Variations 

of life history strategy of the Arctic lamprey Lethenteron camtschaticum from the Utkholok River (Western Kamchatka): 

Journal of Ichthyology, v. 47, no. 1, p. 37–52.

10. Nazarov, D.Y., Kucheryavyi, A.V., Savvaitova, K.A., Gruzdeva, M.A., Kuzishchin, K.V., and Pavlov, D.S., 2011, Population 

structure of Arctic lamprey Lenthenteron camtschaticum from the Kol River (western Kamchatka): Journal of Ichthyology, 

v. 51, no. 4, p. 277–290.

11. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

12. Heard, W.R., 1966, Observations on lampreys in the Naknek River system of southwest Alaska: Copeia, no. 2, p. 332–339.

13. Kucheryavyi, A.V., Savvaitova, K.A., Gruzdeva, M.A., and Pavlov, D.S., 2007, Sexual dimorphism and some special traits 

of spawning behavior of the Arctic lamprey Lethenteron camtschaticum: Journal of Ichthyology, v. 47, no. 7, p. 481–485.

14. Scott, W.B., and Crossman, E.J., 1973, Freshwater fishes of Canada: Fisheries Research Board of Canada, Bulletin 184, 

966 p.

15. McAllister, D.E., 1963, Fishes of the 1960 “Salvelinus” program from western Arctic Canada: National Museum of Canada 

Bulletin, no. 185, p. 17–39.

16. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.

17. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

18. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

19. Bond, W.A., and Erickson, R.N., 1987, Fishery data from Phillips Bay, Yukon, 1985: Winnipeg, Manitoba, Canadian Data 

Report of Fisheries and Aquatic Sciences, Central and Artic Region, Department of Fisheries and Oceans, no. 635, 47 p.

20. Turner, L.M., 1886, Contributions to the natural history of Alaska—Arctic series of publications, no. 2, Washington: U.S. 

Government Printing Office, 226 p.

21. Evermann, B.W., and Goldsborough, E.L., 1907, The fishes of Alaska: Bulletin of the United States Bureau of Fisheries, 

v. 26, p. 219–360.

22. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

23. Mecklenburg, C.W., and Mecklenburg, T.A., 2011, Arctic lamprey—Lethenteron camtschaticum (Tilesius, 1811): Arctic 

Ocean Diversity—Census of Marine Life, http://www.arcodiv.org/Fish/Lethenteron_camtschaticum.

24. Carothers, C., Cotton, S., and Moerlein, K., 2013, Subsistence use and knowledge of salmon in Barrow and Nuiqsut, 

Alaska: U.S. Department of the Interior, Bureau of Ocean Energy Management, OCS Study BOEM 2013-0015, 52 p.
60 Alaska Arctic Marine Fish Ecology Catalog

Spotted Spiny Dogfish to Bering Cisco

Spotted Spiny Dogfish (Squalus suckleyi)

(Girard, 1855)

Family Squalidae 

Note on taxonomy: Meristic, morphometric, and molecular data 

demonstrate that Squalus suckleyi is a distinct species from

S. acanthias (Linnaeus, 1758) [1]. The latter species does not occur 

in the North Pacific, and previous reports of S. acanthias in the 

North Pacific are assumed to represent S. suckleyi. Information 

presented here is only from data or reports of Squalus in North Pacific waters.

Note: Except for geographic range data, all information is from areas outside of the Chukchi and Beaufort Seas.

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: A rare species in the U.S. Chukchi Sea and absent from the U.S. Beaufort Sea. The species has a very limited 

role and little significance in regional food webs.

Physical Description/Attributes: Gray or brown dorsally merging into lighter sides and belly with one or two rows of 

conspicuous white spots on sides. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, 

p. 88) [1] and [2]. Swim bladder: Absent, as with other cartilaginous fishes [1]. Antifreeze glycoproteins in blood serum: 

Unknown. Dorsal spines are venomous [3].

Range: U.S. Chukchi Sea at Kotzebue Sound [1, 4]. Elsewhere in Alaska, from Bering Sea and Aleutian Islands, eastward in the 

Gulf of Alaska. Worldwide, from Koreas and Japan northwards to Bering Sea off Kamchatka Peninsula, Russia, Sea of Okhotsk 

and Sakhalin Island, and from British Columbia, Canada, and Washington south to southern Baja California [2, 5].

Spotted Spiny Dogfish (Squalus suckleyi). Photograph by 

NMFS-Alaska Fisheries Science Center, RACE Division.
Spotted Spiny Dogfish 61

Relative Abundance: Rare in U.S. Chukchi Sea, with one record of occurrence near Kotzebue [1]. Common from Kodiak 

Island, Gulf of Alaska and southward into Baja California, and in Sea of Japan [7–9]. Very rare in northern Bering Sea [1, 10, 

11]. Appears to be increasing in abundance in southern Bering Sea [10]. 

tac14-5222_fig3-2-1m_Squalus_suckleyi

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Spotted Spiny Dogfish

Squalus suckleyi

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Spotted Spiny Dogfish (Squalus suckleyi) within Arctic Outer Continental Shelf 

Planning Areas based on review of published literature and specimens from historical and recent collections 

[4, 6]. 
62 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Very shallow waters to at least 1,236 m [9], typically 250 m or less [5]. Juveniles are born in midwaters at depths 

of 10–140 m [12], and over bottom depths of 50–111 m [13].

3-2-1gSpottedSpinyDogfish

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

500

400

300

200

100

0

Younger juveniles

Some fish to 1,236 m

Squalus suckleyi

Spotted Spiny Dogfish

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Depths where

females give birth

Spawning and eggs

Depth range of mating is uncertain. 

Potential location is based on depth 

range of adults. Eggs and embryos 

are gestated within females.

Older juveniles 

and adults

Benthic and reproductive distribution of Spotted Spiny Dogfish (Squalus suckleyi).

Habitats and Life History

Eggs—Size: 3–4 cm [14]. Time to hatching: Fertilized eggs are contained within candles (a thin membrane 

containing multiple eggs) and incubated within the female’s uterus. Candle membrane dissolves and embryos 

become free within the uterus within 4–6 months [5]. Habitat: In utero [5]. 

Embryos—Age and size: From about 4–6 months to 22 months (<10 to 22.5–30 cm TL) [5, 13]. Habitat:

Embryos are completely dependent on their yolk-sacs and are gestated within the uterus [5, 13]. 

Juveniles—Size: 22.5–26.3 cm at birth to about 60 cm TL [5, 13]. Habitat: Pelagic, in water column, near 

surface and becoming benthic as they grow larger and near sexual maturity [5, 13]. 

Adults—Age and size at first maturity: Based on the most recent study (off British Columbia), a few females 

mature at about 80 cm TL (24 years), 50 percent matured at 93.9 cm (36 years), and almost all fish are mature 

at 110 cm (62 years) [15]. 100 percent of females matured at 119 cm [14]. A few males off British Columbia 

matured at 72 cm TL (15 years), 50 percent at 78 cm TL (19 years), and all at 94 cm [14]. In the North Pacific 

median size and age at maturity is 80–100 cm TL. (35.5 years) for females and 70–80 cm TL 18.5 for males [2]. 

Maximum age: 80 to possibly 100 years [5]. Maximum size: About 140 cm [10]. Habitat: Benthopelagic, in a 

wide depth range [5]. 

Substrate—Unknown. Have been taken over cobble [16]. 

Physical/chemical—Temperature: 0–15 °C [17]; prefers less than 7 °C, often migrating horizontally and 

vertically to follow temperature preference [9]. Salinity: Marine, but can tolerate freshwater for short periods [5].

Behavior

Diel—Migrates closer to surface at night [5, 10] and may be more active at night [16]. 

Seasonal—Makes seasonal feeding migrations, moving north and inshore as waters warm in spring [10]. Highly 

mobile in many areas, though movements are not completely predictable. In the North Pacific, many tagged 

fish were recaptured close to their release site, but some made extensive migrations (as far as 7,000 km) [16]. 

Reproductive—Males mate every year and females every other year. Smaller males mate earlier in the season 

[18]. Because of the female’s long gestation period (22–24 months), she does not release young every year [9, 

18, 19]. Females commonly give birth in shallow bays and estuaries or in mid-water at depths of 50–111 m [13]. 

Schooling—Forms large schools [5]. Sexes tend to segregate into separate schools around time of 

parturition [13]. 

Feeding—Opportunistic feeders [5], congregating in schools where prey is abundant and sensed by smell [20].

Populations or Stocks

There have been no studies.
Spotted Spiny Dogfish 63

Reproduction

Mode—Aplacental viviparous. Internal fertilization [2]. 

Parturition season—September–January, probably peaks in late autumn [14, 18]. 

Fecundity—Litters as high as 20, averaging between 2–12 [9, 12, 14]. Number of pups increases as size of 

female increases [13].

Food and Feeding

Food items—Fishes are a very important, particularly for larger individuals. However, squids, octopuses, 

medusae, ctenophores, crustaceans (for example, shrimps, euphausiids, and amphipods) and polychaetes also are 

often consumed [21–25]. 

Trophic level—4.3 (standard error 0.67) (based on trophic level of S. acanthias) [26].

Biological Interactions

Predators—Various larger sharks (for example, Salmon Sharks, White Sharks, Pacific Sleeper Sharks), bald 

eagles, and marine mammals such as Steller sea lion, northern elephant seal, and sperm whale [21, 27–31]. 

Competitors—Likely various larger cods, flatfishes, and other macrocarnivores.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time is more than 14 years (rm=0.034; K=0.03–0.07; t

m=10–30; tmax=75; 

Fecundity=1) (based on resilience of S. acanthias) [26].

Traditional and Cultural Importance

None reported

Commercial Fisheries

Currently, Spiny Spotted Dogfish are not commercially fished.

Potential Effects of Climate Change

A wider distribution of this species in the Bering Sea occurred after 2000, possibly associated with recent climate 

change [10]. This species would be expected to move northwards into the Chukchi Sea as waters warm.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. If the species becomes more 

common, research needs include: (1) preferred depth ranges for juveniles and adults, (2) growth rates and size at 

maturation, (3) birthing season, (4) seasonal and ontogenetic movements, (5) population studies, (6) prey, and (7) 

predators.
64 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Ebert, D.A., 2003, Sharks, rays, and chimaeras of California: Berkeley, University of California Press, California Natural 

History Guide, v. 71, 297 p. [9]

Ebert, D.A., White, W.T., Goldman, K.J., Compagno, L.J.V., Daly-Engel, T.S., and Ward, R.D., 2010, Resurrection and 

redescription of Squalus suckleyi (Girard, 1854) from the North Pacific, with comments on the Squalus acanthias subgroup 

(Squaliformes: Squalidae): Zootaxa, v. 2612, p. 22–40. [2]

Ketchen, K.S., 1972, Size at maturity, fecundity, and embryonic growth of the spiny dogfish (Squalus acanthias) in British 

Columbia waters: Journal of the Fisheries Research Board of Canada, v. 29, no. 12, p. 1,717–1,723. [14]

Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p. [5]

Tribuzio, C.A., 2004, An investigation of the reproductive physiology of two North Pacific shark species—Spiny Dogfish 

(Squalus acanthias) and salmon shark (Lamna ditropis): Seattle, University of Washington, Master’s thesis, 147 p. [13]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Ebert, D.A., White, W.T., Goldman, K.J., Compagno, L.J.V., Daly-Engel, T.S., and Ward, R.D., 2010, Resurrection and 

redescription of Squalus suckleyi (Girard, 1854) from the North Pacific, with comments on the Squalus acanthias subgroup 

(Squaliformes: Squalidae): Zootaxa, v. 2612, p. 22–40.

3. Halstead, B.W., 1995, Dangerous marine animals that bite, sting, or are non-edible: Centreville, Maryland, Cornell 

Maritime Press, 275 p.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

U.S. Department of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Report NMFS 66, 

151 p.

8. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

9. Ebert, D.A., 2003, Sharks, rays, and chimaeras of California: Berkeley, University of California Press, California Natural 

History Guide, v. 71, 297 p.

10. Orlov, A.M., Savinykh, V.F., Kulish, E.F., and Pelenev, D.V., 2012, New data on the distribution and size composition of the 

North Pacific spiny dogfish Squalus suckleyi (Girard, 1854): Scientia Marina, v. 76, no. 1, p. 111–122.

11. Stevenson, D.E., Orr, J.W., Hoff, G.R., and McEachran, J.D., 2007, Field guide to sharks, skates, and ratfish of Alaska: 

Fairbanks, University of Alaska, Alaska Sea Grant Program, 77 p.

 12. Ketchen, K.S., 1986, The spiny dogfish (Squalus acanthias) in the northeast Pacific and a history of its utilization: Canadian 

Special Publication of Fisheries and Aquatic Sciences, v. 88, 78 p.

13. Tribuzio, C.A., 2004, An investigation of the reproductive physiology of two North Pacific shark species—Spiny Dogfish 

(Squalus acanthias) and salmon shark (Lamna ditropis): Seattle, University of Washington, Master’s thesis, 147 p.
Spotted Spiny Dogfish 65

14. Ketchen, K.S., 1972, Size at maturity, fecundity, and embryonic growth of the spiny dogfish (Squalus acanthias) in British 

Columbia waters: Journal of the Fisheries Research Board of Canada, v. 29, no. 12, p. 1,717–1,723.

15. Saunders, M.W. and McFarlane, G.A., 1993, Age and length of maturity of the female spiny dogfish, Squalus acanthias, in 

the Strait of Georgia, British Columbia, Canada: Environmental Biology of Fishes, v. 38, p. 49–57.

16. Miller, B.S., Simenstad, C.A., Moulton, L.L., Fresh, K.L., Funk, F.C., Karp, W.A., and others, 1977, Puget Sound baseline 

program—Nearshore fish survey: University of Washington, Fishery Research Institute, FRI-UW-7710, 219 p.

17. Orlov, A.M., 2004, Migrations of various fish species between Asian and American waters in the North Pacific Ocean: 

Aqua, Journal of Ichthyology and Aquatic Biology of Fishes, v. 8, no. 3, p. 109–124.

18. Jones, B.C., and Geen, G.H., 1977, Reproduction and embryonic development of spiny dogfish (Squalus acanthias) in the 

Strait of Georgia, British Columbia: Journal of the Fisheries Research Board of Canada, v. 34, no. 9, p. 1,286–1,292.

19. Pratt, H.L., and Carrier, J.C., 2005, Elasmobranch courtship and mating behavior, in Hamlett, W., ed., Reproductive 

behavior and phylogeny of elasmobranchs: Queensland, Australia, Science Publishers, Inc., p. 129–169.

20. Hart, J.L., 1973, Pacific fishes of Canada: Ottawa, Fisheries Research Board of Canada, 740 p.

21. Fraser, C.M., 1923, Ichthyological notes: Contributions to Canadian Biology, New Series, v. 1, no. 14, p. 287–295.

22. Fraser, C.M., 1946, Food of fishes: Transactions of the Royal Society of Canada, v. 45, sec. 5, p. 33–39.

23. Beamish, R.J., and Smith, M.S., 1976, A preliminary report on the distribution, abundance, and biology of juvenile spiny 

dogfish (Squalus acanthias) in the Strait of Georgia and their relationship with other fishes: Environment Canada, Fisheries 

and Marine Service, Technical Report 629, 44 p.

24. Simenstad, C.A., Miller, B.S., Nyblade, C.F., Thornburgh, K., and Bledsoe, L.J., 1979, Food web relationships of northern 

Puget Sound and the Strait of Juan de Fuca—A synthesis of the available knowledge: National Oceanic and Atmospheric 

Administration/Marine Ecosystems Analysis Puget Sound Project, Prepared for Office of Environmental Engineering and 

Technology, United States Environmental Protection Agency, 334 p.

25. Tanasichuk, R.W., Ware, D.M., Shaw, W., and McFarlane, G.A., 1991, Variations in diet, daily ration, and feeding 

periodicity of Pacific hake (Merluccius productus) and spiny dogfish (Squalus acanthias) off the lower west coast of 

Vancouver Island: Canadian Journal of Fisheries and Aquatic Sciences, v. 48, no. 11, p. 2,118–2,128.

26. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

27. Antonelis, G.A., and Fiscus, C.H., 1980, The pinnipeds of the California current: California Cooperative Oceanic Fisheries 

Investigations Reports, v. 21, p. 68–78.

28. Tricas, T.C., and McCosker, J.E., 1984, Predatory behavior of the white shark (Carcharodon carcharias), with notes on 

biology: Proceedings of the California Academy of Sciences, v. 43, no. 14, p. 221–238.

29. Flinn, R.D., Trites, A.W., Gregr, E.J., and Perry, R.I., 2002, Diets of fin, sei, and sperm whales in British Columbia—An 

analysis of commercial whaling records, 1963–1967: Marine Mammal Science, v. 18, no. 3, p. 663–679.

30. Hulbert, L.B., Sigler, M.F., and Lunsford, C.R., 2006, Depth and movement behaviour of the Pacific sleeper shark in the 

north-east Pacific Ocean: Journal of Fish Biology, v. 69, no. 2, p. 406–425.

31. Sigler, M.F., Hulbert, L.B., Lunsford, C.R., Thompson, N.H., Burek, K., O’Corry-Crowe, G., and others, 2006, Diet of 

Pacific sleeper shark, a potential Steller sea lion predator, in the north-east Pacific Ocean: Journal of Fish Biology, v. 69, 

no. 2, p. 392–405.
66 Alaska Arctic Marine Fish Ecology Catalog

Arctic Skate (Amblyraja hyperborea)

(Collett, 1879)

Family Rajidae 

Note: Except for geographic range data, all information is from areas outside of the 

study area.

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Arctic Skate have only rarely been observed in deeper waters of the 

Alaska Beaufort Sea. Its role in benthic ecosystem dynamics, especially over shelf break 

and slope habitats is presently unknown.

Physical Description/Attributes: Brown or grayish brown, often with dark and light 

round spots. Body is flat, with wing-like pectoral fins, mouth on underside; has long ratlike tail with two small dorsal fins near the tip. For specific diagnostic characteristics, see 

Jensen (1948, p. 31–43) [1] and Stehmann and Bürkel (1984, p. 174) [2]. Swim bladder: 

Absent [3]. Antifreeze glycoproteins in blood serum: Unknown.

Range: Continental slope off U.S. Beaufort Sea [4]. Practically circumpolar; polar basins and south to western Canada, Davis 

Strait, Greenland, Iceland, Faroe-Shetland Ridge, Barents Sea and northern Norway [1, 4, 5].

Relative Abundance: Absent from U.S. Beaufort Sea continental shelf, one record from the continental slope about 50 miles 

north-northeast of Brownlow Point at 70°51’N, 145°17’W; absent from Chukchi Sea [4, 7]. Common off east and west 

Greenland, throughout the Norwegian Basin, and in Barents Sea [1, 5].

Arctic Skate (Amblyraja hyperborea), 

continental slope off Barents Sea, 

2011. Photograph by Arve Lynghammar, 

University of Tromsø, Norway.

Geographic distribution of Arctic Skate (Amblyraja hyperborea) within Arctic Outer Continental Shelf Planning 

Areas [6] based on review of published literature and specimens from historical and recent collections [4, 7]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Skate

Amblyraja hyperborea

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Arctic Skate 67

Depth Range: Typically between 300 and 1,500 m [2], with few records as shallow as 200 m [6] or as deep as 2,640 m [8]. The 

one specimen from the slope off the U.S. Beaufort Sea was taken at a depth of 357 m [7].

Habitats and Life History

Eggs—Female lays two egg cases, each with one egg [1]. Size: Egg cases measure 81–125 × 54–77 mm [2]. 

Time to hatching: Unknown. Habitat: Benthic [2]. 

Larvae—Eggs develop through larval stage to juvenile within the egg case [1]. Size at hatching: 15–16 cm [5]. 

Habitat: Benthic [2]. 

Juveniles—Age and size: Unknown. Habitat: Muddy bottoms [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 92 cm and 5.2 kg 

[5]. Habitat: Benthic, in deep water on the continental slopes and basins of the Arctic Ocean [1, 2, 4]. 

Substrate—Muddy bottoms [5]. 

Physical/chemical—Temperature: Mainly between -1.3 [1] and 1.5 °C [2], reported at 4 °C [7]. Salinity: 

Marine [3].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling: Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [1, 2, 5, 9]. 

Spawning season—Unknown. 

Fecundity—Less than 100 [10].

Benthic and reproductive distribution of Arctic Skate (Amblyraja hyperborea). 

Open ocean Shore

Depth, in meters

3,000

2,400

1,800

1,200

600

0

3,000

2,400

1,800

1,200

600

0

Some fish to 2,640 m

Amblyraja hyperborea

Arctic Skate

Benthic distribution

Data from outside U.S. Beaufort-Chukchi Seas

Open ocean No data for this species Shore

Depth, in meters

 Adults and juveniles

Reproductive distribution

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Depths documented

in U.S. Beaufort Sea

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown
68 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food Items—Benthic and pelagic crustaceans such as shrimp, as well on fishes [1, 5]. 

Trophic level—3.84 (standard error 0.58) [10]

Biological Interactions

Predators—Unknown. 

Competitors—Perhaps eelpouts and other benthic feeders.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time is 4.5–14 years (Fecundity assumed to be <100) [10].

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Arctic Skate are not commercially fished.

Potential Effects of Climate Change

Unknown.

Areas for Future Research [B]

Little is known about the ecology and life history of this species in the study area. In particular, research needs 

include: (1) preferred depth ranges for juveniles and adults, (2) growth rates and size at maturity, (3) spawning 

season, (4) seasonal and ontogenetic movements, (5) population studies, (6) prey, and (7) predators. 
Arctic Skate 69

References Cited

Jensen, A.S., 1948, A contribution to the ichthyofauna of Greenland: Spoila Zoologica Musei Hauniensis, v. 9, p. 27–57. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]

Stehmann, M., and Bürkel, D.L., 1984, Rajidae, in Whitehead, P.J.P., and others, eds., Fishes of the North-eastern Atlantic and 

the Mediterranean: Paris, Unesco, p. 163–196. [2]

Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, A., 

Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 2011, 

Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p. [5]

Bibliography

1. Jensen, A.S., 1948, A contribution to the ichthyofauna of Greenland: Spoila Zoologica Musei Hauniensis, v. 9, p. 27–57.

2. Stehmann, M., and Bürkel, D.L., 1984, Rajidae, in Whitehead, P.J.P., and others, eds., Fishes of the North-eastern Atlantic 

and the Mediterranean: Paris, Unesco, p. 163–196.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arctic-marine-fishes.

8. Essipov, V.K., 1937, On the fishes of the Polar Basin and adjacent deep waters: Problemy Arktiki, v. 4, p. 85–97. [In 

Russian.]

9. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

10. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
70 Alaska Arctic Marine Fish Ecology Catalog

Pacific Herring (Clupea pallasii)

Valenciennes, 1847

Family Clupeidae

Colloquial Name: Iñupiat: Uqsruqtuuq [1].

Ecological Role: Based on patterns of abundance, Pacific Herring 

likely are of considerable importance in the U.S. Chukchi Sea and 

of less importance in the U.S. Beaufort Sea. 

Physical Description/Attributes: Moderately compressed body 

with metallic blue-green to olive back with silvery sides and belly. 

For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 134) [2]. Swim bladder: Present 

[2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [3]. Elsewhere in Alaska, occurs in all marine waters. Worldwide, from Korea and 

Japan and the White Sea to Arctic Canada (as far north and east as Viscount Melville Sound and south and east to Bathurst Inlet 

[4]) and along the Pacific Coast south to northern Baja California [2].

Relative Abundance: Common in southeastern and northeastern Chukchi Sea [7, 8], occasionally found along much of U.S. 

Beaufort Sea [9–13]. Occasionally found in Canadian Beaufort Sea to Mackenzie River, common from Tuktoyaktuk Peninsula, 

Northwest Territories [14] to as far east as Darnley Bay in Amundsen Gulf [4].

Pacific Herring (Clupea pallasii) 217 mm TL, northeastern 

Chukchi Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution within Arctic Outer Continental Shelf Planning Areas [5] of Pacific Herring (Clupea 

pallasii), based on review of published literature and specimens from historical and recent collections [3, 6]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Pacific Herring

Clupea pallasii

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Pacific Herring 71

Depth Range: Epipelagic, coastal and offshore, from surface to 250 m, typically 150 m or less. Juveniles usually remain in 

nearshore waters from barely subtidal to at least 30 m [15–17]. Spawning occurs intertidal to at least 10 m [18, 19]. Larvae in 

Canadian Beaufort Sea were most abundant at 20 m or less [20].

Habitats and Life History

Eggs—Size: 1.2–1.8 mm when mature [21]. Time to hatching: 6–21 days [18, 22]. Habitat: Nearshore, on kelp, 

eelgrass, other plant material, and on rocks and other solid surfaces [23]. 

Larvae—Size at hatching: 5.6–7.5 mm SL [21]. Size at juvenile transformation: Metamorphosis starts at 

26 mm TL and completes by 35 mm TL [24]. Days to juvenile transformation: About 2–3 months [24]. Habitat: 

Epipelagic, in ocean currents [24]. Most abundant near surface in estuarine-influenced waters [20, 25]. 

Juveniles—Age and size: 35–150 mm TL, depending on region [24]. Habitat: Epipelagic; often found among 

kelp and eelgrass, and over soft sea floors [15, 17]. 

Adults—Age and size at first maturity: With a few exceptions, depends on water temperatures. Fish mature 

earlier in warmer waters (and lower latitudes) [24, 26]; 2 years in California, 3–5 years in eastern Bering Sea [24, 

27], and 6 years or older in Canadian Beaufort Sea [28]. Fish in California have shorter life spans and smaller 

maximum lengths than do those in the north [18]. 13–26 cm TL, depending on region [29]. Growth patterns are 

highly variable throughout the species’ geographic range as groups of fish living even tens of kilometers apart 

can grow at significantly different rates [7, 22, 24]. Maximum age: As old as 19 years [14], but rarely more than 

15 years [4, 30]. Maximum size: 46 cm TL [2]. Habitat: Epipelagic. 

Substrate—Kelp, eelgrass, other plant material, rocks and other solid surfaces for spawning [23]. 

Physical/chemical—Temperature: -1.7 °C to at least 20 °C [31–33]. Salinity: Marine and brackish waters 

[24]. Occasionally enter rivers [28, 34]. Eggs can survive between 6.1–34.2 parts per thousand [35] and 8-hour 

exposures to air twice daily [36].

Benthic and reproductive distribution for Pacific Herring (Clupea pallasii).

Open ocean

Depth, in meters

500

400

300

200

100

0 Juveniles

Clupea pallasii

Pacific Herring

Benthic distribution

Adults

Some fish to 250 m

Depth, in meters

Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Spawning and eggs

Reproductive distribution

Data from outside U.S. Beaufort-Chukchi Seas 125

100

75

50

25

0

Larvae
72 Alaska Arctic Marine Fish Ecology Catalog

Behavior

Diel—At dawn and dusk, larvae, juveniles, and adults move toward the surface to feed [24]. 

Seasonal—Spawning, over-wintering, and migration patterns are highly variable. For example, within 

Tuktoyaktuk Harbor (Beaufort Sea) fish remain for most of the year, leaving the harbor only for a few months 

during the summer to feed. [28]. Of the 10 known wintering sites in the Tuktoyaktuk Peninsula region, 8 are in 

estuarine coastal habitats, 1 is in the lower Mackenzie River, and 1 in the marine waters of Tuktoyaktuk Harbour 

[37]. At the other extreme, in the eastern Bering Sea large schools of herring winter hundreds of kilometers 

offshore (at depths of 110–130 m) and move into nearshore waters in spring to prepare for summer spawning 

[27]. Use of offshore waters as well as migrations within the U.S. Beaufort and Chukchi seas is unknown.

Elsewhere, there appears to be many migratory and non-migratory, as well as isolated and semi-isolated, 

populations throughout much of the species’ range [24, 38, 39].

Reproductive—Spawning occurs nearshore in marine and brackish waters [18, 19]. During spawning, groups 

of males emit a pheromone-like substance that triggers egg laying [40]. Females lay adhesive eggs on kelp, 

eelgrass, and other plant material, as well as on rocks and other solid surfaces [23]. Eggs are usually deposited 

in layers of one or two eggs, but when spawning runs are heavy, egg deposits may reach 5 cm thick [18]. Off 

California, spawning occurs primarily at night, but has been observed during daylight hours and over all tidal 

stages [23]. Larger and older fish tend to spawn earliest and a female spawns all of her eggs in 1 or 2 days [24]. 

Schooling—Forms schools [24]. Depending on season and location, schools of adults may be found along the 

coast and out to 1,000 km or farther offshore [27]. Schools may remain quite cohesive for extended periods 

as individuals may associate with each other for more than 200 days while moving over 185 km (100 nautical 

miles) [41]. 

Feeding—Generally, feeding is less during winter [28, 42]. Larvae, juveniles, and adults are selective pelagic 

plankton feeders [24].

Populations or Stocks

Coastal sampling and aerial surveys have provided limited information about abundance. No detailed studies 

regarding populations or stocks have been conducted.

Reproduction

Mode—Gonochoristic, oviparous, and iteroparous with external fertilization [24]. 

Spawning season—June–September in the Canadian Beaufort Sea [14, 24, 25] where spawning begins in late 

spring and early summer around the time of ice break up when waters reach at least 2.5 °C [28, 31]. Spawning 

season is highly variable throughout its range, even among groups of fish in such relatively restricted areas such 

as Puget Sound [24]. Generally, spawning occurs earliest (often in the autumn) in the more southern part of 

 the range. 

 Fecundity—Between 9,511 and 77,800 silver-gray eggs. Fecundity is highly variable and egg production at a

 particular body size is lower in high latitudes [26, 43].

Food and Feeding

Food items—Primarily zooplankton, such as mysids, euphausiids, copepods, amphipods, cumaceans, 

polychaetes, crustacean larvae, fish larvae, plant material, foraminifera, small fishes (for example, Arctic Cod, 

Fourhorn Sculpin, and Pacific Sand Lance), and fish larvae [8, 14, 30, 44–46]. 

Trophic level—3.5 [47].

Biological Interactions

Predators—Little is known. Beluga whales in spring near Barrow [48, 49]. Elsewhere, all life stages, from eggs 

to adults, are heavily preyed upon by many species of fishes, seabirds, and marine mammals [16, 50]. 

Competitors—Unknown, although likely to include various whitefishes, ciscoes, Capelin, Arctic Smelt, and 

Arctic Cod.
Pacific Herring 73

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years [51]. 

Traditional and Cultural Importance

Historically, Pacific Herring have been widely used as food as far north as the northeastern Bering Sea [52]. 

Subsistence fisheries in most of the U.S. Chukchi and Beaufort Seas are modest, although some larger catches 

are made in the Chukchi Sea [8, 45] and from the Mackenzie River eastward [4].

Commercial Fisheries

Currently, Pacific Herring are not commercially harvested. The possibility of a fishery on the north side of the 

Seward Peninsula has been suggested.

Potential Effects of Climate Change

Based on this species distributional pattern, increasing marine water temperatures will likely lead to increasing 

abundance in the U.S. Chukchi and Beaufort Seas. However, the introduction, transmission, and effects of novel 

pathogens and parasites associated with climate change elevates the risk of infection to Pacific Herring and its 

marine fish predators in the Chukchi Sea.

Areas for Future Research [A]

Pacific Herring are common in Port Clarence and Kotezebue Sound in the southeastern Chukchi Sea. Basic 

life history information and understanding of population dynamics are lacking. Improved knowledge about 

local patterns of abundance, timing and locations of reproduction, genetics, trophic linkages and energetic 

requirements, and movements and migrations are needed for stock assessments and information about their status 

and trends in time and space. Disease ecology research, including the periodic screening of Pacific Herring and 

its marine predators for the presence of infectious diseases, is recommended.

Remarks

Genetic analyses of Pacific and Atlantic Herrings imply that the ancestor of the Pacific Herring came across the Arctic from the 

Atlantic Ocean about 3 million years ago [53, 54]. 

References Cited

Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special reference 

to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 1119, 

90 p. [28]

Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p. [14]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Miller, D.J., and Schmidtke, J., 1956, Report on the distribution and abundance of Pacific herring (Clupea pallasi) along the 

coast of central and southern California: California Fish and Game, v. 42, no. 3, p. 163–187. [18]
74 Alaska Arctic Marine Fish Ecology Catalog

Stout, H.A., Gastafson, R.G., Lenarz, W.H., McCain, B.B., VanDoornik, D.M., Builder, T.L., and Methot, R.D., 2001, Status 

review of Pacific herring (Clupea pallasi) in Puget Sound, Washington: National Oceanic and Atmospheric Administration 

Technical Memorandum NMFS-NWFSC-45, 175 p. [24]

Bibliography

1. Craig, P.C., 1987, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: U.S. Department of the 

Interior, Minerals Management Service, OCS Study MMS 87-6044, Technical Report No. 129, 63 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Wolotira, R.J., Jr., Sample, T.M., and Morin, M., Jr., 1977, Demersal fish and shellfish resources of Norton Sound, the 

southeastern Chukchi Sea, and adjacent waters in the baseline year 1976: Seattle, Washington, Northwest and Alaska 

Fisheries Center, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Processed Report, 

69 p.

8. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

9. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

10. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

11. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

12. Wiswar, D.W., and Frugé, D.J., 2006, Fisheries investigations in western Camden Bay, Arctic National Wildlife Refuge, 

Alaska, 1987: Alaska Fisheries Data Series, U.S. Fish and Wildlife Service, 2006-10, 49 p.

13. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.

14. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.

15. Rosenthal, R.J., 1980, Shallow water fish assemblages in the northeastern Gulf of Alaska—Habitat evaluation, species 

composition, abundance, spatial distribution and trophic interaction, in Bureau of Land Management, Environmental 

assessment of the Alaskan Continental Shelf, final reports of principal investigators: U.S. Department of Commerce, 

National Oceanic and Atmospheric Administration, p. 451–540.
Pacific Herring 75

16. Hay, D.E., Healey, M.C., Ware, D.M., and Wilimovsky, N.J., 1992, Distribution, abundance, and habitat of prey fish on the 

west coast of Vancouver Island, in Vermeer, K., Butler, R.W., and Morgan, K.H., eds., The ecology, status, and conservation 

of marine and shoreline birds on the west coast of Vancouver Island: Canadian Wildlife Service, Occasional Paper, no. 75, 

p. 22–50.

17. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

18. Miller, D.J., and Schmidtke, J., 1956, Report on the distribution and abundance of Pacific herring (Clupea pallasi) along the 

coast of central and southern California: California Fish and Game, v. 42, no. 3, p. 163–187.

19. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

20. Sareault, J., 2009, Marine larval fish assemblages in the nearshore Canadian Beaufort Sea during July and August: 

Winnipeg, University of Manitoba, Master’s thesis, 146 p.

21. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

22. Wespestad, V.G., and Barton, L.H., 1981, Distribution, migrations, and status of Pacific herring, in Hood, D.W., and 

Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and Resources: National Oceanic and Atmospheric 

Administration, p. 509–524.

23. Eldridge, M.B., and Kaill, W.M., 1973, San Francisco Bay area’s herring resource—A colorful past and a controversial 

future: Marine Fisheries Review, v. 35, no. 11, p. 25–31.

24. Stout, H.A., Gastafson, R.G., Lenarz, W.H., McCain, B.B., VanDoornik, D.M., Builder, T.L., and Methot, R.D., 2001, Status 

review of Pacific herring (Clupea pallasi) in Puget Sound, Washington: National Oceanic and Atmospheric Administration 

Technical Memorandum NMFS-NWFSC-45, 175 p.

25. Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbour, N.W.T.: Canadian Technical 

Report of Fisheries and Aquatic Sciences, no. 1218, 21 p.

26. Hay, D.E., 1985, Reproductive biology of Pacific herring (Clupea harengus pallasi): Canadian Journal of Fisheries and 

Aquatic Sciences, v. 42, suppl. 1, p. 111–126.

27. Dudnik, Y.I., and Usol’tsev, E.A., 1964, The herrings of the eastern part of the Bering Sea, Moiseev, P.A., ed., in Soviet 

fisheries investigations in the Northeastern Pacific, part 1: Jerusalem, Israel Program for Scientific Translations [1968], 

p. 236–240.

28. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 

1119, 90 p.

29. Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and invertebrates 

in west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments 

Division, ELMR Report no. 8, 327 p.

30. Barton, L.H., 1978, Finfish resource surveys in Norton Sound and Kotzebue Sound: Alaska Department of Fish and Game, 

Commercial Fisheries Division, p. 75–313.

31. Gillman, D.V., and Kristofferson, A.H., 1984, Biological data on Pacific herring (Clupea harengus pallasi) from 

Tuktoyaktuk Harbour and the Liverpoor Bay area, Northwest Territories, 1981 to 1983: Winnipeg, Manitoba, Canada, 

Canadian Data Report of Fisheries and Aquatic Sciences, Department of Fisheries and Oceans, Western Region, no. 485, 

22 p.
76 Alaska Arctic Marine Fish Ecology Catalog

32. Grosse, D.J., and Hay, D.E., 1988, Pacific herring, Clupea harengus pallasi, in the northeast Pacific and Bering Sea, in

Wilimovsky, N.J., Incze, L.S., and Westrheim, S.J., eds., Species synopses—Life histories of selected fish and shellfish 

of the northeast Pacific and Bering Sea: Seattle, University of Washington, Washington Sea Grant Program and Fisheries 

Research Institute, p. 34–54.

33. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

34. McAllister, D.E., 1959, Records of marine fishes from fresh water in British Columbia: The Canadian Field-Naturalist, 

v. 73, no. 1, p. 13–14.

35. McMynn, R.G., and Hoar, W.S., 1953, Effects of salinity on the development of the Pacific herring: Canadian Journal of 

Zoology, v. 31, no. 4, p. 417–432.

36. Jones, B.C., 1972, Effect of intertidal exposure on survival and embryonic development of Pacific herring spawn: Journal of 

the Fisheries Research Board of Canada, v. 29, no. 8, p. 1,119–1,124.

37. Sekerak, A.D., Stallard, N., and Griffiths, W.B., 1992, Distribution of fish and fish harvests in the nearshore Beaufort Sea 

and Mackenzie Delta during ice-covered periods, October–June: Environmental Studies Research Funds Report, LGS Ltd. 

No. 117, 524 p.

38. Moser, M., and Hsieh, J., 1992, Biological tags for stock separation in Pacific herring Clupea harengus pallasi in California: 

Journal of Parasitology, v. 78, no. 1, p. 54–60.

39. O’Connell, M., Dillon, M.C., Wright, J.M., Bentzen, P., Merkouris, S.E., and Seeb, J.E., 1998, Genetic structuring among 

Alaskan Pacific herring populations identified using microsatellite variation: Journal of Fish Biology, v. 53, no. 1, 

p. 150–163.

40. Sherwood, N.M., Kyle, A.L., Kreiberg, H., Warby, C.M., Magnus, T.H., Carolsfeld, J., and others, 1991, Partial 

characterization of a spawning pheromone in the herring Clupea harengus pallasi: Canadian Journal of Zoology, v. 69, 

no. 1, p. 91–103.

41. Hay, D.E., and McKinnell, S.M., 2002, Tagging along—Association among individual Pacific herring (Clupea pallasi) 

revealed by tagging: Canadian Journal of Fisheries and Aquatic Sciences, v. 59, no. 12, p. 1,960–1,968.

42. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British, Beaufort Sea Project, Beaufort Sea Technical 

Report, no. 8, 114 p.

43. Paulson, A.C., and Smith, R.L., 1977, Latitudinal variation of Pacific herring fecundity: Transactions of the American 

Fisheries Society, v. 106, no. 3, p. 244–247.

44. Jones, M.L., and Den Beste, J., 1977, Tuft Point and adjacent coastal areas fisheries projects: Calgary, Alberta, Canada, 

Aquatic Environments, Ltd., 152 p.

45. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

46. Lacho, G., 1991, Stomach content analyses of fishes from Tuktoyaktuk Harbour, N.W.T., 1981: Winnipeg, Manitoba, 

Canadian Data Report of Fisheries and Aquatic Sciences, Central and Arctic Region, Department of Fisheries and Oceans, 

no. 853, 15 p.

47. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320. 

48. Seaman, G.A., Lowry, L.F., and Frost, K.J., 1982, Food of beluga whales (Delphinapterus leucas) in western Alaska: 

Cetology, v. 44, p. 1–19.

49. Lowry, L.F., Frost, K.J., and Seaman, G.A., 1986, Investigations of belukha whales in coastal waters of western and 

northern Alaska: Outer Continental Shelf Environmental Program Unit 612, Final Report, p. 359–392.

50. Gerke, B.L., 2002, Spawning habitat characteristics of Pacific herring (Clupea pallasii) in Prince William Sound, Alaska: 

Fairbanks, University of Alaska, Master’s thesis, 112 p.
Pacific Herring 77

51. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

52. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

53. Grant, W.S., 1986, Biochemical genetic divergence between Atlantic, Clupea harengus, and Pacific, C. pallasi, herring: 

Lawrence, Kansas, Copeia, no. 3, p. 714–719.

54. Domanico, M.J., Phillips, R.B., and Schweigert, J.F., 1996, Sequence variation in ribosomal DNA of Pacific (Clupea 

pallasi) and Atlantic herring (Clupea harengus): Canadian Journal of Fisheries and Aquatic Sciences, v. 53, no. 11, 

p. 2,418–2,423.
78 Alaska Arctic Marine Fish Ecology Catalog

Pond Smelt (Hypomesus olidus)

(Pallas, 1814)

Family Osmeridae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort 

Seas. Called “Cigarfish” around Nome and other areas of Norton 

Sound [1].

Ecological Role: The rare occurrence of Pond Smelt in brackish and 

marine waters of the U.S. Chukchi Sea implies a minor ecological 

role in other than freshwater habitats.

Physical Description/Attributes: Grey- or olive-green to yellow-brown dorsally becoming silvery white on belly. Snout 

and operculum are covered with black mottles or spots [2, 3]. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 172) [3]. Swim bladder: Present, physostomous [4]. Antifreeze glycoproteins in blood serum: 

Unknown.

Range: U.S. Chukchi Sea. In Alaska, in drainages northwards from the Copper River, northeastern Gulf of Alaska, to the Kobuk 

River (draining into the Chukchi Sea). Worldwide, from North Korea and Japan to northern Siberia and east through drainages 

of Arctic Canada to Coronation Gulf, Northwest Territories, Canada [3].

Relative Abundance: Absent or rare in coastal waters of the U.S. Chukchi and Beaufort Seas. Elsewhere, common at least as 

far north as Port Clarence, northeastern Bering Sea [1], wherePond Smelt is occasionally found well offshore [6]. Common in 

fresh water and occasional in coastal, brackish conditions in Mackenzie Delta region [8–10]. 

Pond Smelt (Hypomesus olidus) 114 mm, northeastern 

Bering Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Pond Smelt 

Hypomesus olidus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Pond Smelt (Hypomesus olidus) within Arctic Outer Continental Shelf Planning 

Areas [5] based on review of published literature and specimens from historical and recent collections [6, 7]. 
Pond Smelt 79

Depth Range: Nearshore, shallow waters, typically less than 5 m [1, 11]. Taken offshore of Cape Rodney and Sledge Island 

(northeastern Bering Sea) in 2007 by surface trawl fishing to depth of 29 m [6].

324PdSlt

Open ocean Shore

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

Adults and juveniles 

Some fish to 29 m

Hypomesus olidus

Pond Smelt

Benthic distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Depth, in meters

Reproductive distribution

No reproductive distribution in seawater

Anadromous.

Spawning, eggs and larvae

all occur in freshwater

Benthic and reproductive distribution of Pond Smelt (Hypomesus olidus).

Habitats and Life History

Many populations are anadromous, although some stocks are landlocked [3]. 

Eggs—Size: 0.9 mm [12]. Time to hatching: 10–38 days at 5.0–15.0 °C [12, 13]. Habitat: Shallow depths of 

lakes and rivers, on submerged vegetation or rocks [12–14]. 

Larvae—Size at hatching: 4.6 mm long [12, 13]. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic, in freshwater rivers and lakes [12–15]. 

Juveniles—Age and size: As small as 24 mm FL [9, 12, 16]. Habitat: Pelagic in coastal marine and estuarine 

waters, and rivers and lakes [3]. Remain in their natal habitats 1 to 1 year before migrating to coastal waters [9, 

12, 16]. 

Adults—Age and size at first maturity: 1–4 years for anadromous fish [2, 8, 12, 17–19]. In southwestern Bering 

Sea drainages, anadromous fish mature at age-3, whereas non-anadromous type matures at age-1 and age-2 [20]. 

Size is about 10.0 cm FL or more in Asia [2, 12–14]. In the Sea of Okhotsk, females are slightly larger at age 

than males [2]. Fish living in the Sea of Okhotsk grow faster than those in the Mackenzie Delta or a landlocked 

Yukon Lake population [2, 12–14]. Maximum age: About 6 years for anadromous fish in Asia, though few 

survive to that age [12, 20]. Maximum size: 20 cm TL [3]. Habitat: Pelagic, in coastal marine and estuarine 

waters, rivers and lakes [2, 3, 10, 12, 13, 17, 21]. 

Substrate—Taken over sand-gravel in Bristol Bay [22]. 

Physical/chemical—Temperature: As warm as 17 °C [20]. Salinity: Mainly freshwater, occasionally enters 

brackish river deltas and nearshore marine waters [3, 7].

Behavior

Diel—Unknown. Unidentified osmerid larvae in Auke Bay (southeastern Alaska) migrated to surface waters at 

midnight [23]. 

Seasonal—Large downstream migrations to Tuktoyaktuk Harbor occur August and September [9]. Migrations 

upstream may begin while the rivers are still under ice and be as long as 70 km (44 mi) [12]. 

Reproductive—Spawning occurs in rivers and lakes. Some populations in Asia ascend rivers from coastal 

waters in spring, just before spawning, whereas others migrate into fresh waters in autumn and overwinter prior 

to spawning [17]. Spawning takes place at dusk. Eggs are laid on submerged vegetation or rocks in shallow, 

swift-flowing, waters [12–14]. In many, but not all populations, fish die after spawning [10, 12, 17, 19]. 

Surviving fish migrate downstream shortly after spawning [12]. 

Schooling—Forms schools [13]. 

Feeding—Some populations do not feed during spawning season [20] although this is not a universal 

behavior [12].
80 Alaska Arctic Marine Fish Ecology Catalog

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [15]. 

Spawning season—Spawning in North America takes place at least during May–July [10, 19] and as early as 

April in Asia [13]. 

Fecundity—4,820–33,010 adhesive egg, spawned in a single batch (around Sakhalin Island, Russia) [12]. 

Food and Feeding

Food items—Primarily midwater crustaceans (for example, mysids, copepods, amphipods, and isopods), insects, 

snails, and small fishes [10, 18, 20, 24, 25]. 

Trophic level—3.21 (standard error 0.42) [11].

Biological Interactions

Predators—Beluga whales during May and June in Bristol Bay [26]. Inconnu and Northern Pike in North 

American Arctic fresh waters [10]. 

Competitors—Potentially midwater planktivores such as Arctic Cod, Pacific Herring, and Capelin, and other 

coastal fishes.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (t

m=2; tmax=5) [11].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Pond Smelt are not commercially harvested.

Potential Effects of Climate Change

Unclear. It is possible that warming Arctic waters will lead to increased abundance of this species as brackish 

habitats expand. However, it is unknown whether Arctic streams will become suitable spawning habitats for 

successful colonization.
Pond Smelt 81

Areas for Future Research [B]

Little is known about the ecology and life history of this species in the U.S. Chukchi and Beaufort Seas. 

Research needs include: (1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-theyear benthic recruitment; (3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal 

and ontogenetic movements; (6) population studies; (7) prey; and (8) predators.

References Cited

Chereshnev, I., Shestakov, A.V., and Skopets, M.B., 1999, On the distribution of silver smelts of the genus Hypomesus

(Osmeridae) in the northern part of the Sea of Okhotsk: Journal of Ichthyology, v. 39, no. 7, p. 498–503. [2]

DeGraaf, D.A., 1986, Aspects of the life history of the pond smelt (Hypomesus olidus) in the Yukon and Northwest Territories: 

Arctic, v. 39, no. 3, p. 260–263. [10]

Gritsenko, O.F., Churikov, A.A., and Rodionova, S.S., 1984a, The ecology of the pond smelt, Hypomesus olidus (Osmeridae), on 

Sakhalin: Journal of Ichthyology, v. 24, no. 4, p. 91–100. [12]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries investigations 

in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224. [13]

Bibliography

1. Barton, L.H., 1978, Finfish resource surveys in Norton Sound and Kotzebue Sound: Alaska Department of Fish and Game, 

Commercial Fisheries Division, p. 75–313.

2. Chereshnev, I., Shestakov, A.V., and Skopets, M.B., 1999, On the distribution of silver smelts of the genus Hypomesus

(Osmeridae) in the northern part of the Sea of Okhotsk: Journal of Ichthyology, v. 39, no. 7, p. 498–503. 

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Jørgensen, R., 2003, The effects of swimbladder size, condition and gonads on the acoustic target strength of mature 

capelin: ICES Journal of Marine Science, v. 60, no. 5, p. 1,056–1,062.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

8. DeGraaf, D.A., and Machniak, K., 1977, Fisheries investigations along the cross delta pipeline route in the MacKenzie 

Delta, in McCart, P.J., ed., Studies to determine the impact of gas pipeline development on aquatic ecosystems: Arctic Gas, 

Biological Report Series 39, p. 1–169.

9. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 

1119, 90 p.

10. DeGraaf, D.A., 1986, Aspects of the life history of the pond smelt (Hypomesus olidus) in the Yukon and Northwest 

Territories: Arctic, v. 39, no. 3, p. 260–263.
82 Alaska Arctic Marine Fish Ecology Catalog

11. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

12. Gritsenko, O.F., Churikov, A.A., and Rodionova, S.S., 1984a, The ecology of the pond smelt, Hypomesus olidus 

(Osmeridae), on Sakhalin: Journal of Ichthyology, v. 24, no. 4, p. 91–100.

13. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224.

14. Berg, L.S., 1948, Freshwater fishes of the USSR and adjacent countries, volume 1 (4th ed.): Moscow, Academy of Sciences 

of the USSR Zoological Institute, 466 p. [Translated from Russian by Israel Program for Science Translation, Jerusalem, 

505 p.]

15. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

16. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.

17. Hamada, K., 1961, Taxonomic and ecological studies of the genus Hypomesus of Japan: Hokkaido University, Memoirs of 

the Faculty of Fisheries, v. 9, no. 1, p. 1–55.

18. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

19. Harvey, C.J., Ruggerone, G.T., and Rogers, D.E., 1997, Migrations of three-spined stickleback, nine-spined stickleback, and 

pond smelt in the Chignik catchment, Alaska: Journal of Fish Biology, v. 50, no. 5, p. 1,133–1,137.

20. Karpenko, V.I., and Vasilets, P.M., 1996, Biology of smelt (Osmeridae) in the Korf-Karagin coastal area of the southwestern 

Bering Sea, in Mathisen, O.A., and Coyle, K.O., eds., Ecology of the Bering Sea—A review of Russian literature: 

Fairbanks, Alaska, University of Alaska, Alaska Sea Grant Program, AK-SG-96-01, p. 217–235.

21. Reshetnikov, Y.S., Bogutskaya, N.G., Vasil’eva, E.D., Dorofeeva, E.A., Naseka, A.M., Popova, O.A., and others, 1997, An 

annotated check-list of the freshwater fishes of Russia: Journal of Ichthyology, v. 37, no. 9, p. 687–736.

22. Johnson, S.W., Thedinga, J.F., and Lindeberg, M.R., 2012, Nearshore fish atlas of Alaska: National Oceanic and 

Atmospheric Administration Fisheries, accessed February 2012 at http://www.fakr.noaa.gov/habitat/fishatlas/.

23. Haldorson, L.J., Prichett, M., Paul, A.J., and Ziemann, D., 1993, Vertical distribution and migration of fish larvae in a 

northeast Pacific bay: Marine Ecology Progress Series, v. 101, p. 67–80.

24. Martin, D.J., Glass, D.R., Whitmus, C.J., Simenstad, C.A., Milward, D.A., Volk, E.C., Stevenson, M.L., Nunes, P., Savoie, 

M., and Grotefendt, R.A., 1986, Distribution, seasonal abundance, and feeding dependencies of juvenile salmon and 

non-salmonid fishes in the Yukon River Delta: Outer Continental Shelf Environmental Assessment Program, Reports of 

Principal Investigators, U.S. Department of Commerce and U.S. Department of the Interior, 388 p.

25. Maksimenkov, V.V., and Tokranov, A.M., 1993, The diet of the pond smelt, Hypomesus olidus, in the Bol’shaya River 

Estuary (Western Kamchatka): Journal of Ichthyology, v. 33, no. 9, p. 11–21.

26. Lowry, L.F., Frost, K.J., and Seaman, G.A., 1986, Investigations of belukha whales in coastal waters of western and 

northern Alaska: Outer Continental Shelf Environmental Program Unit 612, Final Report, p. 359–392.
Pacific Capelin 83

Pacific Capelin (Mallotus catervarius)

(Pennant, 1784)

Family Osmeridae 

Note: Until recently believed to be a junior synonym of Mallotus 

villosus (Müller, 1776). However, molecular genetic studies 

demonstrate a substantial genetic distance between this species and 

other Arctic mallotus spp. clades [73].

Colloquial Name: Iñupiaq: Panmagriq, Panmaksraq, Pagmaksraq

[1, 2].

Ecological Role: The true abundance of Pacific Capelin is probably 

underestimated in existing survey data, but this species is hypothesized to be a major prey of many fish, birds, and marine 

mammals in the U.S. Chukchi and Beaufort Seas. Although its forage fish status is uncertain, its life history cycle suggests 

an important biological linkage between nearshore and offshore habitats especially in coastal waters influenced by large river 

deltas. It is a wide ranging, high lipid, cold-water fish that is an important part in Arctic and Subarctic food webs.

Physical Description: Elongate and narrow with a blueish, greenish, or yellowish back and silvery sides and belly. For 

specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 171) [3]. Swim bladder: Present [4]. 

Antifreeze glycoproteins in blood serum: Unknown, absent from Mallotus villosus in the Barents Sea [5].

Range: U.S. Chukchi and Beaufort Seas [3]. Elsewhere, Seas of Japan and Okhotsk, Commander and Aleutian Islands, Gulf of 

Alaska to Strait of Juan de Fuca eastwards to at least Davis Strait and southern end of Baffin Island, eastern Canada. Presence in 

Siberian Seas unclear [8].

Relative Abundance: Common, patchily distributed, in U.S. Chukchi and Beaufort seas at least as far east as about Camden 

Bay [9–14].

Pacific Capelin (Mallotus catervarius) 84 mm TL, Semidi 

Islands, western Gulf of Alaska, 2001. Photograph by 

C.W. Mecklenburg, Point Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Pacific Capelin 

Mallotus catervarius

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Pacific Capelin (Mallotus catervarius) within Arctic Outer Continental Shelf Planning 

Areas [7] based on review of published literature and specimens from historical and recent collections [3, 8].
84 Alaska Arctic Marine Fish Ecology Catalog

Depth Distribution: Surface to 200 m [8]. In western U.S. Beaufort Sea, common in intertidal and barely subtidal waters and to 

at least 8 m [14]. In Prince William Sound and the Gulf of Alaska, most abundant in upper 100 m of water column [16]. Reports 

to 725 cm [17] are likely fish caught in trawls much nearer the surface. Larvae are found near the surface [18]. Juveniles are 

reported in very shallow nearshore waters [11, 14, 19]. Spawning occurs in very shallow waters barely subtidal waters [13, 20].

Benthic and reproductive distribution of Pacific Capelin (Mallotus catervarius).

Some fish to 300 m

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Young

juveniles

in study area

Adults and

older juveniles

Mallotus catervarius

Pacific Capelin

Benthic distribution

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Spawning

and eggs 

Spawning

in study area

Larvae and overall spawning data 

from outside Beaufort-Chukchi Seas

 Maximum depth data from

outside U.S. Beaufort-Chukchi Seas

Reproductive distribution

Larvae

Habitats and Life History

Eggs—Time to hatching: Unknown. Time to hatching: Unknown, but in Mallotus villosus, as much as 80 days at 

2 °C, 30 days at 5 °C, and 15 days at 10 °C [22]. Size: Unknown. Once laid, eggs can survive as long as 6 hours 

at temperatures as low as -5 °C [25]. Habitat: Spawning substrate has not been defined. Demersal [26] or buried, 

usually in coarse sand and fine gravel [27, 28]. Occasionally in fine sand [26]. 

Larvae—Size at hatching: About 4 mm [31]. Size at juvenile transformation: 60 mm at start [31]. Larvae are 

found near the surface [18]. After hatching, some appear to remain in substrate for several days [33]. 

Juveniles—Age: Unknown. Size: 75.0–80.0 mm SL [31]. Habitat: Poorly understood. Young-of-the-year live 

from very shallow nearshore waters out to at least 15 km from shore [11, 14, 19]. 

Adults—Age and size at first maturity: Little is known. At Point Lay, U.S. Chukchi Sea, almost all spawning 

fish were 2-year fish with a very small percentage of 3-year fish, and ranged in size from 110 to 155 mm FL [9]. 

Bering Sea fish mature at 2 years [35]. Maximum age: In Canadian Beaufort Sea, at least 5 years [36]. Maximum 

size: Fish in the U.S. Chukchi and Beaufort Seas do not appear to grow much larger than about 160 mm [1, 

9, 13, 36, 37]. Northern Pacific 21.8 cm [74]. Habitat: Poorly understood. Older fish are taken in nearshore 

waters during the spawning season [11, 14, 19]. In a 3-year beach seine study conducted west of Barrow, Pacific 

Capelin were most abundant during the coldest-water year [14]. Their location in winter is unknown. In Bering 

Sea, Pacific Capelin live as much as 560 km from shore, but only where the continental shelf is shallow and 

broad [35].

Physical/chemical—Temperature: Tolerate waters as cold as -2.0 to -1.8 °C and as warm as 14 °C for brief 

periods, but optimal temperatures are about -1.0–6.0 °C [16, 35, 38]. Salinity: Generally, marine and brackish 

waters, but may on occasion enter rivers [41]. 

Behavior

Capelin behavior is poorly understood in U.S. Chukchi and Beaufort Seas. 

Diel—Unknown. Osmerid larvae in southeastern Alaska migrated to the surface at night [42]. 

Seasonal—Unknown. Some Capelin aggregations make extensive migrations to offshore feeding sites [35] 

where single sex schools are formed prior to migrating to spawning grounds [26]. 

Reproductive—Poorly known. Larger fish spawn earlier and males usually reach spawning grounds first [26]. 

Most spawning takes place in marine waters although some occurs in brackish conditions [26] and in very 

shallow, barely subtidal waters [13, 20]. However, there is some evidence that spawning in eastern Bering Sea 
Pacific Capelin 85

and perhaps U.S. Chukchi and Beaufort Seas also may occur somewhat deeper [11, 46], although the maximum 

spawning depth is not known. In eastern Bering Sea and Gulf of Alaska, there is a tendency for spawning to 

occur or at least begin at night and around the highest tides. However, spawning can begin at any time of the day 

or night and has been known to continue over several days [26]. 

Schooling—Capelin school in U.S. Chukchi and Beaufort Seas, but the extent of schools is unknown. In the Gulf 

of Alaska, schools may be more than 1 km long and 20 m or more thick, and aggregations of schools may extend 

to 10 km [47]. 

Feeding—In the southeastern Bering Sea, Capelin feed most heavily in the afternoon and rarely at night [48]. 

Studies in the Chukchi and Barents Seas, North Atlantic Ocean, and off Kamchatka Peninsula, Russia, imply that 

fish feed heavily before and after the spawning season [9, 22, 49]. In the western Gulf of Alaska, fish to 126 mm 

were crepuscular feeders and fish in the Canadian Atlantic switch to diurnal feeding during winter [50].

Population or Stocks

Fish in U.S. Chukchi and Beaufort Seas may form a population that includes Bering Sea and western Pacific 

Ocean fish, but not fish from the Gulf of Alaska or Atlantic Ocean [51].

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external. 

Spawning season—In the U.S. Chukchi and Beaufort Seas, spawning is primarily in July and August [9, 19, 36, 

52], although some may take place in June [53] and early September [54]. 

Fecundity—Unknown. Females release all of their eggs at one time and produce between 5,000 and 22,000 eggs 

[26]. Although not studied in the study area, in other locations most males die after the spawning season [26]. In 

some populations, substantial numbers of females may survive to spawn in the following year [56].

Food and Feeding

Food items—Food habits of larvae unknown. Capelin feed on midwater crustaceans, fish larvae, and other 

planktonic organisms. Limited surveys in the Chukchi and Beaufort Seas have indicated that mysids are the most 

important prey, although calanoid and harpacticoid copepods, euphausiids, amphipods, crustacean larvae, and 

fish eggs and larvae also are consumed [1, 9, 36]. 

Trophic level—3.5 [60].

Biological Interactions

Predators—Besides the seabirds found at Capes Lisburne and Thompson, Capelin are rarely reported in food 

habit studies in the U.S. Chukchi and Beaufort Seas. Ringed seals have eaten Capelin during the winter in the 

U.S. Chukchi Sea [61]. In the Bering Sea, Gulf of Alaska, and eastern Canadian Arctic and northern Atlantic, this 

species is extremely important as food for a very wide range of marine mammals, seabirds, and fishes [63–67]. 

Competitors—Presumably a wide range of water-column, zooplankton feeders, including Arctic Cod and 

Walleye Pollock. 

1980 1990 2000 2010

Resilience

Unknown for this species, but estimated for Mallotus villosus as medium, minimum population doubling time is 

1.4–4.4 years (K=0.3–0.5; t

m=3; tmax=10; Fecundity=6,000) [68].

Traditional and Cultural Importance

Moderate importance in subsistence fisheries. Most fish are taken during spawning runs [2, 69–71].
86 Alaska Arctic Marine Fish Ecology Catalog

Commercial Fisheries

Currently, Pacific Capelin are not commercially harvested.

Potential Effects of Climate Change

Unclear for this species. However, Mallotus villosus have the capacity to respond quickly to climate change [for 

example, water temperature and food availability [72]. 

Areas for Future Research [A]

Although commonly sampled in coastal habitats, very little information exists on the life history of Pacific 

Capelin, particularly in U.S. Chukchi and Beaufort Seas. Because of this, many aspects of the biology of this 

species were inferred from other regions. It is a major forage species elsewhere in the Arctic and in other parts 

of Alaska. The phenology of the species in nearshore waters is brief and linked to reproduction and nursery. 

Early life history stages are likely swept offshore in wind-driven currents and thus the forage significance of the 

species in more poorly studied offshore habitats is not well documented. In particular, although it is clear that 

Pacific Capelin live and spawn (that is, beach versus ocean spawners) in this region, often in large numbers, 

there is a paucity of data on their basic biology, seasonality of their movements and behaviors, and locations of 

overwintering grounds. The basal metabolic and growth rates of Pacific Capelin living in the U.S. Chukchi and 

Beaufort Seas indicate adaptations to cold-water marine environments. The effects of warming temperatures on 

these physiological processes should be determined in laboratory experiments.

References Cited

Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of Alaska, v. 

24, p. 27–54. [2]

Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research Associates, 

Inc., North Slope Borough, Materials Source Division, 105 p. [1]

Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p. [9]

Hart, J.J., 1973, Pacific fishes of Canada: Ottawa, Fisheries Research Board of Canada, Bulletin 180 [74]. 

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1.116 p. [3]

Pahlke, K.A., 1985, Preliminary studies of capelin (Mallotus villosus) in Alaskan waters: Alaska Department of Fish and Game, 

Informational Leaflet No. 250, 64 p. [26]

Thorsteinson, L.K., Jarvela, L.E., and Hale, D.A., 1990, Arctic fish habitat use investigations—Nearshore studies in the Alaskan 

Beaufort Sea, summer 1988: Final Report, Alaska Office, Ocean Assessments Division, National Oceanic and Atmospheric 

Administration, Research Unit 682, 125 p. [11]

Bibliography

1. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

2. Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.
Pacific Capelin 87

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Jørgensen, R., 2003, The effects of swimbladder size, condition and gonads on the acoustic target strength of mature 

capelin: ICES Journal of Marine Science, v. 60, no. 5, p. 1,056–1,062.

5. Raymond, J.A., and Hassel, A., 2000, Some characteristics of freezing avoidance in two osmerids, rainbow smelt and 

capelin: Journal of Fish Biology, v. 57, suppl. A, p. 1–7.

6. Hay, D.E., 1998, Historic changes in capelin and eulachon populations in the Strait of Georgia, in Pauly, D., Pitcher, 

T.J., Preikshot, David, Hearne, J., eds., Back to the future—Reconstructing the Strait of Georgia ecosystem: Vancouver, 

University of British Columbia, The Fisheries Center Research Reports, p. 42–44.

7. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

8. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

9. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

10. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

11. Thorsteinson, L.K., Jarvela, L.E., and Hale, D.A., 1990, Arctic fish habitat use investigations—Nearshore studies in the 

Alaskan Beaufort Sea, summer 1988: Final Report, Alaska Office, Ocean Assessments Division, National Oceanic and 

Atmospheric Administration, Research Unit 682, 125 p.

12. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

13. Wiswar, D.W., and Frugé, D.J., 2006, Fisheries investigations in western Camden Bay, Arctic National Wildlife Refuge, 

Alaska, 1987: Alaska Fisheries Data Series, U.S. Fish and Wildlife Service, 2006-10, 49 p.

14. Johnson, S.W., Thedinga, J.F., Neff, A.D., and Hoffman, C.A., 2010, Fish fauna in nearshore waters of a Barrier Island 

in the western Beaufort Sea, Alaska: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, 

Technical Memorandum, NMFSAFSC-210, 28 p.

15. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

16. Brown, E.D., 2002, Life history, distribution, and size structure of Pacific capelin in Prince William Sound and the northern 

Gulf of Alaska: ICES Journal of Marine Science, v. 59, p. 983–996.

17. Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic marine fishes of Canada: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, no. 2674, 112 p.

18. Jump, C.M., Duffy-Anderson, J.T., and Mier, K.L., 2008, Comparison of the Sameoto, Manta and MARMAP neustonic 

ichthyoplankton samplers in the Gulf of Alaska: Fisheries Research, v. 89, no. 3, p. 222–229.

19. Houghton, J.P., and Whitmus, C.J., 1988, Shallow neritic fish of the central Beaufort Sea: Seattle, Washington, Report 

prepared for Standard Alaska Production Company by Dames and Moore, 17298-002-20.

20. Bendock, T.N., 1977, Beaufort Sea estuarine fishery study: Alaska Department of Fish and Game Annual Report, Contract 

#03-5-022-69, p. 670–729.

21. Huse, G., 1998, Sex-specific life history strategies in capelin (Mallotus villosus): Canadian Journal of Fisheries and Aquatic 

Sciences, v. 55, no. 3, p. 631–638.
88 Alaska Arctic Marine Fish Ecology Catalog

22. Gjøsaeter, H., 1998, The population biology and exploitation of capelin (Mallotus villosus) in the Barents Sea: Sarsia, v. 83, 

no. 6, p. 453–496.

23. Nakashima, B.S., and Taggart, C.T., 2002, Is beach-spawning success for capelin, Mallotus villosus (Müller), a function of 

the beach?: ICES Journal of Marine Science, v. 59, no. 5, p. 897–908.

24. Grégoire, F., Morneau, R., Caron, G., Beaudoin, M., Lévesque, C., Rose, C., and others, 2004, Fécondité du capelan 

(Mallotus villosus) dans l-estuaire de le golfe du Saint-Laurent en 2003: Rapport Technique Canadien Dessciences 

Halieutiques et Aquatiques, Ministère des Pêches et des Océans, Mont-Joli, Québec, no. 2560, 30 p.

25. Power, G., 1997, A review of fish ecology in Arctic North America: American Fisheries Society Symposium, no. 19, 

p. 13–39.

26. Pahlke, K.A., 1985, Preliminary studies of capelin (Mallotus villosus) in Alaskan waters: Alaska Department of Fish and 

Game, Informational Leaflet No. 250, 64 p.

27. Hart, J.L., and McHugh, J.L., 1944, The smelts (Osmeridae) of British Columbia: Fisheries Research Board of Canada, 

v. 64, 27 p.

28. Velikanov, A.Y., 1984, Ecology of reproduction of the fareastern capelin, Mallotus villosus socialis (Osmeridae), along the 

coasts of Sakhalin Island: Journal of Ichthyology, v. 24, no. 3, p. 43–48.

29. Davoren, G.K., Anderson, J.T., and Montevecchi, W.A., 2006, Shoal behaviour and maturity relations of spawning capelin 

(Mallotus villosus) off Newfoundland—Demersal spawning and diel vertical movement patterns: Canadian Journal of 

Fisheries and Aquatic Sciences, v. 63, no. 2, p. 268–284.

30. Jangaard, P.M., 1974, The capelin (Mallotus villosus): Fisheries Research Board of Canada, Bulletin no. 186, p. 1–70.

31. Doyle, M.J., Busby, M.S., Duffy-Anderson, J.T., Picquelle, S.J., and Matarese, A.C., 2002, Aspects of the early life history 

of the capelin (Mallotus villosus) in the northeast Gulf of Alaska—A historical perspective based on larval collections 

October 1977-March 1979: National Oceanic and Atmospheric Administration Technical Memorandum, NMFS-AFSC-132, 

41 p.

32. Collette, B.B., and Klein-MacPhee, G., 2002, Bigelow and Schroeder’s fishes of the Gulf of Maine (3rd ed.): Washington, 

D.C., Smithsonian Institution Press, 882 p.

33. Velikanov, A.Y., 1988, Data on the eggs and larvae of the Pacific capelin (Mallotus villosus socialis) along the shores of 

southern Sakhalin: Journal of Ichthyology, v. 28, p. 86–91.

34. Ochman, S., and Dodson, J.J., 1982, Composition and structure of the larval and juvenile fish community of the Eastman 

River and estuary, James Bay: Naturaliste Canada, v. 109, p. 803–813.

35. Naumenko, E.A., 1996, Distribution, biological condition, and abundance of capelin (Mallotus villosus socialis) in the 

Bering Sea, in Mathisen, O.A., and Coyle, K.O., eds., Ecology of the Bering Sea: Fairbanks, Alaska, University of Alaska, 

Alaska Sea Grant College Program, p. 237–256.

36. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.

37. Wiswar, D.W., West, R.L., and Winkleman, W.N., 1995, Fisheries investigation in Oruktalik Lagoon, Arctic National 

Wildlife Refuge, Alaska, 1986: U.S. Fish and Wildlife Service, Alaska Fisheries Technical Report, no. 30, 38 p. 

38. Moulton, L.L., and Tarbox, K.E., 1987, Analysis of Arctic cod movements in the Beaufort Sea nearshore region, 1978–79: 

Arctic, v. 40, no. 1, p. 43–49.

39. Rose, G.A., 2005, Capelin (Mallotus villosus) distribution and climate—A sea “canary” for marine ecosystem change: ICES 

Journal of Marine Science, v. 62, no. 7, p. 1,524–1,530.

40. Stergiou, K.I., 1989, Capelin Mallotus villosus (Pisces: Osmeridae), glaciations, and speciation—A nomothetic approach to 

fisheries ecology and reproductive biology: Marine Ecology Progress Series, v. 56, p. 211–224.

41. Safronov, S.N., and Nikiforov, S.N., 2003, The list of pisciformes and fishes of the fresh and brackish waters of Sakhalin: 

Journal of Ichthyology, v. 43, no. 1, p. 38–49.
Pacific Capelin 89

42. Haldorson, L.J., Prichett, M., Paul, A.J., and Ziemann, D., 1993, Vertical distribution and migration of fish larvae in a 

northeast Pacific bay: Marine Ecology Progress Series, v. 101, p. 67–80.

43. Vilhjalmsson, H., 2002, Capelin (Mallotus villosus) in the Iceland-East Greenland-Jan Mayen ecosystem: ICES Journal of 

Marine Science, v. 59, no. 5, p. 870–883.

44. Friis-Rødel, E., and Kanneworff, P., 2002, A review of capelin (Mallotus villosus) in Greenland waters: ICES Journal of 

Marine Science, v. 59, no. 5, p. 890–896.

45. Morin, R., and Dodson, J.J., 1986, The ecology of fishes in James Bay, Hudson Bay and Hudson Strait, in Martini, I.P., ed., 

Canadian Inland Seas: Amsterdam, Elsevier Science Publishers B.V., p. 293–323.

46. Barton, L.H., 1978, Finfish resource surveys in Norton Sound and Kotzebue Sound: Alaska Department of Fish and Game, 

Commercial Fisheries Division, p. 75–313.

47. Brown, E.D., Churnside, J.H., Collins, R.L., Veenstra, T., Wilson, J.J., and Abnett, K., 2002, Remote sensing of capelin and 

other biological features in the North Pacific using lidar and video technology: ICES Journal of Marine Science, v. 59, no. 5, 

p. 1,120–1,130.

48. Naumenko, E.A., 1987, Daily feeding rhythm and ration of the capelin, Mallotus villosus socialis (Osmeridae), in the 

southeastern part of the Bering Sea during winter: Journal of Ichthyology, v. 27, no. 1, p. 158–161.

49. Savin, A.B., 2001, Dynamics of main biological indices of capelin Mallotus villosus catervarius (Osmeridae) in its 

wintering prespawning and postspawning aggregations off western Kamchatka: Journal of Ichthyology, v. 41, no. 8, 

p. 589–599.

50. Wilson, M.T., Jump, C.M., and Duffy-Anderson, J.T., 2006, Comparative analysis of the feeding ecology of two pelagic 

forage fishes—Capelin Mallotus villosus and walleye pollock Theragra chalcogramma: Marine Ecology Progress Series, 

v. 317, p. 245–258.

51. Dodson, J.J., Tremblay, S., Colombani, F., Carscadden, J.E., and Lecomte, F., 2007, Trans-Arctic dispersals and the 

evolution of a circumpolar marine fish species complex, the capelin (Mallotus villosus): Molecular Ecology, v. 16, 

p. 5,030–5,043.

52. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

53. Jarvela, L.E., and Thorsteinson, L.K., 1999, The epipelagic fish community of Beaufort Sea coastal waters, Alaska: Arctic, 

v. 52, no. 1, p. 80–94.

54. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

55. Tereshchenko, E.S., 2002, The dynamics of population fecundity in Barents Sea capelin: ICES Journal of Marine Science, 

v. 59, no. 5, p. 976–982.

56. Blackburn, J.E., Jackson, P.B., Warner, I.M., and Dick, M.H., 1981, A survey for spawning forage fish on the east side of the 

Kodiak Archipelago by air and boat during spring and summer 1979: Outer Continental Shelf Environmental Assessment 

Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 309–376.

57. Huse, G., and Gjosaeter, H., 1997, Fecundity of the Barents Sea capelin (Mallotus villosus): Marine Biology, v. 130, no. 2, 

p. 309–313.

58. Flynn, S.R., Nakashima, B.S., and Burton, M.P.M., 2001, Direct assessment of post-spawning survival of female capelin, 

Mallotus villosus: Journal of the Marine Biological Association of the United Kingdom, v. 81, p. 307–312.

59. Christiansen, J.S., Preaebel, K., Siikavuopio, S.I., and Carscadden, J.E., 2008, Facultative semelparity in capelin Mallotus 

villosus (Osmeridae)—An experimental test of a life history phenomenon in a sub-arctic fish: Journal of Experimental 

Marine Biology and Ecology, v. 360, no. 1, p. 47–55.

60. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320. 
90 Alaska Arctic Marine Fish Ecology Catalog

61. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennessee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.

62. Slotte, A., Mikkelsen, N., and Gjøsæter, H., 2006, Egg cannibalism in Barents Sea capelin in relation to a narrow spawning 

distribution: Journal of Fish Biology, v. 69, no. 1, p. 187–202.

63. Frost, K.J., and Lowry, L.F., 1981, Foods and trophic relationships of cetaceans in the Bering Sea, in Hood, D.W., and 

Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and Resources: National Oceanic and Atmospheric 

Administration, p. 825–836.

64. Hunt, G.L., Burgeson, B., and Sanger, G.A., 1981, Feeding ecology of seabirds of the Eastern Bering Sea, in Hood, D.W., 

and Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and Resources: National Oceanic and Atmospheric 

Administration, p. 629–641.

65. Gaston, A.J., Woo, K., and Hipfner, J.M., 2003, Trends in forage fish populations in northern Hudson Bay since 1981, as 

determined from the diet of nestling thick-billed murres Uria lomvia: Arctic, v. 56, no. 3, p. 227–233.

66. Yang, M.-S., Aydin, K., Greig, A., Lang, G.M., and Livingston, P.A., 2005, Historical review of capelin (Mallotus 

villosus) consumption in the Gulf of Alaska and eastern Bering Sea: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration Technical Memorandum NMFS-AFSC-155, 89 p. 

67. Davoren, G.K., May, C., Penton, P., Reinfort, B., Buren, A., Burke, C., and others, 2007, An ecosystem-based research 

program for capelin (Mallotus villosus) in the northwest Atlantic—Overview and results: Journal of the Northwest Atlantic 

Fishery Science, v. 39, p. 35–48.

68. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

69. Murdoch, J., 1884, Fish and fishing at Point Barrow, Arctic Alaska: Transactions of the American Fisheries Society, v. 13, 

no. 1, p. 111–115.

70. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

71. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

72. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

73. Mecklenburg, C.W., and Steinke, D., 2015, Ichthyofaunal baselines in the Pacific Arctic region and RUSALCA study area: 

Oceanography, v. 28, no. 3, p. 158–189.

74. Hart, J.L., 1973, Pacific fishes of Canada: Ottawa, Fisheries Research Board of Canada, Bulletin 180.
Arctic Smelt 91

Arctic Smelt (Osmerus dentex) 

Steindachner & Kner, 1870

Family Osmeridae 

Note on taxonomy: Previously called Osmerus mordax in 

references by authors, as well as O. eperlanus and O. mordax 

dentex populations from the Pacific Arctic are now recognized from 

molecular genetics and morphological studies to be a distinct 

species, O. dentex [1].

Colloquial Name: Iñupiat: Ithuagniq [2]; Ilhuagnig [3, 4]. 

Frequently called Rainbow Smelt and Boreal Smelt. 

Ecological Role: Likely of considerable importance as a prey species, at least in the Chukchi Sea.

Physical Description/Attributes: Elongate, slender body with olive or pale green back, sometimes speckled with black, and a 

silvery belly. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 174, as O. mordax) 

[5]. Swim bladder: Present, physostomous [6]. Antifreeze glycoproteins in blood serum: Present [7]. 

Range: U.S. Chukchi and Beaufort Seas. Elsewhere, White and Barents Seas eastward to Bathurst Inlet, Nunavut, and 

southward to North Korea, Japan, Sea of Okhotsk, and Heceta Head, Oregon [1, 8].

Relative Abundance: Common along all coasts of U.S. Chukchi and Beaufort Seas [11–14]. Common in Canadian Beaufort 

Sea as far east as Liverpool Bay [15–17].

Arctic Smelt (Osmerus dentex), 273 mm, eastern Chukchi 

Sea, 2007. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Arctic Smelt (Osmerus dentex) within Arctic Outer Continental Shelf Planning Areas 

[9] based on review of published literature and specimens from historical and recent collections [1, 5, 10].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Smelt 

Osmerus dentex

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
92 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Primarily in shallow, coastal waters of U.S. Chukchi and Beaufort Seas, common to a depth of about 25 m [18].

In Bering Sea and northeastern Pacific Ocean, nearshore, surface to 150 m, occasionally deeper but deep records probably due 

to fish entering nets nearer the surface than at maximum depth of tow [19]. In late autumn, migrate to bottom depths of 90 m or 

more in southwestern Bering Sea [20].

Habitats and Life History

Anadromous [8]. 

Eggs—Size: 0.8–1.0 mm [21, 22]. Time to hatching: 10–30 days depending on temperature [23–26]. Probably 

over 30 days on Alaskan North Slope in near-freezing waters [2]. Habitat: Freshwater, on gravel, sand, or plants 

in shallow, swift flowing waters (to depths of a few meters). Adheres to substrate until hatching [20, 23, 25, 26]. 

Larvae (fry)—Size at hatching: 5–8 mm SL [20, 27]. Size at juvenile transformation: Reported as post-larval at 

14.7 mm TL [27]. Days to juvenile transformation: Unknown. Habitat: Pelagic in brackish to marine waters [5, 

8]. Soon after hatching in freshwater, larvae are carried downriver and recruit to sheltered, shallow brackish and 

marine waters as small as 10–20 mm FL [15, 16, 28–30]. 

Juveniles—Age and size: A few months to 10 years [23–25, 31, 32]. Habitat: Pelagic in brackish to marine 

waters [5, 8]. Nearshore estuaries, embayments and, at least in southeastern Chukchi Sea, coastal waters [18]. 

Adults—Age and size at first maturity: Highly variable and ranges from 1 to 10 years or more [23–25, 31, 32]. 

Averages between 5–7 years and perhaps 20.0–22.5 cm FL [12, 26, 28, 33, 34]. Growth rates vary between 

areas. Length-weight relationships also vary with location and perhaps with year. Larger males may be heavier 

at length than females [16, 18]. Maximum age: At least 18 years in Arctic and subarctic waters [33], however, 

rarely longer than 15 years [22, 26, 30]. Fish in more temperate waters (specifically, southwestern Bering Sea 

and off Sakhalin Island, Russia) have much shorter life spans, rarely exceeding 6–9 years [20, 24, 25, 34]. 

Maximum size: 31.0 cm FL [8]. Habitat: Pelagic in brackish to marine waters [5, 8]. Nearshore estuaries, 

embayments and, at least in southeastern Chukchi Sea, coastal waters [18]. 

Substrate—Taken over sand-gravel in Bristol Bay [35]. 

Physical/chemical—Temperature: 2.0–13.5 °C. Tolerant of a very wide range [22]. Salinity: Tolerates brackish 

conditions, but typically 22 parts per thousand or greater and will avoid nearshore waters of lower salinities 

[26]. Although most fish enter fresh water only to spawn, landlocked populations are known [36]. 

 Benthic and reproductive distribution of Arctic Smelt (Osmerus dentex).

Some fish to 

150 m

Open ocean Shore

Depth, in meters

250

200

150

100

50

0

250

200

150

100

50

0

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Osmerus dentex

Arctic Smelt

Benthic distribution

Open ocean Shore

Depth, in meters

Larvae

Reproductive distribution

Anadromous.

Spawning and eggs

occur in freshwater

Overall data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Adults and

juveniles

Depth reported as common in 

U.S. Beaufort-Chukchi Seas
Arctic Smelt 93

Behavior

Diel—Enters rivers and spawns at night at least in Asia and eastern North America, [24, 25]. 

Seasonal—Schools of juveniles and adults inhabit nearshore waters during summer [20, 22, 29], although 

significant numbers feed as far as 10 km (6 mi) offshore [37]. Other than for spawning, fish in northeastern North 

America do not make extensive migrations [24], although those in southwestern Bering Sea do move offshore in 

early winter [20]. In U.S. Beaufort and Chukchi Seas, juveniles and adults overwinter under ice in brackish river 

deltas and coastal waters, whereas fish in southwestern Bering Sea retreat offshore to 90–100 m depths during 

early winter, returning to coastal waters in January and February [20]. Many river mouths along U.S. Chukchi 

and Beaufort Seas harbor overwintering populations [30, 32, 38–41]. Larvae and perhaps fertilized eggs are 

carried into marine waters during spring and early summer [23, 28, 30] 

Reproductive—Fish gather near spawning grounds as winter progresses [34]. Spawning takes place in spring, 

just prior to ice break-up [28, 30, 32]. Spawning takes place in many rivers entering U.S. Chukchi and Beaufort 

Seas [33, 34, 42] and in at least one lake (Lake Tasiqpaatchiaq, Alaska) [37]. Most spawning seems to occur in 

lowermost but still freshwater parts of rivers, often very near the mouth [23, 26]. However, fish in some Russian 

waters (for example, Yenisei River, Siberia) may travel upstream more than 1,000 km (621 mi) to spawning 

grounds [43] and some have been taken well upstream on the Mackenzie River in the Arctic Red River area, 

though it is not clear that spawning had occurred there [44]. Occasionally spawns in estuaries and possibly 

coastal marine waters [17, 27, 43]. Sticky and stalked eggs are shed over gravel, sand, or plants in shallow, swift 

flowing waters (to depths of a few meters) and adhere to the substrate until hatching [20, 23, 25, 26]. In Asia, 

adults often leave fresh waters within a few hours of spawning, although some remain in spawning area for 

several weeks [20]. At least some spawn more than once in their lifetimes [26]. 

Schooling—Schooling, water column fish [18]. 

Feeding—Midwater and, to a certain extent, benthic feeders. Feeding is most intense in summer, declines as 

winter progresses, and almost ceases during spring spawning [20, 22, 26, 33, 34].

Populations or Stocks

There have been no studies. Some life history parameters for Arctic Smelt in Simpson Lagoon, Alaska, were 

estimated [34].

Reproduction

Mode—Oviparous [8]. 

Spawning season—March–July, peaking in May–June in the U.S. Chukchi and Beaufort Sea drainages [12, 22, 

29, 34, 45]. May–July in Bering Sea and Asia [20, 25, 46]. 

Fecundity—1,700–207,900 eggs. Females lay eggs in small batches [24, 25].

Food and Feeding

Food items—Small fishes (for example, Arctic Cod, Fourhorn Sculpin, Arctic Cisco, Arctic Smelt, and eelpouts) 

and small crustaceans (for example, mysids, amphipods, isopods, and copepods) but occasionally snails, 

plant material, oligochaetes, penaeid shrimps, fish larvae, and insects [12, 16, 29, 34, 42]. Very young fish eat 

zooplankton and insects [33]. 

Trophic level—4.2 (standard error 0.73) [47].

Biological Interactions

Predators—Dolly Varden and other Arctic Smelt in Canadian Beaufort Sea [16, 30]. May be a major food for 

Beluga Whales between May and July in U.S. Chukchi Sea, at least in Wainwright area, and in eastern Bering 

Sea [48]. Extensively preyed upon by spotted seals in summer near Point Lay [26] and in April in eastern 

Chukchi Sea by ringed seals [49]. In eastern Bering Sea, other predators include harbor seals, Fin and Sei 

Whales [50, 51]. 

Competitors—Other water column piscivores and zooplanktivores such as Arctic Cod and Dolly Varden.
94 Alaska Arctic Marine Fish Ecology Catalog

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (t

m=2–3; tmax=7; Fecundity=1,700) [47].

Traditional and Cultural Importance

For many years, Arctic Smelt have been of great importance to the subsistence fisheries in the Wainwright, 

Alaska, area [52], where during winter and spring fishermen catch large numbers by jigging through the ice as 

these highly valued fish aggregate in the lower Kak River [2, 53]. Arctic Smelt are believed to be one of the few 

resources in the Wainwright area that were regularly sold [53]. During the autumn and winter of 1937, hunting 

was particularly poor around Wainwright and Arctic Smelt saved the local peoples from starvation [26]. Fish 

caught in November are perceived to taste saltier and are less valued than those taken later in the winter [26]. 

Elsewhere in U.S. Chukchi and Beaufort Seas, occasionally taken as bycatch in other subsistence fisheries

[53–55]. Also taken in some numbers in eastern Bering Sea [11] and off Russia [23].

Commercial Fisheries

Currently, Arctic Smelt are not commercially harvested.

Potential Effects of Climate Change

Arctic Smelt reproduce in both Arctic and Boreal waters [1], which makes it difficult to predict how their 

distribution might be affected by climate warming. Like other Arctic marine fish species, they are adapted to life 

in cold waters and changes in temperature could affect physiological functions such as growth and metabolism.

Areas for Future Research [A]

Little offshore research has been conducted in the Arctic and their abundance in offshore waters is unknown 

[26, 32], although likely to be negligible since Arctic Smelt are primarily a shallow-water coastal species. Basic 

life history information is limited and little is known about the larval and juvenile ecology of this species. 

Overwintering areas have not been described and no population studies have been conducted. Bioenergetic 

relationships, including consumption rates by high trophic level organisms need to be described as this species 

is believed to be of major forage importance in certain locales, such as coastally in the southeastern Chukchi Sea 

and near the Colville River Delta. 

References Cited

Belyanina, T.N., 1968, Dynamics of smelt populations in sub-arctic waters: Rapports et procès-verbaux des réunions / Conseil 

permanent international pour l’exploration de la mer, v. 158, p. 74–79. [23]

Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum ReserveAlaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife Management, Barrow, 

Alaska, 94 p. [26]

Gritsenko, O.F., Churikov, A.A., and Rodionova, S.S., 1984b, The reproductive ecology of the Arctic smelt, Osmerus mordax 

dentex (Osmeridae), in the rivers of Sakhalin Island: Journal of Ichthyology, v. 24, no. 3, p. 23–33. [25]

Haldorson, L.J., and Craig, P., 1984, Life history and ecology of a Pacific-Arctic population of rainbow smelt in coastal waters 

of the Beaufort Sea: Transactions of the American Fisheries Society, v. 113, no. 1, p. 33–38. [34]

Karpenko, V.I., and Vasilets, P.M., 1996, Biology of smelt (Osmeridae) in the Korf-Karagin coastal area of the southwestern 

Bering Sea, in Mathisen, O.A., and Coyle, K.O., eds., Ecology of the Bering Sea—A review of Russian literature: Fairbanks, 

Alaska, University of Alaska, Alaska Sea Grant Program, AK-SG-96-01, p. 217–235. [20]
Arctic Smelt 95

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [5]

Bibliography

1. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

2. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

3. Nelson, R.K., 1981, Harvest of the sea—Coastal subsistence in modern Wainwright: North Slope Borough, 112 p.

4. Pedersen, S., and Linn, A., Jr., 2005, Kaktovik 2000–2002 subsistence fishery harvest assessment: U.S. Fish and Wildlife 

Service, Fisheries Resource Monitoring Program, Final Report for FIS Study, Study No. 01-101.

5. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

6. Jørgensen, R., 2003, The effects of swimbladder size, condition and gonads on the acoustic target strength of mature 

capelin: ICES Journal of Marine Science, v. 60, no. 5, p. 1,056–1,062.

7. Power, G., 1997, A review of fish ecology in Arctic North America: American Fisheries Society Symposium, no. 19, 

p. 13–39.

8. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

9. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

10. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

11. Barton, L.H., 1978, Finfish resource surveys in Norton Sound and Kotzebue Sound: Alaska Department of Fish and Game, 

Commercial Fisheries Division, p. 75–313.

12. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

13. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

14. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

15. Jones, M.L., and Den Beste, J., 1977, Tuft Point and adjacent coastal areas fisheries projects: Calgary, Alberta, Canada, 

Aquatic Environments, Ltd., 152 p.

16. Bond, W.A., and Erickson, R.N., 1987, Fishery data from Phillips Bay, Yukon, 1985: Winnipeg, Manitoba, Canadian Data 

Report of Fisheries and Aquatic Sciences, Central and Artic Region, Department of Fisheries and Oceans, no. 635, 47 p.

17. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

18. Wolotira, R.J., Jr., Sample, T.M., and Morin, M., Jr., 1977, Demersal fish and shellfish resources of Norton Sound, the 

southeastern Chukchi Sea, and adjacent waters in the baseline year 1976: Seattle, Washington, Northwest and Alaska 

Fisheries Center, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Processed Report, 

69 p.

19. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.
96 Alaska Arctic Marine Fish Ecology Catalog

20. Karpenko, V.I., and Vasilets, P.M., 1996, Biology of smelt (Osmeridae) in the Korf-Karagin coastal area of the southwestern 

Bering Sea, in Mathisen, O.A., and Coyle, K.O., eds., Ecology of the Bering Sea—A review of Russian literature: 

Fairbanks, Alaska, University of Alaska, Alaska Sea Grant Program, AK-SG-96-01, p. 217–235.

21. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

22. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

23. Belyanina, T.N., 1968, Dynamics of smelt populations in sub-arctic waters: Rapports et procès-verbaux des réunions / 

Conseil permanent international pour l’exploration de la mer, v. 158, p. 74–79.

24. Morrow, J.E., 1980, The freshwater fishes of Alaska: Anchorage, Alaska Northwest Publishing Company, 248 p.

25. Gritsenko, O.F., Churikov, A.A., and Rodionova, S.S., 1984b, The reproductive ecology of the Arctic smelt, Osmerus 

mordax dentex (Osmeridae), in the rivers of Sakhalin Island: Journal of Ichthyology, v. 24, no. 3, p. 23–33.

 26. Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum 

Reserve-Alaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife 

Management, Barrow, Alaska, 94 p.

27. Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbour, N.W.T.: Canadian Technical 

Report of Fisheries and Aquatic Sciences, no. 1218, 21 p.

28. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.

29. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.

30. Bond, W.A., and Erickson, R.N., 1989, Summer studies of the nearshore fish community at Phillips Bay, Beaufort Sea coast, 

Yukon: Winnepeg, Manitoba, Canadian Technical Report of Fisheries and Aquatic Sciences, Central and Arctic Region, 

Department of Fisheries and Oceans, no. 1676, 102 p.

31. Bendock, T.N., 1977, Beaufort Sea estuarine fishery study: Alaska Department of Fish and Game Annual Report, Contract 

#03-5-022-69, p. 670–729.

32. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 

1119, 90 p.

33. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

34. Haldorson, L.J., and Craig, P., 1984, Life history and ecology of a Pacific-Arctic population of rainbow smelt in coastal 

waters of the Beaufort Sea: Transactions of the American Fisheries Society, v. 113, no. 1, p. 33–38.

35. Johnson, S.W., Thedinga, J.F., and Lindeberg, M.R., 2012, Nearshore fish atlas of Alaska: National Oceanic and 

Atmospheric Administration Fisheries, accessed February 2012 at http://www.fakr.noaa.gov/habitat/fishatlas/.

36. McPhail, J.D., and Lindsey, C.C., 1970, Freshwater fishes of northwestern Canada and Alaska: Bulletin of the Fisheries 

Research Board of Canada Bulletin 173, 381 p.

37. Moulton, L.L., Owl Ridge Natural Resource Consultants, written commun., 2011.

38. Schmidt, D.R., Griffiths, W.B., and Martin, L.R., 1987, Importance of anadromous fish overwintering habitat in the 

Sagavanirktok River Delta, Alaska: Anchorage, Alaska, Report by Ecological Research Associates for Standard Alaska 

Production Company and North Slope Borough, 71 p.

39. Bond, W.A., and Erickson, R.N., 1993, Fisheries investigations in coastal waters of Liverpool Bay, Northwest Territories: 

Winnipeg, Manitoba, Canada Department of Fisheries and Oceans, Central and Arctic Region, Canadian Manuscript Report 

of Fisheries and Aquatic Sciences No. 2204, 59 p.
Arctic Smelt 97

40. Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.

41. Sekerak, A.D., Stallard, N., and Griffiths, W.B., 1992, Distribution of fish and fish harvests in the nearshore Beaufort Sea 

and Mackenzie Delta during ice-covered periods, October–June: Environmental Studies Research Funds Report, LGS Ltd. 

No. 117, 524 p.

42. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

43. Berg, L.S., 1948, Freshwater fishes of the USSR and adjacent countries, volume 1 (4th ed.): Moscow, Academy of Sciences 

of the USSR Zoological Institute, 466 p. [Translated from Russian by Israel Program for Science Translation, Jerusalem, 

505 p.]

44. Hatfield, C.T., Stein, J.N., Falk, M.R., and Jessop, C.S., 1972, Fish resources of the Mackenzie River Valley: Winnepeg, 

Environment Canada Fisheries Service, Interim report 1, v. 1.

45. Mann, G.J., 1975, Winter fisheries survey across the Mackenzie Delta, in Craig, P.C., ed., Fisheries investigations in a 

coastal region of the Beaufort Sea: Arctic Gas Biological Report Series, v. 34.

46. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

47. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

48. Lowry, L.F., Frost, K.J., and Seaman, G.A., 1986, Investigations of belukha whales in coastal waters of western and 

northern Alaska: Outer Continental Shelf Environmental Program Unit 612, Final Report, p. 359–392.

49. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.

50. Lowry, L.F., and Frost, K.J., 1981, Feeding and trophic relationships of phocid seals and walruses in the eastern Bering Sea, 

in Hood, D.W., and Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and resources: National Oceanic and 

Atmospheric Administration, p. 813–824.

51. Frost, K.J., and Lowry, L.F., 1981, Foods and trophic relationships of cetaceans in the Bering Sea, in Hood, D.W., and 

Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and Resources: National Oceanic and Atmospheric 

Administration, p. 825–836.

52. Murdoch, J., 1884, Fish and fishing at Point Barrow, Arctic Alaska: Transactions of the American Fisheries Society, v. 13, 

no. 1, p. 111–115.

53. Craig, P.C., 1989b, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers of the 

University of Alaska, v. 24, p. 131–152.

54. George, J.C., and Kovalsky, R., 1986, Observations on the Kupigruak Channel (Colville River) subsistence fishery, October 

1985: Alaska Department of Wildlife Management, 66 p.

55. Moulton, L.L., Field, L.J., and Kovalsky, R., 1991, Predictability in the catch of Arctic cisco in the Colville River, Alaska: 

American Fisheries Society Symposium no. 11, p. 145–156.
98 Alaska Arctic Marine Fish Ecology Catalog

Arctic Cisco (Coregonus autumnalis)

(Pallas, 1776)

Family Salmonidae 

Colloquial Name: Iñupiat: Qaatag, Qaaktaq [1]; Qaaqtaq [2].

Ecological Role: As one of the most common and widely 

distributed coregonids found in Alaskan Beaufort Sea coastal 

waters during summer [3], this species is a prominent member 

of the nearshore fish assemblage. Arctic Cisco is an important 

subsistence resource.

Physical Description/Attributes: Slender body with a dark brown to green back, silver belly, and pale (almost colorless) anal, 

pectoral, and pelvic fins. Unlike other ciscoes, does not have black spots on back or white spots on fins. Lower jaw does not 

protrude beyond upper jaw. Very closely resembles Bering Cisco. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 183) [4]. Swim bladder: Present [5]. Swim bladder ruptures have been documented when 

exposed to explosive-based instantaneous pressure change [6]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Beaufort Sea westward to northeastern Chukchi Sea at Point Lay [7]. Worldwide, along coasts from the White Sea 

east through Siberia and to Mackenzie River, Canada [8]. 

Relative Abundance: Rare from Barrow to Point Lay in U.S. Chukchi Sea [7, 11]. Common along coast of U.S. Beaufort 

Sea eastward from at least Colville River [12]. Common at least as far east as Coppermine River mouth in Coronation Gulf, 

Canada [13].

Arctic Cisco (Coregonus autumnalis). Photograph by 

Kirk Waggoner, MJM Research LCC.

Geographic distribution of Arctic Cisco (Coregonus autumnalis) at sea within Arctic Outer Continental 

Shelf Planning Areas [9] based on review of published literature and specimens from historical and recent 

collections [4, 8, 10]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Cisco

Coregonus autumnalis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Arctic Cisco 99

Depth Range: Nearshore, rarely deeper than about 6 m. Common within a few hundred meters from shore. [12, 14–19]. 

However, in Canadian Beaufort Sea, juveniles have been taken near the surface as far as 50 km offshore [20].

Habitats and Life History

Anadromous. There appear to be some landlocked populations [15]. 

Eggs—Size: 0.5–1.3 mm [21]. Time to hatching: Unknown. Habitat: Gravel beds in fast flowing freshwater 

rivers [15, 22]. 

Larvae—Size at hatching: As small as 19 mm FL [15, 22]. Size at juvenile transformation: About 45–54 mm FL

[12, 15, 23]. Days to juvenile transformation: Unknown. Habitat: Freshwater rivers to nearshore estuaries [22, 

24, 25]. 

Juveniles—Age and size: 0–5 years and 4.5–32.8 cm FL [21, 23]. Habitat: Nearshore brackish or marine waters 

[22, 24, 25]. Occasionally, juveniles have been found well upstream in some river systems (for example, Colville 

and Babbage Rivers) [15, 26]. 

Adults—Age and size at first maturity: As young as 5 years in Russia, where males mature about 1 year earlier 

than females [27]. North American fish mature at a wide range of ages. A few mature as early as 5 years, many 

at 6–8 years, and some at perhaps 11 years or older. A few are mature by 32.8 cm FL and virtually all by about 

40.0 cm FL [15, 21, 26, 28–30]. Length-weight relationships appear to be similar along much of the species’

range. Male and female growth rates are similar [31], but females are larger at older ages than males and may 

also be heavier at length [30]. Fish weigh less at length during years of cold water and heavy ice packs [32]. 

Maximum age: At least 21 years [33]. Males and females may have similar life spans [26, 27]. Maximum size: 

64 cm TL [4]. Habitat: Pelagic, nearshore brackish or marine waters and freshwater rivers [22, 24, 25]. Adults 

generally re-enter fresh water only to spawn and then return to estuarine coastal waters [22, 24, 25]. 

Substrate—Gravel for spawning [15, 22]. Taken over sand-gravel in Chukchi Sea [34]. 

Physical/chemical—Temperature:–1.–13.5 °C [12]. Salinity: 0–30.0 parts per thousand [15, 35], mainly in 

10–25 parts per thousand except when spawning in fresh water [15, 21]. Prefers relatively warm and brackish 

conditions [16, 35–38] but tolerant of cold and saline waters [17, 32, 39–41]. May grow faster in warm and lowsalinity water [37].

Benthic and reproductive distribution of Arctic Cisco (Coregonus autumnalis). 

3-2-7gArcticCisco

Open ocean Shore

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

Some fish to 6 m Larvae depths estimated.

Reported as nearshore estuaries

Adults and juveniles

Coregonus autumnalis

Arctic Cisco

Benthic distribution

Data from U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning and eggs 

occur in freshwater
100 Alaska Arctic Marine Fish Ecology Catalog

Behavior

Diel—Unknown. 

Seasonal—Spawning occurs in autumn and eggs hatch during spring in MacKenzie River, Canada [15, 22]. 

Yolk-sac larvae are flushed downstream into the river delta in late May or early June [15, 22]. Migrations to 

either east or west are passive, depending on strength and direction of winds and currents. The predominant 

westerly winds tend to propel fish along Tuktoyaktuk Peninsula, Canada (at least as far as Liverpool Bay, near 

the Anderson River) [15]. Strong easterlies assist their wind-aided migration westward, often to Colville River 

area, although many are carried only as far as the Yukon Territory coast [39, 42]. Successful year classes that 

reach the Colville River Delta are associated with summers when easterly winds are strong and more-or-less 

continuous, often of 5 km/h or more [17, 29, 43, 44]. Eastward-moving juveniles often stay within 100 m of 

shore although more offshore migrations may occur [15, 17, 45]. Young-of-the-year first occurs off Yukon coast 

(Phillips Bay) between early July and September, and recruit to the Prudhoe Bay-Colville River-Simpson Lagoon 

area between mid-August and late September [12, 15, 23]. 

Juveniles migrate to overwintering grounds as autumn approaches. In Alaska, most fish winter under ice in 

brackish, deep channels of the Colville River, and some in lower parts of the Sagavanirktok River [15, 24, 

46]. However, the Sagavanirktok River may not provide sufficient annual winter refuge to sustain long-term 

populations [40]. An estimated 1.2–1.8 million individuals larger than 250 mm FL overwinter in the Colville 

River Delta [46]. To the east, wintering grounds are in the Mackenzie River Delta (perhaps as far west as 

Herschel Island), as well as in bays and lagoons along the Tuktoyaktuk Peninsula, in Tuktoyaktuk Harbour, and 

as far east as at least the Anderson River [22, 25, 31, 47, 48]. 

Juveniles leave overwintering grounds in summer when waters warm and disperse to feed in the nearshore, some 

moving at an average rate of 2.9 km/d [12]. Younger fish tend to remain in brackish waters and do not venture far 

[39]. Older juveniles migrate farther during summer and are able to tolerate more saline conditions. Regardless 

of size, juveniles always begin to return to overwintering grounds after a few months and are usually in place 

by September [12, 15, 49]. At least in Arctic National Wildlife Refuge region, larger fish tend to move back to 

overwintering grounds earlier than do smaller ones [16].

Populations or Stocks

Fish utilizing different Mackenzie River tributaries may form different genetic stocks [50]. 

Reproduction

Mode—Iteroparous [26]. 

Spawning season—September to early October in Mackenzie River tributaries [15, 51], and September–

December in Russia. 

Fecundity—11,316–30,267 eggs in North America [30] and 7,700–52,000 eggs in Russia [15, 28]. 

Reproductive—Upon maturity, their life cycle is dominated by migrations to and from spawning sites. Adults 

migrate back to the Mackenzie River, spending less time than usual in coastal waters. Autumn spawners enter the 

river from May to early August [22]. Little is known about spawning behaviors or specific conditions. Spawning 

occurs in fast waters over gravel [27, 52]. Most females spawn every other year [21, 26, 27, 51]. Post-spawning 

fish move downstream and overwinter in the Mackenzie River Delta [22, 52]. During the next spring and 

summer they will disperse, with some exceptions, as far westward as Barter Island and at least as far eastward 

as the Anderson River [15, 22, 26]. The presence of a few older individuals (10–15 years) along the North Slope 

as far west as Simpson Lagoon implies that older fish will occasionally make more extensive migrations [12, 16, 

18, 32].
Arctic Cisco 101

Although common along much of the Beaufort Sea coast, most, or perhaps all Arctic Cisco are believed to spawn 

in Mackenzie River tributaries such as Great Bear, Arctic Red, Peel, and Liard Rivers, the latter being more than 

1,700 km from the Beaufort Sea [15, 42, 53]. Evidence for limited spawning in other waterways is discussed 

in the section, “Remarks.” Intertidal or subtidal spawning in estuaries and perhaps the sea has been reported in 

Russian waters [27] but has not been observed in North America. 

Schooling—Forms schools, often in groups of tens to several hundreds. Sometimes schools with Dolly Varden 

[12, 18, 26]. Individuals may stay together in same school for months at a time (specifically, several fish tagged 

on same date at Simpson Lagoon were recaptured together several months later) [12]. 

Feeding—Opportunistic feeders. Feed under ice during winter (at a reduced rate) [46], although food habits 

may change, reflecting differences in food availability [12]. Rarely feeds during spawning migrations [48, 51, 

52, 54].

Food and Feeding

Food items—A wide variety of benthic and water column prey. Important prey include various crustaceans (for 

example, amphipods, copepods, mysids, and cladocerans), insects (particularly chironomids), snails, clams, 

polychaetes, fishes (for example, Fourhorn Sculpin and Arctic Cisco), fish eggs, and occasionally plant material 

[12, 31, 35, 54]. 

Trophic level—3.57 (standard error 0.56) [55].

Biological Interactions

Predators—Dolly Varden, Arctic Smelt, and Arctic Cisco [21, 26]. 

Competitors—Shallow, nearshore species such as Dolly Varden, Arctic Cod, other whitefishes, and sculpins.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time is 4.5–14 years (t

m=6; Fecundity=2,000) [55].

Traditional and Cultural Importance 

Arctic Cisco are widely taken in subsistence fisheries along much of the U.S. Beaufort Sea coast. Juvenile Arctic 

Cisco in particular form the basis for major subsistence in the Colville River Delta. Most of the fish are captured 

under the ice by gill nets during the autumn. Currently, the principal fishing areas on the Colville River are in the 

lower delta and near the village of Nuiqsut [29, 56, 57]. Part of this catch is distributed to other parts of Alaska

[12]. The summer fishery at Kaktovik may catch newly matured fish as they migrate back to the Mackenzie River

[1, 58]. Annual catch records have been collected since 1968.

Commercial Fisheries

A small commercial fishery for Arctic Cisco in the Colville River Delta was terminated in 2010. Currently, there 

is no commercial fishing for Arctic Cisco.

Potential Effects of Climate Change

Unknown, as the effect of climate change on the Mackenzie River system and on the wind patterns that control 

juvenile movements, are unclear. However, von Biela and others (2011) [59] determined that young-of-the-year 

growth increased during years of stronger east winds, as well as reduced sea-ice concentration and Mackenzie 

River discharge, and that there was a time lag of one or 2 years. Generally, Durand and others (2011) [60] predict 

that, at least for anadromous fishes in subarctic rivers, shifts in biology will be effected by spring ice break-up 

and resultant peak flows and surrounding permafrost processes: both of which affect the supply of nutrients and 

(or) sediment to the watershed of climate change on spring break-up intensity.
102 Alaska Arctic Marine Fish Ecology Catalog

Areas for Future Research [A]

Although it is clear that Arctic Cisco frequently use nearshore, shallow waters for feeding and migration, the 

role, if any, of offshore waters has not been completely investigated. The physiological tolerance of young-ofthe-year fish to cold, high salinity water has been suggested but not confirmed in laboratory studies and may be 

an important constraint to recruitment in Alaskan waters. Environmental tolerance experiments including effects 

of different temperature and salinity regimes on the growth and survival of Arctic Cisco are needed to assess the 

species vulnerability to climatic changes. In addition, the location and significance of important habitats in the 

Mackenzie River and the potential for isolated spawning stocks should also be explored. Studies to describe the 

genetic relationships between Arctic and Bering Cisco are needed. Coastal monitoring at key reference locations 

should be designed to track changes in population health (growth, survival, recruitment, and condition).

Remarks

Although it is clear that most, if not all, Beaufort Sea Arctic Cisco spawn in the Mackenzie River, there is evidence that 

spawning may occur on other grounds. Colonell and Gallaway (1997) [53] provide indirect evidence for some spawning west 

of the Colville River. They noted that some subsistence fishermen near Barrow stated that they captured mature Arctic Cisco in 

their autumn fishery. The Colonell and Gallaway (1997) indicated that on several occasions strong westerly winds, anticipated 

to lead to poor recruitment of young Arctic Cisco in the Colville River area (through poor transport from the Mackenzie River 

to the east), led instead to large recruitment. This would imply that westerly currents from some spawning waters west of the 

Colville River carried young fish to the study site. Colonell and Gallaway (1997) [53] also cited three genetic studies that posit 

a genetically differentiated group of Arctic Cisco in western Alaska. Bond and Erickson (1997) [22] reported on the capture of 

young-of-the-year Arctic Cisco at the mouth of the Anderson River in early July. They suggested that these captures were too 

early in the season to be fish that had hatched in the Mackenzie River and then carried eastward for hundreds of kilometers. 

They also noted that capture in Wood Bay (into which the Anderson River empties) of hybrid Arctic Cisco-other coregonids; 

captures that imply that Arctic Cisco were mating with fishes in that region. Lastly, Bickham and others (1997) [61] determined 

that some Arctic Cisco from the western Beaufort Sea carried mitochondrial DNA of Bering Cisco (C. laurettae). As Bering 

Cisco are unknown from the Mackenzie River (the putative site of all Arctic Cisco spawning) it appears that some Arctic Cisco 

spawn to the west of the MacKenzie River.

Arctic Cisco are closely related to Lake Cisco (Coregonus artedi) and Bering Cisco (C. laurettae) [62]. In the Northwest 

Territories, they occasionally hybridize with Least Cisco and Humpback Whitefish [63]. 

References Cited

Bond, W.A., and Erickson, R.N., 1997, Coastal migrations of Arctic ciscoes in the eastern Beaufort Sea, in Reynolds, J., ed., 

Fish ecology in Arctic North America, symposium 19: Bethesda, Maryland, American Fisheries Society, p. 155–164. [22]

Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum ReserveAlaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife Management, Barrow, 

Alaska, 94 p. [15]

Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, Simpson 

Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649. [12]

Griffiths, W.B., Craig, P., Walder, G., and Mann, G., 1975, Fisheries investigations in a coastal region of the Beaufort Sea 

(Nunaluk Lagoon, Yukon), in Craig, P.C., ed., Fisheries investigations in a coastal region of the Beaufort Sea: Arctic Gas, 

Biological Report Series, v. 34, p. 1–129. [26]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [4]
Arctic Cisco 103

Bibliography

1. Craig, P.C., 1989b, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers University 

of Alaska, no. 24, p. 131–152.

2. Pedersen, S., and Linn, A., Jr., 2005, Kaktovik 2000–2002 subsistence fishery harvest assessment: U.S. Fish and Wildlife 

Service, Fisheries Resource Monitoring Program, Final Report for FIS Study, Study No. 01-101.

3. Craig, P.C., 1984, Fish use of coastal waters of the Alaskan Beaufort Sea—A review: Transactions of the American 

Fisheries Society, v. 113, no. 3, p. 265–282.

4. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

5. Moyle, P.B., and Cech, J.J., Jr., 1996, Fishes—An introduction to ichthyology: Upper Saddle River, New Jersey, PrenticeHall, 590 p.

6. Godard, D.R., 2010, Pathological examination of fish exposed to explosive based instantaneous pressure change: Winnipeg, 

Univerity of Monitoba, Master’s thesis, 253 p.

7. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

8. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

9. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

10. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

11. Philo, L.M., George, J.C., and Moulton, L.L., 1993, The occurrence and description of anadromous fish in the Dease Inlet/

Admiralty Bay, Alaska area, 1988–1990: Barrow, Alaska, Department of Wildlife Management, North Slope Borough and 

MJM Research, 150 p.

12. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

13. Gillman, D.V., and Kristofferson, A.H., 1984, Biological data on Pacific herring (Clupea harengus pallasi) from 

Tuktoyaktuk Harbour and the Liverpoor Bay area, Northwest Territories, 1981 to 1983: Winnipeg, Manitoba, Canada, 

Canadian Data Report of Fisheries and Aquatic Sciences, Department of Fisheries and Oceans, Western Region, no. 485, 

22 p.

14. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

15. Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum 

Reserve-Alaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife 

Management, Barrow, Alaska, 94 p.

16. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

17. Schmidt, D.R., Griffiths, W.B., Beaubien, D.K., and Herlugson, C.J., 1991, Movement of young-of-the-year Arctic ciscoes 

across the Beaufort Sea coast, 1985–1988: American Fisheries Society Symposium, no. 11, p. 132–144.
104 Alaska Arctic Marine Fish Ecology Catalog

18. Wiswar, D.W., West, R.L., and Winkleman, W.N., 1995, Fisheries investigation in Oruktalik Lagoon, Arctic National 

Wildlife Refuge, Alaska, 1986: U.S. Fish and Wildlife Service, Alaska Fisheries Technical Report, no. 30, 38 p. 

19. Jarvela, L.E., and Thorsteinson, L.K., 1997, Movements and temperature occupancy of sonically tracked Dolly Varden and 

Arctic ciscoes in Camden Bay, Alaska: American Fisheries Society Symposium, no. 19, p. 165–174.

20. Majewski, A.R., Sareault, J.E., and Reist, J., 2006, Fish catch data from offshore sites in the Mackenzie River estuary and 

Beaufort Sea during the open water season, August 2004 aboard the CCGS Nahidik: Winnipeg, Manitoba, Fisheries and 

Oceans Canada, no. 2771, 42 p.

21. Bond, W.A., and Erickson, R.N., 1987, Fishery data from Phillips Bay, Yukon, 1985: Winnipeg, Manitoba, Canadian Data 

Report of Fisheries and Aquatic Sciences, Central and Artic Region, Department of Fisheries and Oceans, no. 635, 47 p.

22. Bond, W.A., and Erickson, R.N., 1997, Coastal migrations of Arctic ciscoes in the eastern Beaufort Sea, in Reynolds, J., ed., 

Fish ecology in Arctic North America, symposium 19: Bethesda, Maryland, American Fisheries Society, p. 155–164.

23. Moulton, L.L., 1989, Recruitment of Arctic cisco (Coregonus autumnalis) into the Colville Delta, Alaska, in 1985: 

Biological Papers of the University of Alaska, no. 24, p. 107–111.

24. Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.

25. Sekerak, A.D., Stallard, N., and Griffiths, W.B., 1992, Distribution of fish and fish harvests in the nearshore Beaufort Sea 

and Mackenzie Delta during ice-covered periods, October–June: Environmental Studies Research Funds Report, LGS Ltd. 

No. 117, 524 p.

26. Griffiths, W.B., Craig, P., Walder, G., and Mann, G., 1975, Fisheries investigations in a coastal region of the Beaufort Sea 

(Nunaluk Lagoon, Yukon), in Craig, P.C., ed., Fisheries investigations in a coastal region of the Beaufort Sea: Arctic Gas, 

Biological Report Series, v. 34, p. 1–129.

27. Berg, L.S., 1948, Freshwater fishes of the USSR and adjacent countries, volume 1 (4th ed.): Moscow, Academy of Sciences 

of the USSR Zoological Institute, 466 p. [Translated from Russian by Israel Program for Science Translation, Jerusalem, 

505 p.]

28. Craig, P.C., and Mann, G.J., 1974, Life history and distribution of Arctic Cisco (Coregonus autumnalis) along the Beaufort 

Sea Coastline in Alaska and the Yukon Territory, in McCart, P.J., ed., Life histories of anadromous and freshwater fish in the 

western Arctic: Aquatic Environments, Ltd., Arctic Gas Biological Report Series, v. 20, Prepared for Canadian Arctic Gas 

Study Limited and Alaskan Arctic Gas Study Company, 225 p.

29. Fechhelm, R.G., Streever, B., and Gallaway, B.J., 2007, The Arctic cisco (Coregonus autumnalis) subsistence and 

commercial fisheries, Colville River, Alaska—A conceptual model: Arctic, v. 60, no. 4, p. 421–429.

30. Van Gerwen-Toyne, M., Walker-Larsen, J., and Tallman, R.F., 2008, Monitoring spawning populations of migratory 

coregonids in the Peel River, NT—The Peel River fish study 1998-2002: Canadian Manuscript Report of Fisheries and 

Aquatic Sciences no. 2851, 56 p.

31. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 

1119, 90 p.

32. Underwood, T.J., Gordon, J.A., Millard, M.J., Thorpe, L.A., and Osborne, B.M., 1995, Characteristics of selected fish 

populations of Arctic National Wildlife Refuge coastal waters, final report, 1988–91: Fairbanks, Alaska, U.S. Fish and 

Wildlife Service, Fairbanks Fishery Resource Office, Alaska Fisheries Technical Report, no. 28.

33. Craig, P.C., and McCart, P.J., 1975, Fish utilization of nearshore coastal waters between the Colville and Mackenzie rivers 

with an emphasis on anadromous species, in Craig, P.C., ed., Fisheries Investigations in a coastal region of the Beaufort 

Sea: Calgary, Alberta, Canadian Arctic Gas Study Ltd. and Alaskan Arctic Gas Study Co., Biological Report Series 34, 

p. 172–219.

34. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.
Arctic Cisco 105

35. Fechhelm, R.G., Bryan, J.D., Griffiths, W.B., and Martin, L.R., 1997, Summer growth patterns of northern Dolly Varden 

(Salvelinus malma) smolts from the Prudhoe Bay region of Alaska: Canadian Journal of Fisheries and Aquatic Sciences, 

v. 54, no. 5, p. 1,103–1,110.

36. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

37. Hachmeister, L.E., Glass, D.R., and Cannon, T.C., 1991, Effects of solid-fill gravel causeways on the coastal central 

Beaufort Sea environment: American Fisheries Society Symposium, no. 11, p. 81–96.

38. Robertson, S.B., 1991, Habitats versus populations—Approaches for assessing impacts on fish: American Fisheries Society 

Symposium, no. 11, p. 97–108.

39. Griffiths, W.B., Fechhelm, R.G., Gallaway, B.J., and Fissel, D.B., 1988, Yukon fish and effects of causeway: For Indian and 

Northern Affairs Canada, LGL Ltd. and Arctic Science Ltd., 124 p. 

40. Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., and Colonell, J.M., 1996, The 1995 fish and oceanography 

study in Mikkelsen Bay, Alaska: Anchorage, Alaska, Prepared by LGL Alaska Research Associates and Woodward-Clyde 

Consultant for BP Exploration (Alaska) Inc.

41. Jarvela, L.E., and Thorsteinson, L.K., 1999, The epipelagic fish community of Beaufort Sea coastal waters, Alaska: Arctic, 

v. 52, no. 1, p. 80–94.

42. Gallaway, B.J., Griffiths, W.B., Craig, P.C., Gazey, W.J., and Helmericks, J.W., 1983, An assessment of the Colville River 

Delta stock of Arctic cisco—Migrants from Canada?: Biological Papers of the University of Alaska, no. 21, p. 4–23.

43. Fechhelm, R.G., and Fissel, D.B., 1988, Wind-aided recruitment of Canadian Arctic cisco (Coregonus autumnalis) into 

Alaskan waters: Canadian Journal of Fisheries and Aquatic Sciences, v. 45, no. 5, p. 906–910. 

44. Fechhelm, R.G., and Griffiths, W.B., 1990, Effect of wind on the recruitment of Canadian Arctic cisco (Coregonus 

autumnalis) into the central Alaskan Beaufort Sea: Canadian Journal of Fisheries and Aquatic Sciences, v. 47, no. 11, 

p. 2,164–2,171.

45. Gallaway, B.J., Gazey, W.J., Colonell, J.M., Niedoroda, A.W., and Herlugson, C.J., 1991, The Endicott development 

project-preliminary assessment of impacts from the first major offshore oil development in the Alaskan Arctic: American 

Fisheries Society Symposium, no.11, p. 42–80.

46. Schmidt, D.R., Griffiths, W.B., and Martin, L.R., 1987, Importance of anadromous fish overwintering habitat in the 

Sagavanirktok River Delta, Alaska: Anchorage, Alaska, Report by Ecological Research Associates for Standard Alaska 

Production Company and North Slope Borough, 71 p.

47. Kendel, R.E., Johnston, R.A.C., Kozak, M.D., and Lobsiger, U., 1974, Movements, distribution, populations and food 

habits of fish in the western coastal Beaufort Sea: Department of the Environment, Canada, Interim Report of Beaufort Sea 

Project, Study B1, 64 p.

48. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.

49. Bond, W.A., and Erickson, R.N., 1989, Summer studies of the nearshore fish community at Phillips Bay, Beaufort Sea coast, 

Yukon: Winnepeg, Manitoba, Canadian Technical Report of Fisheries and Aquatic Sciences, Central and Arctic Region, 

Department of Fisheries and Oceans, no. 1676, 102 p.

50. Bickham, J.W., Carr, S.M., Hanks, B.G., Burton, D.W., and Gallaway, B.J., 1989, Genetic analysis of population variation 

in the Arctic cisco (Coregonus autumnalis) using electrophoretic, flow cytometric, and mitochondrial DNA restriction 

analyses: Biological Papers of the University of Alaska, no. 24, p. 112–122.

51. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

52. Hatfield, C.T., Stein, J.N., Falk, M.R., and Jessop, C.S., 1972, Fish resources of the Mackenzie River Valley: Winnepeg, 

Environment Canada Fisheries Service, Interim report 1, v. 1.53.
106 Alaska Arctic Marine Fish Ecology Catalog

53. Colonell, J.M., and Gallaway, B.J., 1997, Wind-driven transport and dispersion of age-0 Arctic ciscoes along the Alaska 

Beaufort coast: American Fisheries Society Symposium, no. 19, p. 90–103.

54. Jones, M.L., and Den Beste, J., 1977, Tuft Point and adjacent coastal areas fisheries projects: Calgary, Alberta, Canada, 

Aquatic Environments, Ltd., 152 p.

55. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

56. George, J.C., and Kovalsky, R., 1986, Observations on the Kupigruak Channel (Colville River) subsistence fishery, October 

1985: Alaska Department of Wildlife Management, 66 p.

57. Moulton, L.L., Field, L.J., and Kovalsky, R., 1991, Predictability in the catch of Arctic cisco in the Colville River, Alaska: 

American Fisheries Society Symposium no. 11, p. 145–156.

58. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

59. von Biela, V.R., Zimmerman, C.E., and Moulton, L.L., 2011, Long-term increases in young-of-the-year growth of Arctic 

cisco Coregonus autumnalis and environmental influences: Journal of Fish Biology, v. 78, no. 1, p. 39–56.

60. Durand, J.R., Lusardi, R.A., Nover, D.M., Suddeth, R.J., Carmona-Catot, G., Connell-Buck, C.R., and others, 2011, 

Environmental heterogeneity and community structure of the Kobuk River, Alaska, in response to climate change: 

Ecosphere, v. 2, no. 4.

61. Bickham, J.W., Patton, J.C., Minzenmayer, S., Moulton, L.L., and Gallaway, B.J., 1997, Identification of Arctic and Bering 

ciscoes in the Colville River Delta, Beaufort Sea coast, Alaska, in Reynolds, J., ed., Fish ecology in Arctic North America: 

Bethesda, Maryland, American Fisheries Society, Symposium no. 19, p. 224–228.

62. Reist, J.D., Maiers, L.D., Bodaly, R.A., Vuorinen, J.A., and Carmichael, T.J., 1998, The phylogeny of new- and oldworld coregonine fishes as revealed by sequence variation in a portion of the d-loop of mitochondrial DNA: Archiv für 

Hydrobiologie, Special Issues Advances in Limnology, v. 50, p. 323–339.

63. Reist, J.D., Vuorinen, J.A., and Bodily, R.A., 1992, Genetic and morphological identification of coregonid hybrid fishes 

from Arctic Canada: Polskie Archiwum Hydrobiologii, v. 39, no. 3/4, p. 551–562.
Bering Cisco 107

Bering Cisco (Coregonus laurettae)

Bean, 1881

Family Salmonidae 

Colloquial Name: Iñupiat: Qaaktaq, tipuk [1].

Ecological Role: Although data are lacking, this is a schooling 

species, and may be of some ecological importance in the 

nearshore of the U.S. Chukchi Sea and perhaps western part of 

U.S. Beaufort Sea. 

Physical Description/Attributes: Elongate, slightly compressed 

body with brownish to dark green back and silvery belly. There may 

be black dots with faint halos on the body, or white spots on the fins, or both. The anal, pectoral, and pelvic fins are pale. For 

specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 184) [2]. Swim bladder: Present [3]. 

Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea east to U.S. Beaufort Sea at Oliktok Point (just east of the Colville River). Elsewhere in Alaska, 

southward to the Kenai Peninsula, northern Gulf of Alaska. Worldwide, Chukotka Peninsula, eastern Siberia, Russia [2].

Relative Abundance: Patchily abundant (common on rare occasions) as far eastward as the Colville River, as well as such 

locations as the Barrow, Wainwright, and Kotzebue regions [7–9].

Bering Cisco (Coregonus laurettae). Photograph by 

R.J. Brown, U.S. Fish and Wildlife Service.

Geographic distribution of Bering Cisco (Coregonus laurettae) at sea within 2008–09 lease areas [4] based on 

review of published literature and specimens from historical and recent collections [5, 6]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Bering Cisco

Coregonus laurettae

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
108 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Mainly shallow nearshore waters, although taken as much as 20–30 km offshore in Yukon River plume [10, 11].

Habitats and Life History

Anadromous [12]. 

Eggs—Size: Specific size unknown. 2.3–3.0 mm for whitefish in general [12]. Time to hatching: Specific time 

unknown. 150–200 days for northern whitefish in general [12]. Habitat: Benthic, in gravel beds of fast-flowing 

rivers [11, 12]. 

Alevins (larvae)—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile

transformation: Unknown. Habitat: Pelagic, in freshwater rivers to nearshore estuaries [11]. 

Juveniles—Age and size: Minimum size unknown. Matures at about 4 years and 310 mm [11, 13]. Habitat:

Nearshore marine and brackish waters [11].

Adults—Age and size at first maturity: 4 years and as small as 310 mm (Yukon River) [11, 13]. Maximum age:

At least 13 years [12], and at least 8 years in Colville River area [9]. Maximum size: 48 cm [2]. Habitat: Pelagic,

mainly in marine and estuarine nearshore waters. Fast-flowing rivers for spawning [11]. 

Substrate—Sand and gravel for spawning [11, 12]. 

Physical/chemical—Temperature: Unknown. Salinity: Fresh to full seawater. The most marine-tolerant of all

coregonids within the study area range [13].

Behavior

Diel—Unknown. 

Seasonal—Eggs hatch in spring. Alevins (larvae) likely move downstream soon after and enter coastal waters 

where they spend their first years [11]. Upon maturity, adults migrate up river. Anadromous fish, probably on 

spawning runs, have been found at least 2,000 km [14] or perhaps as much as 2,150 km [15] upstream in the 

Yukon River. Spawning occurs in Yukon, Kuskokwim, and Susitna Rivers [11, 12, 16]. Yukon River spawning 

migrations are continuous throughout the summer with major pulses varying from year to year [12]. Spawning

occurs in autumn [11]. Juveniles and adults overwinter beneath ice of river deltas and other coastal waters [7,

11, 17]. 

Reproductive—Spawns annually [12]. Broadcast spawners over gravel beds in fast-flowing rivers [11]. Returns

to sea after spawning [11]. 

Schooling—Forms schools [11]. 

Feeding—Feeding likely occurs in nearshore waters, especially near river mouths and brackish estuaries [11].

Does not feed during spawning migrations [11]. 

Populations or Stocks

There have been no studies within the study area.

Benthic and reproductive distribution of Bering Cisco (Coregonus laurette).

3-2-8g_BeringCisco

Open ocean Shore

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

Adult and juvenile depths estimated.

Reported as shallow and nearshore 

Larvae depths estimated.

Reported as shallow and nearshore 

Coregonus laurette

Bering Cisco

Benthic distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning and eggs 

occur in freshwater
Bering Cisco 109

Reproduction

Mode—Gonochoristic, oviparous, iteroparous with external fertilization [11]. 

Spawning season—Autumn; early to mid-October in Yukon and Kuskokwim River drainages [11, 12]. 

Fecundity—In Yukon River, 20,210–34,166 orange, non-adhesive eggs [12].

Food and Feeding

Food items—Mysids as well as harpacticoid copepods, isopods, gammarid amphipods, crangonid shrimps, 

insects, and small fishes [18]. 

Trophic level—3.79 (standard error 0.59) [19].

Biological Interactions

Predators—Unknown. 

Competitors—Unknown.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [19]. 

Traditional and Cultural Importance

There are subsistence fisheries for Bering Cisco wherever the species is found. The species is most important 

near Wainwright [20] and Kotzebue Sound [7], although occasional anomalously large runs are known from the 

Colville River region [8]. Bering Cisco are taken during open water seasons and under the ice by gillnets and 

hook and line [20, 21]. This very oily species is most often roasted, salted, or frozen [7]. 

Commercial Fisheries

Currently, Bering Cisco are not commercially harvested. Historically, there has been no commercial fishery for 

this species [22] until 2008, when a fishery was initiated at the mouth of the Yukon River to supply a New York 

kosher market with smoked fish [12]. 

Potential Effects of Climate Change

Unknown. However, Durand and others (2011) estimate that, at least for anadromous fishes in sub-arctic rivers 

various biological shifts will be caused by the timing of spring ice break-up (and thus peak flow timing) and 

various permafrost processes that influence nutrient and sediment supply [23]

Areas for Future Research [B]

Little is known about the ecology and life history of this species in the study area. Research needs include: 

(1) depth, location, and timing of spawning; (2) size and age of fish at hatching and transformation; 

(3) preferred depth ranges and locations for juveniles and adults; (4) spawning season; (5) seasonal and 

ontogenetic movements; (6) population studies; (7) prey; and (8) predators. 
110 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Alt, K.T., 1973a, Contributions to the biology of the Bering cisco (Coregonus laurettae) in Alaska: Journal of the Fisheries 

Research Board of Canada, v. 30, no. 12, p. 1,885–1,888. [13]

Brown, R.J., Brown, C., Braem, N.M., Carter, W.K., III, Legere, N., and Slayton, L., 2012, Whitefish biology, distribution, 

and fisheries in the Yukon and Kuskokwim river drainages in Alaska—A synthesis of available information: U.S. Fish and 

Wildlife Service, Alaska Fisheries Data Series Number 2012-4, 316 p. [12]

Committee on the Status of Endangered Wildlife in Canada, 2004, COSEWIC assessment and update status report on the Bering 

cisco Coregonus laurettae in Canada: Ottawa, Canada, Committee on the Status of Endangered Wildlife in Canada, 19 p. [11]

Georgette, S., and Shiedt, A., 2005, Whitefish—Traditional ecological knowledge and subsistence fishing in the Kotzebue 

Sound, Alaska: Alaska Department of Fish and Game, Technical Report No. 290, 148 p. [7]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Bibliography

1. Craig, P.C., 1989, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers University 

of Alaska, no. 24, p. 131–152.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

6. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

7. Georgette, S., and Shiedt, A., 2005, Whitefish—Traditional ecological knowledge and subsistence fishing in the Kotzebue 

Sound, Alaska: Alaska Department of Fish and Game, Technical Report No. 290, 148 p.

8. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

9. Moulton, L.L., Owl Ridge Natural Resource Consultants, written commun., 2011.

10. Schmidt, D.R., Griffiths, W.B., Beaubien, D.K., and Herlugson, C.J., 1991, Movement of young-of-the-year Arctic ciscoes 

across the Beaufort Sea coast, 1985–1988: American Fisheries Society Symposium, no. 11, p. 132–144.

11. Committee on the Status of Endangered Wildlife in Canada, 2004, COSEWIC assessment and update status report on the 

Bering cisco Coregonus laurettae in Canada: Ottawa, Canada, Committee on the Status of Endangered Wildlife in Canada, 

19 p.

12. Brown, R.J., Brown, C., Braem, N.M., Carter, W.K., III, Legere, N., and Slayton, L., 2012, Whitefish biology, distribution, 

and fisheries in the Yukon and Kuskokwim river drainages in Alaska—A synthesis of available information: U.S. Fish and 

Wildlife Service, Alaska Fisheries Data Series Number 2012-4, 316 p.
Bering Cisco 111

13. Alt, K.T., 1973a, Contributions to the biology of the Bering cisco (Coregonus laurettae) in Alaska: Journal of the Fisheries 

Research Board of Canada, v. 30, no. 12, p. 1,885–1,888.

14. Brown, R.J., Bickford, N., and Severin, K., 2007, Otolith trace element chemistry as an indicator of anadromy in Yukon 

River drainage coregonine fishes: Transactions of the American Fisheries Society, v. 136, no. 3, p. 678–690.

15. DeGraaf, D.A., 1981, First Canadian record of Bering cisco (Coregonus laurettae) from the Yukon River at Dawson, Yukon 

Territory: The Canadian Field-Naturalist, v. 95, no. 1, p. 365.

16. Bickham, J.W., Patton, J.C., Minzenmayer, S., Moulton, L.L., and Gallaway, B.J., 1997, Identification of Arctic and Bering 

ciscoes in the Colville River Delta, Beaufort Sea coast, Alaska, in Reynolds, J., ed., Fish ecology in Arctic North America: 

Bethesda, Maryland, American Fisheries Society, Symposium no. 19, p. 224–228.

17. Craig, P.C., 1989, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.

18. Martin, D.J., Glass, D.R., Whitmus, C.J., Simenstad, C.A., Milward, D.A., Volk, E.C., Stevenson, M.L., Nunes, P., Savoie, 

M., and Grotefendt, R.A., 1986, Distribution, seasonal abundance, and feeding dependencies of juvenile salmon and 

non-salmonid fishes in the Yukon River Delta: Outer Continental Shelf Environmental Assessment Program, Reports of 

Principal Investigators, U.S. Department of Commerce and U.S. Department of the Interior, 388 p.

19. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

20. Nelson, R.K., 1981, Harvest of the sea—Coastal subsistence in modern Wainwright: North Slope Borough, 112 p.

21. Moulton, L.L., Field, L.J., and Kovalsky, R., 1991, Predictability in the catch of Arctic cisco in the Colville River, Alaska: 

American Fisheries Society Symposium no. 11, p. 145–156.

22. Alaska Department of Fish and Game, 2008, 2008 Lower Yukon whitefish fishery outlook, regulations, and management 

strategies: Alaska Department of Fish and Game, Division of Commercial Fisheries, news release, no. 51.

23. Durand, J.R., Lusardi, R.A., Nover, D.M., Suddeth, R.J., Carmona-Catot, G., Connell-Buck, C.R., and others, 2011, 

Environmental heterogeneity and community structure of the Kobuk River, Alaska, in response to climate change: 

Ecosphere, v. 2, no. 4.
112 Alaska Arctic Marine Fish Ecology Catalog

Broad Whitefish to Dolly Varden

Broad Whitefish (Coregonus nasus) 

(Pallas, 1776)

Family Salmonidae 

Colloquial Name: Iñupiat—Aanaakliq, Aanaaliq [1, 2]; Qalupiaq, 

Qausriluk, Qausiluk, Sigguilaq, Siyyuilaq [3, 57].

Ecological Role: This species rarely ventures into marine waters, 

preferring fresh and nearshore brackish conditions. It is one of 

the most prominent members of the coastal fish community and is 

important in seasonal food webs during ice-free periods. 

Physical Description/Attributes: Laterally compressed with a 

rounded to flat head, a broad maxilla, short and blunt gill rakers, and very thick scales. These scales develop prominent tubercles 

during the spawning season. Colors are olive-brown to nearly black on back, silvery, and white to yellowish belly. Fins of small 

fish are pale and in older individuals are dark [3, 4]. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg 

and others, 2002, p. 185) [4]. Swim bladder: Present [5]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [4]. Elsewhere in Alaska, found in Bering Sea drainages south to Kuskokwim Bay, 

southwestern Alaska. Worldwide, Arctic coasts from Siberia eastward to the Perry River, Nunavut, eastern Canada [4].

Broad Whitefish (Coregonus nasus). Photograph by R.J. 

Brown, U.S. Fish and Wildlife Service.
Broad Whitefish 113

Geographic distribution of Broad Whitefish (Coregonus nasus), at sea within Alaska Outer Continental Shelf Planning 

Areas [6] based on review of published literature and specimens from historical and recent collections [4, 7, 8].

tac14-5222_fig3-3-1m_Coregonus_nasus

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Broad Whitefish

Coregonus nasus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Relative Abundance: Common in parts of U.S. Beaufort Sea and southward in U.S. Chukchi Sea to at least the Kotzebue Sound

[3, 9]. Relatively rare nearshore between Smith Bay and Cape Halkett in Beaufort Sea [10], along ANWR and Yukon coasts 

[11–16], and along northeast Chukchi Sea coast [17], reflecting this species’ intolerance of cold and saline waters. Elsewhere, 

common eastward in Canadian Beaufort Sea to at least the Coppermine River [18]. 
114 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Shallow, nearshore waters [4, 19, 20].

Habitats and Life History

Exhibits a wide range of life history strategies. Anadromous, migrating from fresh to brackish waters for a few 

months in summer, to lacustrine, never leaving fresh water [21–24]. Migratory fish have been variously termed 

“facultative anadromous” [14], “semi-anadromous” [23], and “diadromous” [25]. Those entering U.S. Chukchi 

and Beaufort seas are primarily freshwater residents, only using the coastal zone as a migration corridor and 

an alternate feeding habitat under suitable (low-salinity) conditions [26]. This account focuses on those fish that 

migrate into the shallow waters of the Chukchi and Beaufort Seas. Lacustrine and riverine types are discussed 

elsewhere [23, 27–29]. The early life history of this species is best known for fish produced in the Mackenzie 

River watershed, however even here there are some uncertainties. 

Eggs—Size: 1.7–2.3 mm [30]. Time to hatching: Specific time unknown; 150–200 days for northern whitefish in 

general [31]. Spawning is in autumn and eggs hatch during spring [3, 27, 32, 33]. Habitat: Gravel beds in fastflowing freshwater rivers [27].

Fry—Size at hatching: Unknown. Days to juvenile transformation: Unknown. Habitat: Freshwater rivers to 

nearshore estuaries [14, 16, 27, 34]. 

Juveniles—Age and size: 0–5 years and 4.5–32.8 cm [35, 36]. Habitat: Primarily freshwater rivers and lakes 

to nearshore estuaries [14, 16, 34]. In the Mackenzie River, when young-of-the-year fish are flushed out of the 

river, they mainly are carried eastward along the Tuktoyaktuk Peninsula; some move into the outer or inner delta 

regions or eastward along the Yukon coast. Those reaching the Tuktoyaktuk Peninsula ascend rivers and spend up 

to 4 years in lakes. Larger, but still immature, fish may over winter in lakes or coastal waters, and use the coast, 

creeks, and lakes of the Tuktoyaktuk Peninsula as summer feeding grounds [12, 13, 16, 23, 27, 37]. Outcome 

of fish not transported along the Tuktoyaktuk Peninsula is unknown; although those carried westward along the 

Yukon coast may not survive due to its cold and highly saline waters [12, 13, 16, 23, 27, 37]. 

Adults—Age and size at first maturity: Highly variable throughout its range; from 3 to at least 21 years, mostly 

at 6–9 years (30–40 cm long) [13, 20, 22, 23, 27, 29, 37, 38]. Maximum age: Between 30 and 38 or more years 

[2, 27]. Maximum size: 83.5 cm [2]. Habitat: Pelagic, in fresh and nearshore brackish (rarely marine) waters [21, 

23, 24, 26, 28]. 

Substrate—Sand and gravel beds for spawning [27, 31]. 

Physical/chemical—Temperature: At least 0–16 °C [24]. Spawning occurs at about 0 °C [13]. Salinity: 0–30 

parts per thousand, but rare in higher salinity waters. Juveniles cope with salinities greater than 15–20 parts per 

thousand for only short periods, whereas larger fish are more tolerant of brackish conditions [20, 22, 26, 39].

Benthic and reproductive distribution of Broad Whitefish (Coregonus nasus). 

3-3-1g_BroadWhitefish

Open ocean Shore

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

Adult and juvenile depths estimated.

Reported as shallow and nearshore Larvae depths estimated.

Reported as nearshore estuaries 

Coregonus nasus

Broad Whitefish

Benthic distribution

Data from U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning and eggs 

occur in freshwater
Broad Whitefish 115

Behavior

Diel—Unknown. Seasonal—In spring eggs hatch under ice (perhaps in April and May) [23, 27] and young-ofthe-year are flushed out of the river into the river’s estuary during break-up [12, 13, 16, 23, 27, 37].

In June fish migrate into the smaller estuarine river deltas of the U.S. Beaufort Sea [19] [20], and generally do 

not migrate far [12, 40, 41]. For instance, Sagavanirktok River fish remain in the river delta until they are at 

least 3 years old [41]. In that population, the somewhat older fish are more tolerant of saline conditions and 

may migrate farther along coasts, typically moving between the Sagavanirktok and Colville Rivers [25]. At the 

extreme, a fish tagged in Prudhoe Bay was recaptured in Kaktovik Lagoon, 175 km to the east [42]. However, 

once mature, many adult fish also exhibit restricted movements [26]. At least in the Mackenzie River, once a fish 

has spawned it remains within the river’s influence for the rest of its life and never returns to its Tuktoyaktuk 

Peninsula nursery grounds [23]. Fish destined to spawn begin to enter rivers mostly in July and August [19, 

27, 38]. Mature fish preparing to spawn tend to move to staging areas in river mouths earlier than do mature, 

non-spawning individuals. Regardless of maturation state, by September all broad whitefish have returned to 

rivers, with spawning fish migrating earliest [27, 37]. Fish overwinter in a wide range of habitats. Depending 

on location, overwintering grounds include nearshore brackish waters, river deltas, deep pools in rivers, and a 

variety of lakes [16, 19, 23, 27, 43]. In Alaska, overwintering in nearshore waters is not known to occur. 

Reproductive—Spawning occurs in fresh water, likely in their natal streams [23]. In some river systems, fish 

may travel great distances to spawning sites. For instance, Yukon River fish may migrate at least 1,700 km [44]. 

In the Mackenzie River, fish first congregate in the river delta for a few months before moving upstream to the 

spawning grounds, perhaps waiting for water temperatures to be to near 0 °C before migrating [13, 23, 27]. In 

U.S. Chukchi and Beaufort Seas, spawning occurs in many rivers, from the Sagavanirktok River to at least the 

Meade River [12, 19], and in the Kobuk and Selawik Rivers in the Kotzebue region [3]. In most northern Alaskan 

rivers, spawning occurs in the lower parts of the waterways in deeper pools [19, 27], except the Colville River 

where fish may spawn a significant distance from the mouth [27]. With the possible exception of the Canning 

River, none of the “mountain” rivers or streams of the eastern Alaskan and Yukon coasts harbor spawning 

populations [12, 25, 45]. However, scattered non-spawning fish have been captured in a number of Canadian 

rivers west of the Mackenzie River [12]. Upon reaching maturity, spawning may occur annually or every other 

year [31]. Broad Whitefish are broadcast spawners that release gametes into the water column over gravel beds; 

eggs sink to the bottom after fertilization [27, 46]. 

Schooling—Unknown. 

Feeding—Opportunistic predators [27]. Adults undertaking spawning migrations only occasionally feed [47]. 

Mature individuals or current-year spawners spend the summer prior to spawning feeding in delta habitats, 

peninsula lakes, or brackish water environments [23]. In the Alaska Beaufort Sea, nearshore brackish habitats are 

annually colonized by a rich benthic invertebrate community that forms the base of the coastal food web.

Populations or Stocks

Information about population sizes of Broad Whitefish occurring in U.S. Chukchi and Beaufort Sea drainages 

does not exist. However, life history parameters have been developed for fish found in the Sagavanirktok and 

Colville Rivers. Genetic distinctions among fish from the Mackenzie River and its tributaries appear likely. 

Groups of fish may have both distinct spawning and overwintering grounds [16]. Fish of the Mackenzie River 

Basin also likely are genetically distinct from those living in other watersheds [32, 48–50]. Investigators have 

reported genetic differences in fish found in the Sagavanirktok and Colville Rivers. 

Reproduction

Mode—Gonochoristic, oviparous, and iteroparous with external fertilization [27]. 

Spawning season—Mainly in October and November, at freeze-up under the ice [3, 21, 23, 27]. 

Fecundity—10,070–117,687 pale-colored eggs [3, 33].

Food and Feeding

Food items—Zooplankton, for young-of-the-year; a wide range of invertebrates including chironomids and 

other insects, clams, amphipods, snails, polychaetes, oligochaetes, mysids, isopods, and plants for larger 

individuals [27, 28, 47, 51, 52]. 

Trophic level—3.28 (standard error 0.44) [53].
116 Alaska Arctic Marine Fish Ecology Catalog

Biological Interactions

Predators—Not well understood. Ringed seals are probably a major predator; brown bears in the Mackenzie 

River occasionally feed on them [54]. 

Competitors—Likely other opportunistic feeders, such as other whitefish species, Dolly Varden, Arctic Cisco, 

Least Cisco, Bering Cisco, Humpback Whitefish, Arctic Flounder, Fourhorn Sculpin, and Arctic Cod.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (K=0.10–0.30; t

m=7; tmax=15; Fecundity=10,000) [53].

Traditional and Cultural Importance

Among the most important food fishes in the Arctic, taken in large numbers in the Kotzebue region on the 

Chukchi Sea, and from about Barrow, Alaska, to the Coppermine River, Northwest Territories, Canada, in 

the Beaufort Sea [1–3, 18, 37, 55]. Depending on location, and apparently on local preferences, most Broad 

Whitefish are frozen and either aged or dried [2, 3, 55]. 

Commercial Fisheries

Currently, Broad Whitefish are not commercially harvested. Repeated efforts to create viable, large-scale 

commercial fisheries for this species have failed, although they were caught and sold as by-catch in the 

Arctic Cisco fishery on the Colville River [20, 39]. The Iñupiat currently harvest and sale in Barrow under the 

subsistence barter and trade category rather than as an actual commercial fishing operation.

Potential Effects of Climate Change

Unknown. Generally, Durand and others (2011) [60] predict that, at least for anadromous fishes in subarctic 

rivers, shifts in biology will be effected by spring ice break-up and resultant peak flows and surrounding 

permafrost processes: both of which affect the supply of nutrients and (or) sediment to the watershed of climate 

change on spring break-up intensity.

Areas for Future Research [A]

The Broad Whitefish is a dominant species in brackish waters of the Alaska Beaufort Sea and has been well 

studied in the central Beaufort Sea near Prudhoe Bay. Population parameters including condition factors, genetic 

information, and habitat use are reasonably well known for this area. Coastal monitoring of populations should 

be continued to track population growth, survival, and recruitment patterns and changes in the coastal fish 

assemblage as indicators of environmental change.

References Cited

Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum ReserveAlaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife Management, Barrow, 

Alaska, 94 p. [27]

Carothers, C., Cotton, S., and Moerlein, K., 2013, Subsistence use and knowledge of salmon in Barrow and Nuiqsut, Alaska: 

U.S. Department of the Interior, Bureau of Ocean Energy Management, OCS Study BOEM 2013-0015, 52 p. [57]

Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of Alaska, 

v. 24, p. 27–54. [14]

Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, Simpson 

Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649. [20]
Broad Whitefish 117

Georgette, S., and Shiedt, A., 2005, Whitefish—Traditional ecological knowledge and subsistence fishing in the Kotzebue 

Sound, Alaska: Alaska Department of Fish and Game, Technical Report No. 290, 148 p. [3]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [4]

Philo, L.M., George, J.C., and Moulton, L.L., 1993b, The occurrence and description of anadromous fish in the Dease Inlet/

Admiralty Bay, Alaska area, 1988–1990: Barrow, Alaska, Department of Wildlife Management, North Slope Borough and 

MJM Research, 150 p. [22]

Reist, J.D., and Chang-Kue, K.T.J., 1997, The life history and habitat usage of broad whitefish in the lower Mackenzie River 

Basin, in Tallman, R.F., and Reist, J.D., eds., The proceedings of the broad whitefish workshop—The biology, traditional 

knowledge and scientific management of broad whitefish (Coregonus nasus (Pallas)) in the Lower Mackenzie River: 

Canadian Technical Report of Fisheries and Aquatic Sciences, no. 2193, p. 63–84. [23]

Bibliography

1. Craig, P.C., 1989b, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers University 

of Alaska, no. 24, p. 131–152.

2. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

3. Georgette, S., and Shiedt, A., 2005, Whitefish—Traditional ecological knowledge and subsistence fishing in the Kotzebue 

Sound, Alaska: Alaska Department of Fish and Game, Technical Report No. 290, 148 p.

4. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

5. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

8. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

9. Ellis, D.V., 1962, Observations on the distribution and ecology of some Arctic fish: Arctic, v. 15, no. 3, p. 179–189.

10. Moulton, L.L., Owl Ridge Natural Resource Consultants, written commun., 2011.

11. Kendel, R.E., Johnston, R.A.C., Kozak, M.D., and Lobsiger, U., 1974, Movements, distribution, populations and food 

habits of fish in the western coastal Beaufort Sea: Department of the Environment, Canada, Interim Report of Beaufort Sea 

Project, Study B1, 64 p.

12. Griffiths, W.B., Fechhelm, R.G., Gallaway, B.J., and Fissel, D.B., 1988, Yukon fish and effects of causeway: For Indian and 

Northern Affairs Canada, LGL Ltd. and Arctic Science Ltd., 124 p. 

13. Reist, J.D., and Bond, W.A., 1988, Life history characteristics of migratory coregonids of the lower Mackenzie River, 

Northwest Territories, Canada: Finnish Fisheries Research, v. 9, p. 133–144.

14. Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.
118 Alaska Arctic Marine Fish Ecology Catalog

15. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

16. Sekerak, A.D., Stallard, N., and Griffiths, W.B., 1992, Distribution of fish and fish harvests in the nearshore Beaufort Sea 

and Mackenzie Delta during ice-covered periods, October–June: Environmental Studies Research Funds Report, LGS Ltd. 

No. 117, 524 p.

17. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

18. Gillman, D.V., and Kristofferson, A.H., 1984, Biological data on Pacific herring (Clupea harengus pallasi) from 

Tuktoyaktuk Harbour and the Liverpoor Bay area, Northwest Territories, 1981 to 1983: Winnipeg, Manitoba, Canada, 

Canadian Data Report of Fisheries and Aquatic Sciences, Department of Fisheries and Oceans, Western Region, no. 485, 

22 p.

19. Bendock, T.N., 1977, Beaufort Sea estuarine fishery study: Alaska Department of Fish and Game Annual Report, Contract 

#03-5-022-69, p. 670–729.

20. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

21. Craig, P.C., and Wells, J., 1980, Fisheries investigations in the Chandalar River region, northeast Alaska: Prepared for 

Northwest Alaskan Pipeline Company by LGL Ecological Research Associates, Fluor Northwest, Inc., Summary Report 

prepared for Northwest Alaskan Pipeline Company.

22. Philo, L.M., George, J.C., and Moulton, L.L., 1993b, The occurrence and description of anadromous fish in the Dease Inlet/

Admiralty Bay, Alaska area, 1988–1990: Barrow, Alaska, Department of Wildlife Management, North Slope Borough and 

MJM Research, 150 p.

23. Reist, J.D., and Chang-Kue, K.T.J., 1997, The life history and habitat usage of broad whitefish in the lower Mackenzie 

River Basin, in Tallman, R.F., and Reist, J.D., eds., The proceedings of the broad whitefish workshop—The biology, 

traditional knowledge and scientific management of broad whitefish (Coregonus nasus (Pallas)) in the Lower Mackenzie 

River: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 2193, p. 63–84.

24. Brown, R.J., 2004, A biological assessment of whitefish species harvested during the spring and fall in the Selawik River 

Delta Selawik National Wildlife Refuge, Alaska: Alaska Fisheries Technical Report no. 77.

25. Gallaway, B.J., and Fechhelm, R.G., 1997, Anadromous and amphidromous fishes, in Truett, J.C., and Johnson, S.R., eds., 

The natural history of an Arctic oil field: San Diego, Academic Press, p. 349–369.

26. Gallaway, B.J., Fechhelm, R.G., Griffiths, W.B., and Cole, J.G., 1997, Population dynamics of broad whitefish in the 

Prudhoe Bay region, Alaska: American Fisheries Society Symposium, no. 19, p. 194–207.

27. Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum 

Reserve, Alaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife 

Management, 94 p.

28. Philo, L.M., George, J.C., and Moulton, L.L., 1993a, The occurrence and description of anadromous and freshwater fish 

in Teshekpuk Lake, Alaska, 1990–1992: Barrow, Alaska, Department of Wildlife Management, North Slope Borough and 

MJM Research, 98 p.

29. Tallman, R.F., Abrahams, M.V., and Chudobiak, D.H., 2002, Migration and life history alternatives in a high latitude 

species, the broad whitefish, Coregonus nasus Pallas: Ecology of Freshwater Fish, v. 11, p. 101–111.

30. Riehl, R., 1980, Micropyle of some salmonins and coregonins: Environmental Biology of Fishes, v. 5, no. 1, p. 59–66.

31. Brown, R.J., Brown, C., Braem, N.M., Carter, W.K., III, Legere, N., and Slayton, L., 2012, Whitefish biology, distribution, 

and fisheries in the Yukon and Kuskokwim river drainages in Alaska—A synthesis of available information: U.S. Fish and 

Wildlife Service, Alaska Fisheries Data Series Number 2012-4, 316 p.
Broad Whitefish 119

32. Reist, J.D., and Chang-Kue, K.T.J., 1997, The life history and habitat usage of broad whitefish in the lower Mackenzie 

River Basin, in Tallman, R.F., and Reist, J.D., eds., The proceedings of the broad whitefish workshop—The biology, 

traditional knowledge and scientific management of broad whitefish (Coregonus nasus [Pallas]) in the Lower Mackenzie 

River: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 2193, p. 63–84.

33. Van Gerwen-Toyne, M., Walker-Larsen, J., and Tallman, R.F., 2008, Monitoring spawning populations of migratory 

coregonids in the Peel River, NT—The Peel River fish study 1998-2002: Canadian Manuscript Report of Fisheries and 

Aquatic Sciences no. 2851, 56 p.

34. Bond, W.A., and Erickson, R.N., 1997, Coastal migrations of Arctic ciscoes in the eastern Beaufort Sea, in Reynolds, J., ed., 

Fish ecology in Arctic North America, symposium 19: Bethesda, Maryland, American Fisheries Society, p. 155–164.

35. Bond, W.A., and Erickson, R.N., 1987, Fishery data from Phillips Bay, Yukon, 1985: Winnipeg, Manitoba, Canadian Data 

Report of Fisheries and Aquatic Sciences, Central and Artic Region, Department of Fisheries and Oceans, no. 635, 47 p.

36. Moulton, L.L., 1989, Recruitment of Arctic cisco (Coregonus autumnalis) into the Colville Delta, Alaska, in 1985: 

Biological Papers of the University of Alaska, no. 24, p. 107–111.

37. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, 

no. 1119, 90 p.

38. Alt, K.T., and Kogl, D.R., 1973, Notes on the whitefish of the Colville River, Alaska: Journal of the Fisheries Research 

Board of Canada, v. 30, no. 4, p. 554–556.

39. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

40. Craig, P.C., and McCart, P.J., 1975, Fish utilization of nearshore coastal waters between the Colville and Mackenzie Rivers 

with an emphasis on anadromous species, in Craig, P.C., ed., Fisheries Investigations in a coastal region of the Beaufort 

Sea: Calgary, Alberta, Canadian Arctic Gas Study Ltd. and Alaskan Arctic Gas Study Co., Biological Report Series 34, 

p. 172–219.

41. Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., and Colonell, J.M., 1996, The 1995 fish and oceanography 

study in Mikkelsen Bay, Alaska: Anchorage, Alaska, Prepared by LGL Alaska Research Associates and Woodward-Clyde 

Consultant for BP Exploration (Alaska) Inc., 102 p. plus apps.

42. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

43. Schmidt, D.R., Griffiths, W.B., and Martin, L.R., 1987, Importance of anadromous fish overwintering habitat in the 

Sagavanirktok River Delta, Alaska: Anchorage, Alaska, Report by Ecological Research Associates for Standard Alaska 

Production Company and North Slope Borough, 71 p.

44. Brown, R.J., Bickford, N., and Severin, K., 2007, Otolith trace element chemistry as an indicator of anadromy in Yukon 

River drainage coregonine fishes: Transactions of the American Fisheries Society, v. 136, no. 3, p. 678–690.

45. Bond, W.A., and Erickson, R.N., 1989, Summer studies of the nearshore fish community at Phillips Bay, Beaufort Sea coast, 

Yukon: Winnepeg, Manitoba, Canadian Technical Report of Fisheries and Aquatic Sciences, Central and Arctic Region, 

Department of Fisheries and Oceans, no. 1676, 102 p.

46. Carter, W.K., III, 2010, Life history and spawning movements of broad whitefish in the middle Yukon River: Fairbanks, 

University of Alaska, Master’s thesis, 63 p.

47. Hatfield, C.T., Stein, J.N., Falk, M.R., and Jessop, C.S., 1972, Fish resources of the Mackenzie River Valley: Winnepeg, 

Environment Canada Fisheries Service, Interim report 1, v. 1.

48. Tallman, R.F., 1997, Interpopulation variation in growth rates of broad whitefish: American Fisheries Society Symposium, 

no. 19, p. 184–193.
120 Alaska Arctic Marine Fish Ecology Catalog

49. Harris, L.N., and Taylor, E.B., 2010, Genetic population structure of broad whitefish, Coregonus nasus, from the Mackenzie 

River, Northwest Territories—Implications for subsistence fishery management: Canadian Journal of Fisheries and Aquatic 

Sciences, v. 67, no. 6, p. 905–918.

50. Harris, L.N., and Taylor, E.B., 2010, Pleistocene glaciations and contemporary genetic diversity in a Beringian fish, the 

broad whitefish, Coregonus nasus (Pallas)—Inferences from microsatellite DNA variation: Journal of Environmental 

Biology, v. 23, no. 1, p. 72–86.

51. Furniss, R.A., 1975, Inventory and cataloging of Arctic area waters: Alaska Department of Fish and Game, Federal Aid in 

Fish Restoration, Annual Report of Performance, Project F-9-7, Study G-1-1, v. 16, 47 p.

52. Fechhelm, R.G., Bryan, J.D., Griffiths, W.B., and Martin, L.R., 1997, Summer growth patterns of northern Dolly Varden 

(Salvelinus malma) smolts from the Prudhoe Bay region of Alaska: Canadian Journal of Fisheries and Aquatic Sciences, 

v. 54, no. 5, p. 1,103–1,110. 

53. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

54. Barker, O.E., and Derocher, A.E., 2009, Brown bear (Ursus arctos) predation of broad whitefish (Coregonus nasus) in the 

Mackenzie Delta region, Northwest Territories: Arctic, v. 62, p. 312–316.

55. Moulton, L.L., Field, L.J., and Kovalsky, R., 1991, Predictability in the catch of Arctic cisco in the Colville River, Alaska: 

American Fisheries Society Symposium no. 11, p. 145–156.

56. Durand, J.R., Lusardi, R.A., Nover, D.M., Suddeth, R.J., Carmona-Catot, G., Connell-Buck, C.R., and others, 2011, 

Environmental heterogeneity and community structure of the Kobuk River, Alaska, in response to climate change: 

Ecosphere, v. 2, no. 4.

57. Carothers, C., Cotton, S., and Moerlein, K., 2013, Subsistence use and knowledge of salmon in Barrow and Nuiqsut, 

Alaska: U.S. Department of the Interior, Bureau of Ocean Energy Management, OCS Study BOEM 2013-0015, 52 p.
Humpback Whitefish 121

Humpback Whitefish (Coregonus pidschian) 

(Gmelin, 1789)

Family Salmonidae 

Colloquial Name: Iñupiat—Piqutuuq, Pikuktuuq [1]; Ikkuiyiq, 

Iqalupiaq, Iqalutchiaq, Qaalgiq [2].

Note: American biologists have generally referred to 

anadromous and Alaska-dwelling individuals of this species as 

“humpback” whitefish (Coregonus pidschian). Anadromous fish 

in northern Canada usually have been called “lake” whitefish 

(C. clupeaformis), by Canadian researchers. Based on both 

morphological and genetic studies, McDermid and others (2005)

[3] determined that, in North America, individuals of this group could be assigned to one of three subspecies (1) Humpback 

Whitefish (Coregonus clupeaformis pidschian), (2) Mississippian Lake Whitefish (Coregonus clupeaformis clupeaformis), and 

(3) Alaska Whitefish (Coregonus clupeaformis nelsonii). In particular, fish referable to the Humpback Whitefish subspecies were 

found in waters from the Alaska Peninsula through the U.S. Chukchi and Beaufort Sea drainages, and eastward to at least the 

lower Mackenzie River. For geographic ranges of all three subspecies, see McDermid and others (2005) [3]. In this account, 

data are included only from populations living within those ranges.

Taxonomy: The correct species name for this complex is pidschian, because it is the oldest name, used by Gmelin in 1789 when 

naming and describing the species. The name clupeaformis is more recent, used by Mitchill in 1818. In zoological nomenclature 

the oldest name has priority.

Ecological Role: This species rarely ventures into marine waters, and prefers freshwaters and nearshore brackish waters. It 

likely is of considerable ecological importance only within this relatively proscribed area.

Physical Description/Attributes: Dark brown to dark blue back, silver sides, and white belly prominent nuchal hump [4]. Both 

sexes (more prominently in males) develop “nuptial tubercles” on head and scales [2]. For specific diagnostic characteristics, 

see Fishes of Alaska (Mecklenburg and others, 2002, p. 188) [4]. Swim bladder: Present [5]. Antifreeze glycoproteins in blood 

serum: Unknown.

Range: Coastal waters of U.S. Chukchi and Beaufort Seas. Elsewhere in Alaska, found in Bering Sea south to Bristol Bay [6]. 

Worldwide, along Arctic coasts from Siberia, Russia, west to Kara Sea, and eastward along Alaskan and Canadian coasts to 

Hudson Bay and New England (as C. clupeaformis of Canadian authors) [4, 7].

Humpback Whitefish (Coregonus pidschian). Photograph by 

R.J. Brown, U.S. Fish and Wildlife Service.
122 Alaska Arctic Marine Fish Ecology Catalog

Relative Abundance: Common in coastal waters of southeastern U.S. Chukchi Sea, but is uncommon in northeastern Chukchi 

Sea [8]. Common in coastal waters of U.S. Beaufort Sea, except is uncommon between about Smith Bay and Cape Halkett

[9] and off the Arctic National Wildlife Refuge [10, 11]. Elsewhere, fairly common in Canadian Beaufort Sea along the outer 

Mackenzie River Delta [12]. 

tac14-5222_fig3-3-2m_Coregonus_pidschian

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Humpback Whitefish

Coregonus pidschian

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Humpback Whitefish (Coregonus pidschian), at sea within Arctic Outer Continental 

Shelf Planning Areas [13] based on review of published literature and specimens from historical and recent 

collections [4, 14, 15]. 
Humpback Whitefish 123

Depth Range: Shallow, nearshore waters [16, 17].

Benthic and reproductive distribution of Humpback Whitefish (Coregonus pidschian).

3-3-2g_HumpbackWhitefish

Open ocean Shore No reproductive distribution in seawater

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

Adult and juvenile depths estimated. Reported

as shallow, nearshore, and brackish water

Coregonus pidschian

Humpback Whitefish

Benthic distribution

Data from U.S. Beaufort-Chukchi Seas

Open ocean Shore

Depth, in meters

Reproductive distribution

Anadromous. Spawning, 

eggs and larvae all 

occur in freshwater

Habitats and Life History

Populations are amphidromous and freshwater [18, 19]. Amphidromous fish tend to remain in estuaries, within 

the influence of freshwater drainages [12, 16, 17]. 

Eggs—Size: 1.2–1.6 mm [20] and 144–257 eggs per gram ovary weight [21]. Time to hatching: About 6 months, 

based on autumn spawning and late winter to spring hatching [22, 23]. Size of eggs: Unhatched. Habitat: 

Freshwater; gravel beds in shallow and moderately swift waters [24]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Freshwater [18]. 

Juveniles—Age and size: On average, to about 6 years and about 30.0 cm [16, 18, 23, 25, 26]. Habitat: Fresh 

and brackish waters [23, 26]; tend to remain in estuaries, within the influence of freshwater drainages [12, 

16, 17]. 

Adults—Age and size at first maturity: Primarily at 7–10 years [16, 18, 23, 25]. However, in some populations, 

maturation occurs at a later age. For example, in Dease Inlet/Admiralty Bay region, 50 percent of fish are 

mature at about 11 years and not all fish mature until perhaps 14 years [26, 27]. Most fish are mature between 

300 and 350 mm [23, 26, 28]. Maximum age: 37 years [23]. Maximum size: 54 cm [39]. Habitat: Pelagic, in 

fresh and nearshore brackish waters; tend to remain in estuaries, within the influence of freshwater drainages

[12, 16, 17]. 

Substrate—Gravel to sand and silt for spawning [24, 29]. 

Physical/chemical—Temperature: 0–16 °C [19]. Salinity: At least 0–28 parts per thousand [17]. 
124 Alaska Arctic Marine Fish Ecology Catalog

Behavior

Diel—Unknown. 

Seasonal—Young fish are carried downstream to lower parts of rivers and into brackish coastal waters [22, 23]. 

Smaller fish tend to stay in or near river mouths, whereas larger individuals are able to tolerate somewhat colder 

and more saline waters and venture somewhat farther along the coast [23, 26]. However, most fish tend to remain 

within the brackish water lens [17, 24]. In Mackenzie River area (including the Tuktoyaktuk Peninsula), juvenile 

fish may remain in coastal channels, lakes, and other quiet waters for several years before returning to the river 

[24]. Mature fish return to rivers beginning in June and this migration perhaps peaks in August [25, 30, 31]. 

Reproductive—Although spawning may occur within a few kilometers of river mouths [23], spawning 

migrations may be quite extensive (as far as 1,700 km from the mouth in the Yukon River) [32]. Spawning tends 

to occur in relatively slow- or moderate-moving rivers (that is, Colville River, Alaska, and Mackenzie River, 

Canada). In Beaufort Sea drainages, most spawning takes place between Barrow and the Sagavanirktok River

and in the Mackenzie River [18, 23, 33]. Although Humpback Whitefish have been captured in most of the rivers 

flowing into the Beaufort Sea, it is not known whether fish spawn in such fast-flowing rivers as the Canning 

River, Alaska, or if these are only seasonal visits. Similarly, in the Chukchi Sea, Humpback Whitefish have not 

been found in numerous rivers, including the Kokolik, Utukok, Kukpowruk, and Kuk, Alaska [23], although they 

do spawn in the Kobuk River [2]. Spawning occurs over gravel or sand, in shallow waters. Eggs are broadcast 

over the river floor and lodge in crevices [24]. After spawning, mature fish migrate down river and overwinter in 

brackish waters of river mouths and perhaps in fresh water [18, 24, 34]. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

At least some distinct populations may exist among drainages running into the Beaufort and Chukchi Seas [3].

Reproduction 

Mode—Gonochoristic, oviparous, and iteroparous with external fertilization [23]. 

Spawning season—Primarily September–October, but may extend into November and December [2, 22, 29]. At 

least some mature individuals spawn only every other year [19, 26]. 

Fecundity—5,000–122,000 eggs, with higher numbers in larger fish [21, 26, 27, 29]. 

Food and Feeding

Food items—Benthic and epibenthic prey including clams, crustaceans (for example, isopods, amphipods, 

copepods, and mysids), insects, snails, fishes, and plants [26, 28]. 

Trophic level— 3.2 (standard error 0.40) [35].

Biological Interactions

Predators—Unknown. 

Competitors—Likely other benthic and epibenthic predators including other whitefish species, sculpins, Arctic 

Flounder, and Dolly Varden.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (t

m=3–14; tmax=14; Fecundity=8,000) [35].
Humpback Whitefish 125

Traditional and Cultural Importance 

A very important part of several subsistence fisheries, particularly those in the Kotzebue Sound [2]. Humpback 

Whitefish also are taken in some numbers in the Barrow area and in the lower Colville River [36, 37]. In 

Kotzebue Sound, their ease of scaling and lower oil content makes them a preferred species for paniqtuq (dried 

fish) and also for the aged and frozen product (quaq). Most fish are taken in seines or gillnets, but many are 

captured in ditches (qargisat) that are dug to divert migrating fish [2].

Commercial Fisheries

Currently, Humpback Whitefish are not commercially harvested.

Potential Effects of Climate Change

Unknown. Generally, Durand and others (2011) [60] predict that, at least for anadromous fishes in subarctic 

rivers, shifts in biology will be effected by spring ice break-up and resultant peak flows and surrounding 

permafrost processes: both of which affect the supply of nutrients and (or) sediment to the watershed of climate 

change on spring break-up intensity.

Areas for Future Research [A]

Research needs for this species in the study area include: spawning behavior, spawning locations and early life 

histories, as well as studies on populations and predators. The vulnerability of the species to climate change 

effects on hydrology, timing of key life history events, and habitat use requires investigation. The projected 

expansion of brackish water habitats could greatly change this species occurrence in nearshore areas.

References Cited

Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum Reserve, 

Alaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife Management, 94 p. 

[23]

Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of Alaska, 

v. 24, p. 27–54. [18]

Evans, C.L., Reist, J.D., and Minns, C.K., 2002, Life history characteristics of freshwater fishes occurring in the Northwest 

Territories and Nunavut, with major emphasis on riverine habitat requirements: Canadian Manuscript Report of Fisheries and 

Aquatic Sciences, no. 2614, 169 p. [24]

Georgette, S., and Shiedt, A., 2005, Whitefish—Traditional ecological knowledge and subsistence fishing in the Kotzebue 

Sound, Alaska: Alaska Department of Fish and Game, Technical Report No. 290, 148 p. [2]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [4]

Philo, L.M., George, J.C., and Moulton, L.L., 1993a, The occurrence and description of anadromous fish in the Dease Inlet/

Admiralty Bay, Alaska area, 1988–1990: Barrow, Alaska, Department of Wildlife Management, North Slope Borough and 

MJM Research, 150 p. [26]

Bibliography

1. George, J.C., and Nageak, B.P., 1986, Observations on the Colville River subsistence fishery at Nuiqsut, Alaska for the 

period 4 July–1 November 1984: Alaska Department of Wildlife Manangement, 70 p.

2. Georgette, S., and Shiedt, A., 2005, Whitefish—Traditional ecological knowledge and subsistence fishing in the Kotzebue 

Sound, Alaska: Alaska Department of Fish and Game, Technical Report No. 290, 148 p.
126 Alaska Arctic Marine Fish Ecology Catalog

3. McDermid, J.I., Reist, J.D., and Bodaly, R.A., 2005, Phylogeography and postglacial dispersal of whitefish (Coregonus 

clupeaformis complex) in Northwestern North America: Advances in Limnology, v. 60, p. 91–109.

4. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

5. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

6. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

7. Hunter, J.G., Leach, S.T., McAllister, D.E., and Steigerwalk, M.B., 1984, A distributional atlas of records of the marine 

fishes of Arctic Canada in the National Museums of Canada and Arctic Biological Station: National Museums of Canada 

Syllogues, no. 52.

8. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

9. Moulton, L.L., Owl Ridge Natural Resource Consultants, written commun., 2011.

10. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

11. Wiswar, D.W., West, R.L., and Winkleman, W.N., 1995, Fisheries investigation in Oruktalik Lagoon, Arctic National 

Wildlife Refuge, Alaska, 1986: U.S. Fish and Wildlife Service, Alaska Fisheries Technical Report, no. 30, 38 p. 

12. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

13. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

14. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

15. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

16. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.

17. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

18. Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.

19. Brown, R.J., 2004, A biological assessment of whitefish species harvested during the spring and fall in the Selawik River 

Delta Selawik National Wildlife Refuge, Alaska: Alaska Fisheries Technical Report no. 77.

20. Riehl, R., 1980, Micropyle of some salmonins and coregonins: Environmental Biology of Fishes, v. 5, no. 1, p. 59–66.

21. Dupuis, A.W., and Sutton, T.M., 2011, Reproductive biology of female humpback whitefish Coregonus pidschian in the 

Chatanika River, Alaska: Journal of Applied Ichthyology, v. 27, p. 1,365–1,370.
Humpback Whitefish 127

22. Craig, P.C., and Wells, J., 1980, Fisheries investigations in the Chandalar River region, northeast Alaska: Prepared for 

Northwest Alaskan Pipeline Company by LGL Ecological Research Associates, Fluor Northwest, Inc., Summary Report 

prepared for Northwest Alaskan Pipeline Company.

23. Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum 

Reserve, Alaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife 

Management, 94 p.

24. Evans, C.L., Reist, J.D., and Minns, C.K., 2002, Life history characteristics of freshwater fishes occurring in the Northwest 

Territories and Nunavut, with major emphasis on riverine habitat requirements: Canadian Manuscript Report of Fisheries 

and Aquatic Sciences, no. 2614, 169 p.

25. Alt, K.T., and Kogl, D.R., 1973, Notes on the whitefish of the Colville River, Alaska: Journal of the Fisheries Research 

Board of Canada, v. 30, no. 4, p. 554–556.

26. Philo, L.M., George, J.C., and Moulton, L.L., 1993b, The occurrence and description of anadromous fish in the Dease Inlet/

Admiralty Bay, Alaska area, 1988–1990: Barrow, Alaska, Department of Wildlife Management, North Slope Borough and 

MJM Research, 150 p.

27. Moulton, L.L., Philo, L.M., and George, J.C., 1997, Some reproductive characteristics of least ciscoes and humpback 

whitefish in Dease Inlet, Alaska: American Fisheries Society Symposium, no. 19, p. 119–126.

28. Hatfield, C.T., Stein, J.N., Falk, M.R., and Jessop, C.S., 1972, Fish resources of the Mackenzie River Valley: Winnepeg, 

Environment Canada Fisheries Service, Interim report 1, v. 1.

29. Brown, R.J., Brown, C., Braem, N.M., Carter, W.K., III, Legere, N., and Slayton, L., 2012, Whitefish biology, distribution, 

and fisheries in the Yukon and Kuskokwim river drainages in Alaska—A synthesis of available information: U.S. Fish and 

Wildlife Service, Alaska Fisheries Data Series Number 2012-4, 316 p.

30. Jones, M.L., and Den Beste, J., 1977, Tuft Point and adjacent coastal areas fisheries projects: Calgary, Alberta, Canada, 

Aquatic Environments, Ltd., 152 p.

31. Gwich’in Renewable Resource Board, 2008, Fisheries: Gwich’in Renewable Resource Board, accessed November 12, 

2008, at http://www.grrb.nt.ca/.

32. Brown, R.J., Bickford, N., and Severin, K., 2007, Otolith trace element chemistry as an indicator of anadromy in Yukon 

River drainage coregonine fishes: Transactions of the American Fisheries Society, v. 136, no. 3, p. 678–690.

33. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

34. Schmidt, D.R., Griffiths, W.B., and Martin, L.R., 1987, Importance of anadromous fish overwintering habitat in the 

Sagavanirktok River Delta, Alaska: Anchorage, Alaska, Report by Ecological Research Associates for Standard Alaska 

Production Company and North Slope Borough, 71 p.

35. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

36. Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.

37. Moulton, L.L., Field, L.J., and Kovalsky, R., 1991, Predictability in the catch of Arctic cisco in the Colville River, Alaska: 

American Fisheries Society Symposium no. 11, p. 145–156.

38. Durand, J.R., Lusardi, R.A., Nover, D.M., Suddeth, R.J., Carmona-Catot, G., Connell-Buck, C.R., and others, 2011, 

Environmental heterogeneity and community structure of the Kobuk River, Alaska, in response to climate change: 

Ecosphere, v. 2, no. 4.

39. Brown, R.J., 2009, Distribution and demographics of whitefish species in the Upper Koyukuk River drainage, Alaska, with 

emphasis on seasonal migrations and important habitats of broad whitefish and humpback whitefish: U.S. Fish and Wildlife 

Service, Alaska Fisheries Technical Report No. 104.
128 Alaska Arctic Marine Fish Ecology Catalog

Least Cisco (Coregonus sardinella)

Valenciennes, 1848

Family Salmonidae 

Colloquial Name: Iñupiat—Iqalusaaq [1]; Anuutituuq, Qalusraaq, 

Qalutchiaq [2].

Ecological Role: This species rarely ventures into marine waters, 

preferring fresh and nearshore brackish conditions. Least Cisco are 

seasonally common in nearshore waters and are important in coastal 

food webs. 

Physical Description/Attributes: Brownish to dark green back and silvery belly; back and dorsal fin have dark spots and 

fish more than 15 cm long and have dusky or black pelvic fins. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 182) [3]. Swim bladder: Present [4]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas. Along U.S. Chukchi Sea, found at least in Utukok, Kokolik, Kuk, Kukpowruk, and 

Utukok Rivers [5] and rivers running into Kotzebue Sound [2]. In U.S. Beaufort Sea, found in most rivers of North Slope region, 

from Inaru to Canning River, but are not presumed to live in or enter the rivers between Canning River and Babbage River in 

Yukon Territory [6–10]. Elsewhere in Alaska, south to Bristol Bay in eastern Bering Sea [3, 11]. Worldwide, Siberia, Russia, 

west to the White Sea and east to Southampton Island, Hudson Bay, Canada, northeast to Viscount Melville Sound, Beaufort 

Sea [12]. 

Least Cisco (Coregonus sardinella). Photograph by 

R.J. Brown, U.S. Fish and Wildlife Service.
Least Cisco 129

Relative Abundance: Common in U.S. Chukchi and Beaufort Sea coastal waters north from at least Kotzebue Sound [2] but 

relatively uncommon along some of the northeastern Chukchi Sea coast and in U.S. Beaufort Sea between Smith Bay and Cape 

Halkett, and between Canning River–Camden Bay region and near Phillips Bay, reflecting this species’ tendency to avoid high 

salinity and cold marine waters [5, 6, 16–20]. Common in Canadian Beaufort Sea east to as close as the Tuktoyaktuk Peninsula, 

Canada [21]. 

tac14-5222fig3-3-3mCoregonussardinella

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Least Cisco

Coregonus sardinella

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Least Cisco (Coregonus sardinella), at sea within Arctic Outer Continental Shelf 

Planning Areas [13] based on review of published literature and specimens from historical and recent 

collections [3, 14, 15]. 
130 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Shallow, nearshore waters [5, 22–24].

Benthic and reproductive distribution of Least Cisco (Coregonus sardinella). 

Habitats and Life History

In Alaska, Least Cisco has three life history patterns: (1) amphidromous (feeding in brackish or marine systems 

during the summer), (2) exclusively freshwater, found in lakes, and (3) a resident dwarf form of freshwater 

variety. Amphidromous and freshwater forms can occur together [25]. Information in this account is about the 

amphidromous form. For information on freshwater residents, see [1, 5, 24, 26–28]. 

Eggs—Size: 0.9–2.0 mm for mature eggs [29, 30]. Time to hatching: Based on autumn spawning and late winter 

and spring hatching, about 6 months [5, 31]. Habitat: Shallow waters of rivers, in gravel and sand [5, 31–33]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Some fish as small as 25 mm 

migrate to sea [34]. Days to juvenile transformation: Unknown. Habitat: Hatches in freshwater rivers [5]. In 

Siberia, amphidromous fish move downstream to sea soon after hatching [35]. 

Juveniles—Age and size: 0 to at least 3 years and to at least 220 mm [24, 31, 36]. Habitat: Fresh and nearshore 

brackish (rarely marine waters) [5]. 

Adults—Age and size at first maturity: Highly variable, age ranges from 4 to 18 years [24, 31]. Most are mature 

by perhaps 6–10 years [5, 16, 21, 25, 29, 30]; size at least a few by about 220 mm and all by about 350 mm [24, 

36]. Maximum age: 28 years [24]. Maximum size: 47 cm [3]. Habitat: Fresh and nearshore brackish (rarely 

marine waters) [5]. 

Substrate—Gravel to sand in freshwater for spawning [5, 37]. 

Physical/chemical—Temperature: –1.7–16 °C [25, 29]. Salinity: 0–32 parts per thousand [25, 29], primarily 

20 parts per thousand or less [5]. Juveniles tend to remain in warmer and less saline waters, whereas older 

fishes are more tolerant of colder temperatures and increased salinity [5, 8, 16, 22–24].

3-3-3g_LeastCisco

Open ocean Shore

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

Adult and juvenile depths estimated. Reported 

as shallow, nearshore, brackish waters

Larvae depths estimated.

Reported as nearshore estuaries 

Coregonus sardinella

Least Cisco

Benthic distribution

Data from U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning and eggs 

occur in freshwater
Least Cisco 131

Behavior

Diel—Unknown. 

Seasonal—Fish enter U.S. Beaufort Sea following ice breakup in June and July, and begin to disperse and feed 

along the coast [21, 29, 30, 34, 38]. Young fish may remain in rivers for several years before entering the sea 

[16]. Due to higher temperature and lower salinity preferences, juveniles do not migrate as far as adults, who 

disperse farther along the coast [5, 8, 16, 22–24]. Most Least Cisco found along the North Slope originate from 

spawning and overwintering grounds in the Colville River [8]. Colville River fish migrate farther east in years 

of strong westerly winds. Similarly, Mackenzie River fish move westward to a greater extent in years with strong 

easterly winds [8, 39, 40]. Along the North Slope, larger fish can swim faster and reach distant feeding grounds 

before smaller individuals [41]. Given the short summer season, Least Cisco can travel substantial distances 

when conditions are favorable. For instance a fish tagged in Simpson Lagoon was recaptured off Barrow, 300 km 

to the west [29]. However, fish leaving the Colville River generally migrate eastward along the Arctic National 

Wildlife Refuge coast, because nearshore marine waters between Smith Bay and Cape Halkett may block 

westward movements [20, 40]. Return migrations to spawning and overwintering grounds begin between July 

and September [8, 29, 41]. Both juveniles and adults overwinter in brackish waters of river deltas (for example, 

Colville River, Alaska, and Mackenzie River, Canada, are the largest overwintering areas along the Beaufort 

Sea), open coast (for example, Tuktoyaktuk Harbor), and freshwater lakes of the Tuktoyaktuk Peninsula, Canada 

[21, 41, 42]. Except for the Sagavanirktok River (where some overwintering occurs in delta waters), Least Cisco 

do not seem to either spawn or overwinter in any waterway between the Colville and Mackenzie Rivers [41]. 

Reproductive—Spawning occurs under ice in the shallow waters of rivers over gravel and sand [5, 31–33]. 

During spawning, males and females move toward the surface, perpendicular to the current, while the eggs are 

released and fertilized [43]. Sea-run fish spawn in lower reaches of rivers or, as in the Yukon-Koyukuk River 

system, may migrate at least as much as 1,600 km upstream [44]. In U.S. Chukchi and Beaufort Sea drainages, 

spawning by the Least Cisco is known to occur in various rivers including the Kobuk River [2], Colville River

[45], and Mackenzie River and its tributaries [42]. Individuals have been variously reported to spawn annually 

[25], in alternate years, or perhaps for some fish, less often [5, 16, 24, 28]. After spawning, fish quickly migrate 

downstream to freshwater overwintering grounds [22]. 

Schooling—In coastal waters, forms schools that can maintain their integrity for months [29]. 

Feeding—Generalists, feeding throughout the water column with planktonic organisms dominating [28]. Feeds 

both during winter and during spawning migrations, although feeding intensity in both instances may be low [7, 

29, 33, 42, 46].

Populations or Stocks

Catch statistics from the Colville River and coastal monitoring in Prudhoe Bay provide useful life history 

information and indices of population abundance. Genetic research (phylogeography and population structure) is 

ongoing (University of Alaska, Fairbanks).

Reproduction

Mode—Gonochoristic, oviparous, and iteroparous with external fertilization [5]. 

Spawning season—September–November [2, 5]. 

Fecundity—10,505–100,939 eggs in North America [24, 47].

Food and Feeding

Food items—Primarily crustaceans (for example, copepods, mysids, isopods, and amphipods) and insects, along 

with some polychaetes, clams, snails, fishes, fish eggs, fish larvae, and plant material [5, 24, 28, 29, 48, 49]. 

Trophic level—3.24 (standard error 0.50) [50]. 
132 Alaska Arctic Marine Fish Ecology Catalog

Biological Interactions

Predators—Dolly Varden and Arctic Smelt [34, 42], ice seals, and various seabirds such as the red throated loon.

Competitors—Other generalist feeders including various whitefish species and Dolly Varden, Arctic Flounder, 

and Fourhorn Sculpin. Lease Cisco is an important prey for loons and other predators in the coastal Chukchi Sea.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (K=0.40; t

m=2–4; tmax=11; Fecundity=2,500) [50].

Traditional and Cultural Importance

Historically, Least Cisco have been of some importance in subsistence fisheries, particularly before the advent of 

snowmobiles, when local populations depended on dogs for transportation. At that time, this small species was 

widely caught and used for dog food. Currently, along many rivers, Least Cisco are only occasionally harvested 

for human and dog food [1, 25, 29, 51]. Subsistence fishermen in the Kotzebue Sound area often consume the 

eggs of freshly caught fish [2].

Commercial Fisheries

Currently, Least Cisco are not commercially harvested.

Potential Effects of Climate Change

Unknown. A general prediction is that, at least for anadromous fishes in subarctic rivers, shifts in biology will 

be effected by spring ice break-up and resultant peak flows and surrounding permafrost processes: both of 

which affect the supply of nutrients and (or) sediment to the watershed of climate change on spring break-up 

intensity [52].

Areas for Future Research [A]

Least Cisco populations are best studied in the western (for example, Dease Inlet) and central (Prudhoe Bay) 

regions of the southeastern Beaufort Sea. They are an important member of the coastal fish assemblage during 

ice-free months. Monitoring of key parameters relative to population growth, survival, and recruitment, relative 

abundance, and condition, should be continued at key reference locations. 

References Cited

Brown, R.J., 2004, A biological assessment of whitefish species harvested during the spring and fall in the Selawik River Delta 

Selawik National Wildlife Refuge, Alaska: Alaska Fisheries Technical Report no. 77. [25]

Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum Reserve, 

Alaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife Management, 

94 p. [5]

Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, Simpson 

Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649. [29]

Georgette, S., and Shiedt, A., 2005, Whitefish—Traditional ecological knowledge and subsistence fishing in the Kotzebue 

Sound, Alaska: Alaska Department of Fish and Game, Technical Report No. 290, 148 p. [2]
Least Cisco 133

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Philo, L.M., George, J.C., and Moulton, L.L., 1993b, The occurrence and description of anadromous fish in the Dease Inlet/

Admiralty Bay, Alaska area, 1988–1990: Barrow, Alaska, Department of Wildlife Management, North Slope Borough and 

MJM Research, 150 p. [24]

Bibliography

1. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

2. Georgette, S., and Shiedt, A., 2005, Whitefish—Traditional ecological knowledge and subsistence fishing in the Kotzebue 

Sound, Alaska: Alaska Department of Fish and Game, Technical Report No. 290, 148 p.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

5. Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum 

Reserve, Alaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife 

Management, 94 p.

6. Kendel, R.E., Johnston, R.A.C., Kozak, M.D., and Lobsiger, U., 1974, Movements, distribution, populations and food 

habits of fish in the western coastal Beaufort Sea: Department of the Environment, Canada, Interim Report of Beaufort Sea 

Project, Study B1, 64 p.

7. Schmidt, D.R., Griffiths, W.B., and Martin, L.R., 1987, Importance of anadromous fish overwintering habitat in the 

Sagavanirktok River Delta, Alaska: Anchorage, Alaska, Report by Ecological Research Associates for Standard Alaska 

Production Company and North Slope Borough, 71 p.

8. Griffiths, W.B., Fechhelm, R.G., Gallaway, B.J., and Fissel, D.B., 1988, Yukon fish and effects of causeway: For Indian and 

Northern Affairs Canada, LGL Ltd. and Arctic Science Ltd., 124 p. 

9. Moulton, L.L., and George, J.C., 1997, Freshwater fishes in the Arctic oil-field region and coastal plain of Alaska, in Truett, 

J.C., and Johnson, J.A., eds., The natural history of an Arctic oil field: San Diego, California, Academic Press, p. 327–348.

10. Wiswar, D.W., and Frugé, D.J., 2006, Fisheries investigations in western Camden Bay, Arctic National Wildlife Refuge, 

Alaska, 1987: Alaska Fisheries Data Series, U.S. Fish and Wildlife Service, 2006-10, 49 p.

11. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

12. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

13. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

14. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

15. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.
134 Alaska Arctic Marine Fish Ecology Catalog

16. Bendock, T.N., 1977, Beaufort Sea estuarine fishery study: Alaska Department of Fish and Game Annual Report, Contract 

#03-5-022-69, p. 670–729.

17. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

18. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

19. Wiswar, D.W., West, R.L., and Winkleman, W.N., 1995, Fisheries investigation in Oruktalik Lagoon, Arctic National 

Wildlife Refuge, Alaska, 1986: U.S. Fish and Wildlife Service, Alaska Fisheries Technical Report, no. 30, 38 p. 

20. Moulton, L.L., Owl Ridge Natural Resource Consultants, written commun., 2011.

21. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

22. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.

23. Schmidt, D.R., Griffiths, W.B., Beaubien, D.K., and Herlugson, C.J., 1991, Movement of young-of-the-year Arctic ciscoes 

across the Beaufort Sea coast, 1985–1988: American Fisheries Society Symposium, no. 11, p. 132–144.

24. Philo, L.M., George, J.C., and Moulton, L.L., 1993b, The occurrence and description of anadromous fish in the Dease Inlet/

Admiralty Bay, Alaska area, 1988–1990: Barrow, Alaska, Department of Wildlife Management, North Slope Borough and 

MJM Research, 150 p.

25. Brown, R.J., 2004, A biological assessment of whitefish species harvested during the spring and fall in the Selawik River 

Delta Selawik National Wildlife Refuge, Alaska: Alaska Fisheries Technical Report no. 77.

26. Cohen, D.M., 1954, Age and growth studies on two species of whitefishes from Point Barrow, Alaska: Stanford Ichthyology 

Bulletin, v. 4, p. 168–187.

27. Wohlschlag, D.E., 1952, Growth peculiarities of the cisco, Coregonus sardinella (Valenciennes), in the vicinity of Point 

Barrow, Alaska: Stanford Ichthyology Bulletin, v. 4, p. 189–209.

28. Mann, G.J., 1974, Life history types of the least cisco (Coregonus sardinella, Valenciennes) in the Yukon Territory, North 

Slope and eastern Mackenzie River Delta drainages, in McCart, P.J., ed., Life histories of three species of freshwater fishes 

in Beaufort Sea drainages, Yukon Territory: Arctic Gas Biological Report Series, v. 18, no. 3, 132 p.

29. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

30. Bond, W.A., and Erickson, R.N., 1987, Fishery data from Phillips Bay, Yukon, 1985: Winnipeg, Manitoba, Canadian Data 

Report of Fisheries and Aquatic Sciences, Central and Artic Region, Department of Fisheries and Oceans, no. 635, 47 p.

31. Reist, J.D., and Bond, W.A., 1988, Life history characteristics of migratory coregonids of the lower Mackenzie River, 

Northwest Territories, Canada: Finnish Fisheries Research, v. 9, p. 133–144.

32. Berg, L.S., 1948, Freshwater fishes of the USSR and adjacent countries, volume 1 (4th ed.): Moscow, Academy of Sciences 

of the USSR Zoological Institute, 466 p. [Translated from Russian by Israel Program for Science Translation, Jerusalem, 

505 p.]

33. Hatfield, C.T., Stein, J.N., Falk, M.R., and Jessop, C.S., 1972, Fish resources of the Mackenzie River Valley: Winnepeg, 

Environment Canada Fisheries Service, Interim report 1, v. 1.

34. Bond, W.A., and Erickson, R.N., 1989, Summer studies of the nearshore fish community at Phillips Bay, Beaufort Sea coast, 

Yukon: Winnepeg, Manitoba, Canadian Technical Report of Fisheries and Aquatic Sciences, Central and Arctic Region, 

Department of Fisheries and Oceans, no. 1676, 102 p.

35. Scott, W.B., and Crossman, E.J., 1973, Freshwater fishes of Canada: Fisheries Research Board of Canada, Bulletin 184, 

966 p.
Least Cisco 135

36. Alt, K.T., and Kogl, D.R., 1973, Notes on the whitefish of the Colville River, Alaska: Journal of the Fisheries Research 

Board of Canada, v. 30, no. 4, p. 554–556.

37. Brown, R.J., Brown, C., Braem, N.M., Carter, W.K., III, Legere, N., and Slayton, L., 2012, Whitefish biology, distribution, 

and fisheries in the Yukon and Kuskokwim river drainages in Alaska—A synthesis of available information: U.S. Fish and 

Wildlife Service, Alaska Fisheries Data Series Number 2012-4, 316 p.

38. Gallaway, B.J., Fechhelm, R.G., and Gazey, W.J., 1997, Estimating abundance of Arctic anadromous fish in the coastal 

Beaufort Sea during summer—Is an open population model necessary?: American Fisheries Society Symposium, no. 19, 

p. 274–286.

39. Fechhelm, R.G., Baker, J.S., Griffiths, W.B., and Schmidt, D.R., 1989, Localized movement patterns of least cisco 

(Coregonus sardinella) and Arctic cisco (C. autumnalis) in the vicinity of a solid-fill causeway: Biological Papers of the 

University of Alaska, no. 24, p. 75–106.

40. Gallaway, B.J., and Fechhelm, R.G., 1997, Anadromous and amphidromous fishes, in Truett, J.C., and Johnson, S.R., eds., 

The natural history of an Arctic oil field: San Diego, Academic Press, p. 349–369.

41. Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., and Colonell, J.M., 1996, The 1995 fish and oceanography 

study in Mikkelsen Bay, Alaska: Anchorage, Alaska, Prepared by LGL Alaska Research Associates and Woodward-Clyde 

Consultant for BP Exploration (Alaska) Inc., 102 p. plus apps.

42. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, 

no. 1119, 90 p.

43. Morrow, J.E., 1980, The freshwater fishes of Alaska: Anchorage, Alaska Northwest Publishing Company, 248 p.

44. Brown, R.J., Bickford, N., and Severin, K., 2007, Otolith trace element chemistry as an indicator of anadromy in Yukon 

River drainage coregonine fishes: Transactions of the American Fisheries Society, v. 136, no. 3, p. 678–690.

45. Craig, P.C., and McCart, P.J., 1975, Fish utilization of nearshore coastal waters between the Colville and Mackenzie Rivers 

with an emphasis on anadromous species, in Craig, P.C., ed., Fisheries Investigations in a coastal region of the Beaufort 

Sea: Calgary, Alberta, Canadian Arctic Gas Study Ltd. and Alaskan Arctic Gas Study Co., Biological Report Series 34, 

p. 172–219.

46. Mann, G.J., 1975, Winter fisheries survey across the Mackenzie Delta, in Craig, P.C., ed., Fisheries investigations in a 

coastal region of the Beaufort Sea: Arctic Gas Biological Report Series, v. 34.

47. Van Gerwen-Toyne, M., Walker-Larsen, J., and Tallman, R.F., 2008, Monitoring spawning populations of migratory 

coregonids in the Peel River, NT—The Peel River fish study 1998-2002: Canadian Manuscript Report of Fisheries and 

Aquatic Sciences no. 2851, 56 p.

48. Furniss, R.A., 1975, Inventory and cataloging of Arctic area waters: Alaska Department of Fish and Game, Federal Aid in 

Fish Restoration, Annual Report of Performance, Project F-9-7, Study G-1-1, v. 16, 47 p.

49. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.

50. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

51. George, J.C., and Kovalsky, R., 1986, Observations on the Kupigruak Channel (Colville River) subsistence fishery, October 

1985: Alaska Department of Wildlife Management, 66 p.

52. Durand, J.R., Lusardi, R.A., Nover, D.M., Suddeth, R.J., Carmona-Catot, G., Connell-Buck, C.R., and others, 2011, 

Environmental heterogeneity and community structure of the Kobuk River, Alaska, in response to climate change: 

Ecosphere, v. 2, no. 4.
136 Alaska Arctic Marine Fish Ecology Catalog

Pink Salmon (Oncorhynchus gorbuscha) 

(Walbaum, 1792)

Family Salmonidae 

Colloquial Name: Iñupiat—Amaqtuuq [1].

Ecological Role: The seeming relatively recent increase in 

abundance of this species north and eastward to at least the Barrow 

area implies that this species is becoming of some ecological 

importance in coastal waters of the U.S. Chukchi Sea. 

Physical Description/Attributes: At sea, metallic-blue to blue-green on back, silvery on sides, and white on belly with black, 

oval spots on the backs and upper sides and on both lobes of the caudal fins. For specific diagnostic characteristics, see Fishes 

of Alaska (Mecklenburg and others, 2002, p. 205) [2]. Swim bladder: Present [3]. Antifreeze glycoproteins in blood serum: 

Unknown.

Range: U.S. Chukchi and Beaufort Seas [4, 5]. North Korea and Japan and northern Siberia to the Beaufort Sea coast of Alaska 

and western Canada, eastward to Sachs Harbor, Banks Island, [5] and south to La Jolla, California [6].

Relative Abundance: Small runs occur in numerous rivers and streams running into the Chukchi Sea to about Barrow and 

perhaps in the Colville River [4, 10]; common in the Chukchi Sea (the most abundant of the salmon species) and, in the Beaufort 

Sea, at least as far east as Simpson Lagoon and in smaller numbers along the Alaska National Wildlife Refuge [4, 11–14]. 

Pink Salmon (Oncorhynchus gorbuscha), 211 mm, 

southeastern Chukchi Sea, 2007. Photograph by C.W. 

Mecklenburg, Point Stephens Research.

Geographic distribution of Pink Salmon (Oncorhynchus gorbuscha), at sea within Arctic Outer Continental 

Shelf Planning Areas [7] based on review of published literature and specimens from historical and recent 

collections [2, 8, 9]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Pink Salmon

Oncorhynchus gorbuscha

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Pink Salmon 137

Depth Range: Epipelagic, coastal and offshore, from surface to at least 74 m [15], usually from 0 to 10 m [16]. Young juveniles 

are neritic, occurring from nearshore waters as shallow as a few centimeters deep to the edge of the continental shelf as they 

mature [17].

Habitats and Life History

Anadromous. 

Eggs—Size: 4.0–7.9 mm [17]. Time to hatching: 5–8 months [17]. Length at hatching: About 19–24 mm [18]. 

Habitat: Benthic, buried under gravel in shallow, relatively fast-flowing waters of streams and rivers, often near 

the ocean [16, 17, 19, 20]. 

Alevins (larvae)—Size: 19–28 mm [16, 18]. Habitat: Under gravel for a few weeks to several months in shallow, 

relatively fast-flowing waters of streams and rivers, often near the ocean [16, 17, 19, 20]. 

Fry—Size at emergence: About 28–34 mm [18]. Time as fry: Relatively short, as little as a few days [17]. 

Habitat: Benthopelagic in shallow waters of streams and rivers [16, 17]. 

Smolt and ocean phase—Age and size at smoltification: as young as a few days and 3.0–45.0 cm [16, 17]. 

Habitat: Epipelagic in ocean, over various habitats and often far from shore [16, 17, 20].

Adults— Age and size at first maturity: On average, 1.5 years and 45–55 cm [17]. Maximum age: 2 years 

[16]. Maximum size: 76 cm [2]. Habitat: Mature adults return to freshwater rivers and streams to spawn [17]. 

Substrate—Over various substrates while in ocean [16]. Gravel for spawning [17]. 

Physical/chemical—Temperature: In ocean, at least 3–15 °C, perhaps primarily at 4–11 °C [21]. Salinity: 

Tolerates fresh to marine waters, depending on life stage [17].

Behavior

Diel—Migrations of fry to sea take place at night [17]. 

Seasonal—Unknown in U.S. Chukchi and Beaufort Seas, although it is speculated that juveniles enter the 

northeastern Chukchi Sea during the late spring thaw (June to early July) [11]. Vertical distribution at sea may 

change with season, with fish remaining in deep water during the night early in the year and then moving to 

shallow water later in the year [17]. 

Reproductive—Spawning occurs in freshwater streams and rivers, often very close to the ocean. Females 

prepare a gravel nest and lay their eggs, which are fertilized by the males. All adults die after spawning [16]. 

Schooling—Newly emerged fry may or may not school; fry do school when they reach the lower reaches of 

rivers [17]. Juveniles school in nearshore waters during early phases of their seaward migration [17]. Schooling 

behavior during ocean phases of migration may be less structured and include intermingling with other 

stocks [17]. 

Feeding—At sea, may vertically migrate (ascending during the night) when following prey [17]. Juveniles feed 

primarily during the day with peak feeding at dusk [17].

Benthic and reproductive distribution of Pink Salmon (Oncorhynchus gorbuscha). 

Open ocean Shore

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

No reproductive distribution in seawater

Smolts in nearshore 

estuaries

Juveniles and pre-spawning adults

Some fish to 74 m

Oncorhynchus gorbuscha

Pink Salmon

Benthic distribution

Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning, eggs, alevins

and fry all occur in freshwater
138 Alaska Arctic Marine Fish Ecology Catalog

Populations or Stocks

There have been no studies in the U.S. Chukchi and Beaufort Seas. 

Reproduction

Mode—Gonochoristic, oviparous, and semelparous with external fertilization [22].

Spawning season—July–October and usually earlier in more northern waters [17, 23]. 

Fecundity—854–1,549, Simpson Lagoon [4].

Food and Feeding

Food items—In the U.S. Chukchi Sea, fishes, planktonic crustaceans (for example, mysids, calanoid copepods, 

and gammarid amphipods), fish eggs, and fish larvae [11, 12]. 

Trophic level— 4.19 (standard error 0.71) [24].

Biological Interactions

Predators—Dolly Varden, Arctic Smelt, Starry Flounder, Walleye Pollock, spotted seal, beluga whale, and orca 

[25–29]. 

Competitors—Likely other salmon species, as well as such gadids as Arctic Cod. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (t

m=2; tmax=3; Fecundity=800) [24].

Traditional and Cultural Importance 

Taken in subsistence fisheries in the U.S. Chukchi Sea [10, 30, 31] and are caught in relatively small numbers in 

subsistence fisheries along the U.S. Beaufort Sea at least are far east as the Colville River [32].

Commercial Fisheries

Pink Salmon are captured in small numbers in salmon fisheries for Chum Salmon in Kotzebue Sound.

Potential Effects of Climate Change

Warming temperatures likely will allow this species to become more abundant in the U.S. Chukchi and Beaufort 

Seas. Whether this species will be able to colonize additional freshwater habitats successfully on Alaska’s North 

Slope is hypothesized, but is not known.
Pink Salmon 139

Areas for Future Research [A]

Little is known about the ecology and life history of this species in the study area. Human use of this species in 

coastal villages seems to be increasing. Research needs include: preferred habitats (including depth ranges for 

juveniles and adults), spawning season, seasonal and ontogenetic movements, population studies, and predators. 

The success of colonization east of Point Hope will depend on the presence of suitable spawning habitats and 

thermal regimes for developing embryos. The timing of seaward migrations and quality of nearshore marine 

environments will be critical to population viability. The potential for colonization of perennial springs in the 

eastern Beaufort Sea and other prospective spawning areas in the Colville and Sagavanirktok Rivers should be 

examined in habitat-based monitoring. Salmon catches at Point Barrow and other villages should be monitored 

and biological investigations of salmon interactions with traditional fish foods, such as Arctic and Least cisco and 

Dolly Varden, should be undertaken.

References Cited

Craig, P.C., and Haldorson, L.J., 1986, Pacific salmon in the North American Arctic: Arctic, v. 39, no. 1, p. 2–7. [4]

Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates in 

west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments Division, 

ELMR Report no. 8, 327 p. [16]

Heard, W.R., 1991, Life history of pink salmon (Oncorhynchus gorbuscha), in Groot, C., and Margolis, L., eds., Pacific salmon 

life histories: Vancouver, British Columbia, UBC Press, p. 121–230. [17]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Bibliography

1. Carothers, C., 2010, Subsistence use and knowledge of Beaufort salmon populations: 2010 Annual Report, Cooperative 

Agreement M09AC15378.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

4. Craig, P.C., and Haldorson, L.J., 1986, Pacific salmon in the North American Arctic: Arctic, v. 39, no. 1, p. 2–7. 

5. Stephenson, S.A., 2006, A review of the occurrence of Pacific Salmon (Oncorhynchus spp.) in the Canadian Western Arctic: 

Arctic, v. 59, no. 1, p. 37–46.

6. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

7. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

8. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

9. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.
140 Alaska Arctic Marine Fish Ecology Catalog

10. Craig, P.C., 1989b, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers University 

of Alaska, no. 24, p. 131–152.

11. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

12. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

13. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

14. Irvine, J.R., Macdonald, R.W., Brown, R.J., Godbout, L., Reist, J.D., and Carmack, E.C., 2009, Salmon in the Arctic and 

how they avoid lethal low temperatures: North Pacific Anadromous Fish Commission, Bulletin no. 5, p. 39–50.

15. Walker, R.V., Sviridov, V.V., Urawa, S., and Azumaya, T., 2007, Spatio-temporal variation in vertical distributions of Pacific 

salmon in the ocean: North Pacific Anadromous Fish Commission, Bulletin no. 4, p. 193–201.

16. Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates 

in west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments 

Division, ELMR Report no. 8, 327 p.

17. Heard, W.R., 1991, Life history of pink salmon (Oncorhynchus gorbuscha), in Groot, C., and Margolis, L., eds., Pacific 

salmon life histories: Vancouver, British Columbia, UBC Press, p. 121–230.

18. Beacham, T.D., and Murray, C.B., 1990, Temperature, egg size, and development of embryos and alevins of five species of 

Pacific salmon—A comparative analysis: Transactions of the American Fisheries Society, v. 119, no. 6, p. 927–945.

19. Craig, P.C., and Wells, J., 1980, Fisheries investigations in the Chandalar River region, northeast Alaska: Prepared for 

Northwest Alaskan Pipeline Company by LGL Ecological Research Associates, Fluor Northwest, Inc., Summary Report 

prepared for Northwest Alaskan Pipeline Company.

20. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

21. Brodeur, R.D., 1988, Zoogeography and trophic ecology of the dominant epipelagic fishes in the northern North Pacific: 

University of Tokyo, Bulletin of the Ocean Research Institute, v. 26, no. 2, p. 1–27.

22. Moyle, P.B., 2002, Inland fishes of California: Berkeley, University of California Press, 517 p.

23. Bonar, S.A., Pauley, G.B., and Thomas, G.L., 1989, Species profiles—Life histories and environmental requirements of 

coastal fishes and invertebrates (Pacific Northwest)—Pink salmon: U.S. Fish and Wildlife Service, Biological Report no. 82 

(11.88), TR EL-82-4, 17 p.

24. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

25. Armstrong, R.H., and Winslow, P.C., 1968, An incidence of Walleye Pollock feeding on salmon young: Transactions of the 

American Fisheries Society, v. 97, no. 2, p. 202–203.

26. Dobrynina, M.V., Gorshkov, S.A., and Kinas, N.M., 1989, Effect of density of juvenile pink salmon, Oncorhynchus 

gorbuscha, on their vulnerability to predators in the Utka River (Kamchatka): Journal of Ichthyology, v. 29, no. 1, 

p. 148–155.

27. Rugh, D.J., Shelden, K.E.W., and Withrow, D.E., 1997, Spotted seals, Phoca largha, in Alaska: Marine Fisheries Review, 

v. 59, no. 1, p. 1–18.

28. Huntington, H.P., 2000, Traditional knowledge of the ecology of belugas, Delphinapterus leucas, in Cook Inlet, Alaska: 

Marine Fisheries Review, v. 62, p. 134–140.
Pink Salmon 141

29. Ford, J.K.B., and Ellis, G.M., 2006, Selective foraging by fish-eating killer whales Orcinus orca in British Columbia: 

Marine Ecology Progress Series v. 316, p. 185–199.

30. Braund, S.R., and Burnham, D.C., 1984, Subsistence economics and marine resource use patterns, in Truett, J.C., ed., The 

Barrow Arch environment and possible consequences of planned offshore oil and gas development: Outer Continental Shelf 

Environmental Assessment Program, National Oceanic and Atmospheric Administration/Ocean Assessment Division, 

p. 167–215.

31. Menard, J., 2008, 2008 Kotzebue Sound salmon fisheries management plan: Alaska Department of Fish and Game, 

Regional Information Report No. 3A08-02.

32. George, J.C., and Nageak, B.P., 1986, Observations on the Colville River subsistence fishery at Nuiqsut, Alaska for the 

period 4 July–1 November 1984: Alaska Department of Wildlife Manangement, 70 p.
142 Alaska Arctic Marine Fish Ecology Catalog

Chum Salmon (Oncorhynchus keta) 

(Walbaum, 1792)

Family Salmonidae 

Colloquial Name: Iñupiat—Iqalugruaq [1].

Ecological Role: The abundance of this species north and eastward 

to at least the Barrow area implies that this species is likely of 

some ecological importance. Especially important member of the 

nearshore marine in Kotzebue Sound.

Physical Description/Attributes: At sea, Chum Salmon have dark 

blue backs with silvery sides and bellies and no distinct black spots, 

fewer than 30 gill rakers, and gill rakers that are smooth, fairly 

short, stout, and widely spaced. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, 

p. 208) [2]. Swim bladder: Present [3]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas. Arctic coasts of Siberia west to the Laptev Sea and east across Alaskan and Canadian 

Arctic to Kugluktuk, Nunavut; southward through Bering Sea to Korea and southern Japan, and to Del Mar, southern California 

near the United States–Mexico border [4, 5].

Relative Abundance: The most common salmon species in the U.S. Chukchi and Beaufort Seas [5]. Spawning runs occur in 

a number of Chukchi Sea drainages and at least in a few Beaufort Sea drainages, such as the Mackenzie River, perhaps the 

Colville River, and others [8, 9].

Chum Salmon (Oncorhynchus keta), 209 mm, northeastern 

Chukchi Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Chum Salmon (Oncorhynchus keta), at sea within Arctic Outer Continental 

Shelf Planning Areas [6] based on review of published literature and specimens from historical and recent 

collections [2, 7]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Chum Salmon

Oncorhynchus keta

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Chum Salmon 143

Depth Range: Epipelagic, coast and offshore, from surface to at least 750 m [10], primarily less than 40 m [10–12].

Habitats and Life History

Anadromous. 

Eggs—Size: At least 6.0–10.0 mm [13, 14]. Time to hatching: Highly variable, 52–182 days [14]. Size at 

hatching: 20–25.0 mm [13, 15]. Habitat: Benthic, buried under gravel of streams and rivers [13, 14]. 

Alevins (larvae)—Size range: 20–35.0 mm [13] Habitat: Under gravel of streams and rivers until yolk sac 

absorbed (30–50 days) [14, 16]. 

Fry—Size at emergence: About 30–39 mm [15]. Length of time as fry: 1 month or more. May remain in fresh 

water for several weeks or may immediately migrate downstream and enter marine or estuarine waters [14, 17]. 

Habitat: Demersal, in streams and rivers [13, 14]. 

Smolt (juveniles) and ocean phase—Size when entering ocean: Between 30–70.0 mm, depending on distance 

from spawning grounds [13]. Habitat: Juveniles spend their first few months to a year in nearshore, marine 

waters, often inhabiting eelgrass or algal beds [18], and then two to five winters offshore [19]. 

Adults: Age and size at first maturity: Highly variable, 2–6 years, primarily 3–5 years, and 60–80 cm [14]. 

Maximum age: 7 years [16]. Maximum size: 109 cm [14]. Habitat: Adults return to freshwater streams and rivers 

to spawn [13, 14]. 

Substrate—Gravel for spawning [14]. 

Physical/chemical—Temperature: At sea, -0.8–22.5 °C [20], most common between 2 and 11 °C [21]. Salinity: 

Fresh water to marine, depending on life stage [14]. Fry are euryhaline and can withstand daily fluctuations of 

0–27 parts per thousand [14].

Behavior

Diel—Fry typically emerge from gravel and migrate downstream at night [14]. At sea, many individuals may 

ascend into surface waters at night and return to depth during the day [20]. 

Seasonal—Fry emerge from gravel beds in spring [17]. At-sea behavior in U.S. Chukchi and Beaufort Seas is 

unknown. It is speculated that fish entering the Beaufort Sea from various drainages may overwinter either in 

deeper waters of the Beaufort Sea or in the North Pacific [9]. 

Reproductive—Spawning occurs in freshwater in streams and rivers. Females prepare a gravel nest and lay their 

eggs, which are fertilized by a male [14]. All adults die after spawning [13]. 

Schooling—Fry do not strongly school [14]. Juveniles in estuaries and nearshore waters may form loose 

aggregations [14].

Feeding—Juveniles in estuaries feed most intensely at high tides when marsh is submerged [14].

Benthic and reproductive distribution of Chum Salmon (Oncorhynchus keta).

3-3-5g_ChumSalmon

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

1,000

800

600

400

200

0

Juveniles and pre-spawning adults

Some fish to 750 m

Onchorynchus keta

Chum Salmon

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Data from outside U.S. Beaufort-Chukchi Seas

Anadromous.

Spawning, eggs, alevins

and fry all occur in freshwater

No reproductive distribution in seawater 

Smolts in nearshore

estuaries
144 Alaska Arctic Marine Fish Ecology Catalog

Populations or Stocks

The State of Alaska monitors catch and escapement of Chum Salmon in the major river drainages in Kotzebue 

Sound. Genetic analysis of samples of summer run Chum Salmon from six watersheds indicates a Kotzebue 

Sound complex that is related to but distinct from Norton Sound, Yukon, and Kuskokwim regions [22].

Reproduction

Mode—Gonochoristic, oviparous, and semelparous with external fertilization [23].

Spawning season—Unknown in U.S. Chukchi and Beaufort Seas watersheds except for stocks from Kotzebue 

Sound. In the Mackenzie River, Canada, spawning migrations are during July and August with spawning in 

autumn [24]. In other regions, spawning is in June–January [17]. 

Fecundity—At least 900–8,000 eggs [17].

Food and Feeding

Food items—Zooplankton (for example, copepods, euphausiids, mysids, pteropods, and fish larvae) as well as 

insects and small amounts of fishes [25–27]. 

Trophic level—3.47 (standard error 0.48) [28].

Biological Interactions

Predators—A wide variety of fishes, sea birds, and mammals. Examples include Pacific Halibut, Pacific Sleeper 

Shark, Walleye Pollock, rhinoceros auklet, pigeon guillemot, marbled murrelet, pelagic cormorant, northern fur 

seal, beluga whale, and orca [16, 29–31] [32–35]. 

Competitors—Likely other planktivores (for example, salmonids, Pacific Herring, and Polar and Saffron cods). 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (K=0.27–0.45; t

m=2–5; tmax=6) [28].

Traditional and Cultural Importance 

Taken with some frequency in subsistence fisheries along the Chukchi Sea (that is, Kotzebue Sound) and near 

Barrow [36–40]. Chum Salmon also are taken in the Mackenzie River and its tributaries and in the Colville River 

[37, 41].

Commercial Fisheries

The Alaska Department of Fish and Game manages small Chum Salmon fisheries in Kotzebue Sound. Market 

conditions and availability of fish buyers have affected harvest levels in recent years. Estimated low salmon 

abundance in 2014 resulted in the termination of the fishery.

Potential Effects of Climate Change

An increase in the Arctic population with climate change has been hypothesized [42]. Availability of freshwater 

habitats and sufficient thermal conditions for incubation are potential limiting factors.
Chum Salmon 145

Areas for Future Research [A]

The potential colonization rivers and streams of Arctic Alaska is of research interest. Like other salmonids, 

limitations associated with potential source populations, incubation in freshwater habitats (temperature and 

oxygen), timing of seaward migration and related quality of nearshore marine habitats for juveniles, dispersal 

routes and migration corridors, and distance from natal habitats to oceanic rearing grounds will affect 

colonization success. Potential interactions and outcomes of Chum Salmon and other nearshore fishes is an 

expressed concern to residents. Regular surveys of prospective habitats for colonizing fish and evidence of 

successful reproduction should be established.

References Cited

Bakkala, R.G., 1970, Synopsis of biological data on the chum salmon, Oncorhynchus keta (Walbaum, 1792): U.S. Fish and 

Wildlife Sevice, FAO Fisheries Synopsis no. 41, 89 p. [17]

Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates in 

west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments Division, 

ELMR Report no. 8, 327 p. [13]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Pauley, G.B., Bowers, K.L., and Thomas, G.L., 1988, Species profiles—Life histories and environmental requirements of 

coast fishes and invertebrates (Pacific Northwest)—Chum salmon: U.S. Fish and Wildlife Service and U.S. Army Corps of 

Engineers, Biological Report no. 82 (11.81), TR EL-82-4, 17 p. [16]

Salo, E.O., 1991, Life history of chum salmon (Oncorhynchus keta), in Groot, C., and Margolis, L., eds., Pacific salmon life 

histories: Vancouver, British Columbia, UBC Press, p. 233–309. [14]

Bibliography

1. Carothers, C., 2010, Subsistence use and knowledge of Beaufort salmon populations: 2010 Annual Report, Cooperative 

Agreement M09AC15378.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

4. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

5. Stephenson, S.A., 2006, A review of the occurrence of Pacific Salmon (Oncorhynchus spp.) in the Canadian Western Arctic: 

Arctic, v. 59, no. 1, p. 37–46.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

8. Craig, P.C., and Haldorson, L.J., 1986, Pacific salmon in the North American Arctic: Arctic, v. 39, no. 1, p. 2–7. 
146 Alaska Arctic Marine Fish Ecology Catalog

9. Irvine, J.R., Macdonald, R.W., Brown, R.J., Godbout, L., Reist, J.D., and Carmack, E.C., 2009, Salmon in the Arctic and 

how they avoid lethal low temperatures: North Pacific Anadromous Fish Commission, Bulletin no. 5, p. 39–50.

10. Walker, R.V., Sviridov, V.V., Urawa, S., and Azumaya, T., 2007, Spatio-temporal variation in vertical distributions of Pacific 

salmon in the ocean: North Pacific Anadromous Fish Commission, Bulletin no. 4, p. 193–201.

11. Ogura, M., and Ishida, Y., 1995, Homing behavior and vertical movements of four species of Pacific salmon (Oncorhynchus

spp.) in the central Bering Sea: Canadian Journal of Fisheries and Aquatic Sciences, v. 52, no. 3, p. 532–540.

12. Ishida, Y., Yano, A., Ban, M., and Ogura, M., 2001, Vertical movement of a chum salmon Oncorhynchus keta in the western 

North Pacific Ocean as determined by a depth-recording archival tag: Fisheries Science, v. 67, no. 6, p. 1,030–1,035.

13. Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates 

in west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments 

Division, ELMR Report no. 8, 327 p.

14. Salo, E.O., 1991, Life history of chum salmon (Oncorhynchus keta), in Groot, C., and Margolis, L., eds., Pacific salmon life 

histories: Vancouver, British Columbia, UBC Press, p. 233–309.

15. Beacham, T.D., and Murray, C.B., 1990, Temperature, egg size, and development of embryos and alevins of five species of 

Pacific salmon—A comparative analysis: Transactions of the American Fisheries Society, v. 119, no. 6, p. 927–945.

16. Pauley, G.B., Bowers, K.L., and Thomas, G.L., 1988, Species profiles—Life histories and environmental requirements of 

coast fishes and invertebrates (Pacific Northwest)—Chum salmon: U.S. Fish and Wildlife Service and U.S. Army Corps of 

Engineers, Biological Report no. 82 (11.81), TR EL-82-4, 17 p.

17. Bakkala, R.G., 1970, Synopsis of biological data on the chum salmon, Oncorhynchus keta (Walbaum, 1792): U.S. Fish and 

Wildlife Sevice, FAO Fisheries Synopsis no. 41, 89 p.

18. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

19. Beamish, R.J., McFarlane, G.A., and King, J.R., 2005, Migratory patterns of pelagic fishes and possible linkages between 

open ocean and coastal ecosystems off the Pacific coast of North America: Deep-Sea Research II, v. 52, p. 739–755.

20. Walker, R.V., Myers, K.W., Davis, N.D., Aydin, K.Y., Friedland, K.D., Carlson, H.R., and others, 2000, Diurnal variation 

in thermal environment experienced by salmonids in the North Pacific as indicated by data storage tags: Fisheries 

Oceanography, v. 9, no. 2, p. 171–186.

21. Brodeur, R.D., 1988, Zoogeography and trophic ecology of the dominant epipelagic fishes in the northern North Pacific: 

University of Tokyo, Bulletin of the Ocean Research Institute, v. 26, no. 2, p. 1–27.

22. Beacham, T.D., Le, K.D., Wetklo, M., McIntosh, B., Ming, T., and Miller, K.D., 2009, Population structure and stock 

identification of Chum Salmon from Western Alaska determined with microsatellite DNA and major histocompatibility 

complex variation: American Fisheries Society Symposium 70, p. 1–20.

23. Moyle, P.B., 1976, Inland fishes of California (1st ed.): Berkeley, University of California Press, 517 p.

24. Hatfield, C.T., Stein, J.N., Falk, M.R., and Jessop, C.S., 1972, Fish resources of the Mackenzie River Valley: Winnepeg, 

Environment Canada Fisheries Service, Interim report 1, v. 1.

25. Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, Fisheries Research Institute, FRI-UW-7719.

26. Pearcy, W.G., Brodeur, R.D., Shenker, J.M., Smoker, W.W., and Endo, Y., 1988, Food habits of Pacific salmon and steelhead 

trout, midwater trawl catches and oceanographic conditions in the Gulf of Alaska, 1980–1985: University of Tokyo, Bulletin 

of the Ocean Research Institute, v. 26, no. 2, p. 29–78.

27. Brodeur, R.D., 1990, A synthesis of the food habits and feeding ecology of salmonids in marine waters of the North Pacific: 

Seattle, University of Washington, Fisheries Research Institute, FRI-UW-9016, 38 p.
Chum Salmon 147

28. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

29. Armstrong, R.H., and Winslow, P.C., 1968, An incidence of Walleye Pollock feeding on salmon young: Transactions of the 

American Fisheries Society, v. 97, no. 2, p. 202–203.

30. Harry, G.Y., and Hartley, J.R., 1981, Northern fur seals in the Bering Sea, in Hood, D.W., and Calder, J.A., eds., The Eastern 

Bering Sea shelf—Oceanography and resources: U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, p. 847–867.

31. Vermeer, K., and Westrheim, S.J., 1984, Fish changes in diets of nestling rhinoceros auklets and their implications, in

Nettleship, D.N., Sanger, G.A., and Springer, P.F., eds., Marine birds—Their feeding ecology and commercial fisheries 

relationships—Proceedings of the Pacific Seabird Group Symposium: Ottawa, Ontario, Canadian Wildlife Service Special 

Publication, p. 96–105.

32. Ford, J.K.B., Ellis, G.M., Barrett-Lennard, L.G., Morton, A.B., Palm, R.S., and Balcomb, K.C., III, 1998, Dietary 

specialization in two sympatric populations of killer whales (Orcinus orca) in coastal British Columbia and adjacent waters: 

Canadian Journal of Zoology, v. 76, p. 1,456–1,471.

33. Huntington, H.P., 2000, Traditional knowledge of the ecology of belugas, Delphinapterus leucas, in Cook Inlet, Alaska: 

Marine Fisheries Review, v. 62, p. 134–140.

34. Sigler, M.F., Hulbert, L.B., Lunsford, C.R., Thompson, N.H., Burek, K., O’Corry-Crowe, G., and others, 2006, Diet of 

Pacific sleeper shark, a potential Steller sea lion predator, in the north-east Pacific Ocean: Journal of Fish Biology, v. 69, 

no. 2, p. 392–405.

35. Yang, M.-S., Dodd, K., Hibpshman, R., and Whitehouse, A., 2006, Food habits of groundfishes in the Gulf of Alaska 

in 1999 and 2001: U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical 

Memorandum, NMFS-AFSC-164, 189 p.

36. Craig, P.C., 1989b, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers University 

of Alaska, no. 24, p. 131–152.

37. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

38. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

39. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

40. Menard, J., 2008, 2008 Kotzebue Sound salmon fisheries management plan: Alaska Department of Fish and Game, 

Regional Information Report No. 3A08-02.

41. George, J.C., and Nageak, B.P., 1986, Observations on the Colville River subsistence fishery at Nuiqsut, Alaska for the 

period 4 July–1 November 1984: Alaska Department of Wildlife Manangement, 70 p.

42. Kaeriyama, M., 2008, Ecosystem-based sustainable conservation and management of Pacific salmon, in Tsukamoto, K., and 

others, eds., Fisheries for global welfare and environment: 5th World Fisheries Congress 2008, Terrapub, p. 371–380.
148 Alaska Arctic Marine Fish Ecology Catalog

Coho Salmon (Oncorhynchus kisutch) 

(Walbaum, 1792)

Family Salmonidae 

Note: Exception for geographic range data, all information is from 

areas outside of the Chukchi and Beaufort Seas. 

Colloquial Name: Iñupiat—Iqalugruaq [1].

Ecological Role: Likely of minor subsistence importance in 

southeastern U.S. Chukchi Sea and occasional occurrence in 

coastal captures farther north.

Physical Description/Attributes: At sea, metallic-blue on back, silvery flanks, and white belly, small black spots on back and 

sides and on the upper lobe of the caudal fin, and white gums at the base of the teeth on the lower jaw. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 206) [2]. Swim bladder: Present [3]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [2, 4, 5]. Worldwide, North Korea and Japan to the Mackenzie River, Canada, and to 

Bahia Camalu, northern Baja California, Mexico [2, 6].

Relative Abundance: Uncommon in southeastern Chukchi Sea (where it returns to rivers to spawn) and rare in northeastern 

Chukchi and Beaufort Seas east to Mackenzie River, Canada [4, 5, 10].

Coho salmon (Oncorhynchus kisutch), freshwater 

spawning stage. Photograph by Thomas Kline, http://www.

salmonography.com.

Coho Salmon

Oncorhynchus kisutch

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Coho Salmon (Oncorhynchus kisutch), at sea within Arctic Outer Continental 

Shelf Planning Areas [7] based on review of published literature and specimens from historical and recent 

collections [2, 8, 9]. 
Coho Salmon 149

Depth Range: Epipelagic, coastal and offshore, from surface to 247 m [11], primarily 40 m or less [12].

Habitats and Life History

Anadromous. 

Eggs—Size: 4.5–7.9 mm [13]. Time to hatching: 5–12 weeks. Length at hatching: About 19–24 mm [14]. 

Habitat: Benthic, buried under gravel of streams and rivers [15]. 

Alevins (larvae)—Size range: 19–30 mm [14, 15]. Habitat: Benthic, remain under gravel for 3–26 weeks until 

yolk sac is absorbed [11, 15]. 

Fry—Size at emergence: About 27–30 mm [14, 15]. Length of time as fry: 1–4 years, typically 1 year [15]. 

Habitat: Benthic, in streams and rivers [15]. 

Smolt and ocean phase—Age and size at smoltification: 1–4 years and 8–17 cm [15]. Habitat: Epipelagic [13, 

15]. Young fish newly entered into ocean waters often reside near shore, frequently in eelgrass beds, but also in 

kelp or over bare sea floor [16]. Generally, do not migrate far offshore. Some spend their entire marine lives in 

inshore waters [17]. 

Adults—Age and size at first maturity: A few males mature at 1 year, but most fish at 3–4 years and at about 

60–70 cm [15]. Maximum age: About 4 years [15]. Maximum size: 108 cm [2]. Habitat: Mature adults return to 

rivers and streams to spawn [13, 15]. 

Substrate—Gravel for spawning [15]. 

Physical/chemical—Temperature: At sea, 5–16.2 °C, primarily 7 °C and greater [15, 18, 19]. Salinity: Fresh to 

marine waters depending on life stage [15].

Behavior

Diel—Fry and smolts migrate primarily at night [15]. At sea, many or most individuals may ascend into surface 

waters at night and return to depth during day [19]. 

Seasonal—Fry emerge from under gravel as much as 6 months after eggs were laid [15]. Young fish may spend 

as much as 4 years in fresh water before migrating to the sea, although a 1-year residency (including a winter) is 

more typical [15]. Most fish enter seawater in spring and many remain nearshore for at least a few months before 

moving somewhat offshore. At sea, fish entering from California to British Columbia tend to move northwards. 

Generally, Coho Salmon do not migrate far offshore and some spend their entire marine lives in inshore waters 

[17]. Utilization of offshore waters as well as migrations in U.S. Chukchi and Beaufort Seas is unknown. Coho 

Salmon spend 18 months or more at sea before entering freshwater to spawn [15]. 

Reproductive—Spawning occurs in freshwater streams and rivers. Females prepare a gravel nest and lay their 

eggs, which are fertilized by a male [15]. All adults die after spawning [13]. 

Schooling—Smolts and marine individuals form schools [15]. 

Feeding—Juveniles and adults are carnivorous, opportunistic feeders [13].

Benthic and reproductive distribution of Coho Salmon (Oncorhynchus kisutch). 

3-3-6g_CohoSalmon

Some fish to 247 m

Open ocean Shore

Depth, in meters

250

200

150

100

50

0

250

200

150

100

50

0

Oncorhynchus kisutch

Coho Salmon

Benthic distribution

Open ocean No reproductive distribution in seawater Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning, eggs, alevins

and fry all occur in freshwater

 Data from outside 

U.S. Beaufort-Chukchi Seas

Juveniles and pre-spawning adults

Smolts in nearshore

estuaries
150 Alaska Arctic Marine Fish Ecology Catalog

Populations or Stocks

There have been no studies in the U.S. Chukchi and Beaufort Seas.

Reproduction

Mode—Gonochoristic, oviparous, and semelparous with external fertilization [20].

Spawning season—October–March, primarily November–January [15]. 

Fecundity—At least 1,724–7,600 eggs [15], varies between years, increasing to the north [21, 22].

Food and Feeding

Food items—Various fishes and squids, and such small water column zooplankters as euphausiids, hyperiid 

amphipods, and crustacean larvae. Diets change seasonally and between years [23]. 

Trophic level—4.2 [24].

Biological Interactions

Predators—Fishes, sea birds, and marine mammals [11].

Competitors—Possibly gadids such as Arctic Cod and other salmonids.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (K=0.98; t

m=2–4; Fecundity=1,400) [24].

Traditional and Cultural Importance

Only rarely, but apparently increasing numbers harvested in subsistence fisheries in U.S. Chukchi and Beaufort 

Seas [10].

Commercial Fisheries

Currently, Coho Salmon are not commercially harvested.

Potential Effects of Climate Change

Warming temperatures likely will allow this species to colonize Arctic marine and freshwater ecosystems. It can 

be expected that Coho Salmon will eventually become more abundant, especially in the southeastern Chukchi 

Sea. Because of thermal requirements for incubation, it is unclear whether the viable populations will be able to 

establish in watersheds of the Chukchi and Beaufort Seas.
Coho Salmon 151

Areas for Future Research [B]

Coho Salmon are well studied in the State of Alaska. Specific information from the region is lacking with respect 

to (1) preferred habitat, including depth ranges for juveniles and adults; (2) spawning season; (3) seasonal and 

ontogenetic movements; (4) population studies; (5) prey; and (6) predators. Surveys of Chukchi and Beaufort Sea 

watersheds for suitable spawning and rearing habitats, evidence of spawning, or successful reproduction, should 

be considered.

References Cited

Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates in 

west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments Division, 

ELMR Report no. 8, 327 p. [13]

Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p. [11]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Sandercock, F.K., 1991, Life history of Coho salmon (Oncorhynchus kisutch), in Groot, C., and Margolis, L., eds., Pacific 

salmon life histories: Vancouver, British Columbia, UBC Press, p. 397–445. [15]

Bibliography

1. Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum 

Reserve, Alaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife 

Management, 94 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

4. Craig, P.C., and Haldorson, L.J., 1986, Pacific salmon in the North American Arctic: Arctic, v. 39, no. 1, p. 2–7. 

5. Stephenson, S.A., 2006, A review of the occurrence of Pacific Salmon (Oncorhynchus spp.) in the Canadian Western Arctic: 

Arctic, v. 59, no. 1, p. 37–46.

6. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

7. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

8. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

9. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

10. Menard, J., 2008, 2008 Kotzebue Sound salmon fisheries management plan: Alaska Department of Fish and Game, 

Regional Information Report No. 3A08-02.
152 Alaska Arctic Marine Fish Ecology Catalog

11. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

12. Walker, R.V., Sviridov, V.V., Urawa, S., and Azumaya, T., 2007, Spatio-temporal variation in vertical distributions of Pacific 

salmon in the ocean: North Pacific Anadromous Fish Commission, Bulletin no. 4, p. 193–201.

13. Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates 

in west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments 

Division, ELMR Report no. 8, 327 p.

14. Beacham, T.D., and Murray, C.B., 1990, Temperature, egg size, and development of embryos and alevins of five species of 

Pacific salmon—A comparative analysis: Transactions of the American Fisheries Society, v. 119, no. 6, p. 927–945.

15. Sandercock, F.K., 1991, Life history of Coho salmon (Oncorhynchus kisutch), in Groot, C., and Margolis, L., eds., Pacific 

salmon life histories: Vancouver, British Columbia, UBC Press, p. 397–445.

16. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

17. Beamish, R.J., McFarlane, G.A., and King, J.R., 2005, Migratory patterns of pelagic fishes and possible linkages between 

open ocean and coastal ecosystems off the Pacific coast of North America: Deep-Sea Research II, v. 52, p. 739–755.

18. Brodeur, R.D., 1988, Zoogeography and trophic ecology of the dominant epipelagic fishes in the northern North Pacific: 

University of Tokyo, Bulletin of the Ocean Research Institute, v. 26, no. 2, p. 1–27.

19. Walker, R.V., Myers, K.W., Davis, N.D., Aydin, K.Y., Friedland, K.D., Carlson, H.R., and others, 2000, Diurnal variation 

in thermal environment experienced by salmonids in the North Pacific as indicated by data storage tags: Fisheries 

Oceanography, v. 9, no. 2, p. 171–186.

20. Moyle, P.B., 2002, Inland fishes of California (2nd ed.): Berkeley, University of California Press, 517 p.

21. Beacham, T.D., 1982, Fecundity of Coho salmon (Oncorhynchus kisutch) and chum salmon (O. keta) in the northeast 

Pacific Ocean: Canadian Journal of Zoology, v. 60, no. 6, p. 1,463–1,469.

22. Fleming, I.A., and Gross, M.R., 1990, Latitudinal clines—A trade-off between egg number and size in Pacific salmon: 

Ecology, v. 71, no. 1, p. 1–11.

23. Brodeur, R.D., 1990, A synthesis of the food habits and feeding ecology of salmonids in marine waters of the North Pacific: 

Seattle, University of Washington, Fisheries Research Institute, FRI-UW-9016, 38 p.

24. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Sockeye Salmon 153

Sockeye Salmon (Oncorhynchus nerka) 

(Walbaum, 1792)

Family Salmonidae 

Note: Except for geographic range data, all information is from 

areas outside the Chukchi and Beaufort Seas. 

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Of limited ecological importance in the U.S. 

Chukchi and Beaufort Seas. Harvested in small numbers in 

Kotzebue Sound; therefore, this fish may play a seasonal role in 

pelagic ecosystems in the southeastern Chukchi Sea.

Physical Description/Attributes: At sea, head blue to greenblue, rest of body silvery. Usually no black spots on back and fins, occasionally small spots on caudal or dorsal fin. For specific 

diagnostic characteristics and spawning coloration, see Fishes of Alaska (Mecklenburg and others, 2002, p. 209) [1]. Swim 

bladder: Present [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [3]. Worldwide, northern Japan and the Sea of Okhotsk to the Klamath River, northern 

California, and Arctic Canada coasts to Bathurst Inlet, Nunavut and northward to the Sachs River estuary, Banks Island [3].

Relative Abundance: Common in U.S. Chukchi Sea northward to about Kotzebue Sound area (where it spawns), occasionally 

farther north in U.S. Chukchi and Beaufort Seas and to Sachs River, Canada [3, 7]. Generally, Sockeye Salmon are uncommon 

in the U.S. Chukchi and Beaufort Seas north of about Kotzebue Sound.

Sockeye Salmon (Oncorhynchus nerka), 237 mm, southwestern 

Gulf of Alaska, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Sockeye Salmon

Oncorhynchus nerka

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Sockeye Salmon (Oncorhynchus nerka), at sea within Arctic Outer Continental 

Shelf Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [1, 5, 6].
154 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Epipelagic, coastal and offshore, from surface to 90 m, but mostly 40 m and less [8, 9].

Habitats and Life History

Anadromous. 

Eggs—Size: 5.0–6.6 mm [10]. Size at hatching: Unknown, probably 20–25 mm TL [11]. Time to hatching: 

57–171 days [10]. Habitat: Benthic, buried in gravel in waterways entering lakes or in shallow waters of 

lakes [10]. 

Alevins (larvae)—Size range: Become fry at about 25–30 mm [10, 11]. Habitat: Benthic, buried in the substrate 

of lakes and rivers [10]. 

Fry—Size at emergence: 25–31 mm [10]. Length of time as fry: Several weeks to 3 years (although streamreared juveniles and juveniles that migrate to sea within a few weeks are known) [10]. Habitat: Lakes and 

streams [10]. 

Smolt and ocean phase—Age and size at smoltification: Less than 1 year to 3 years and at a minimum of 

60 mm [10]. Habitat: Following smoltification, epipelagic in ocean for 1–4 years (mostly 2–3 years), often far 

from shore [10]. In North American waters, may migrate almost as far west as Kamchatka Peninsula, Russia, and 

fish from Asia intermingle with those from North America [10, 12]. 

Adults—Age and size at spawning: On average 3–5 years and 50–60 cm [10]. Maximum age: At least 6 

years [10], a non-native individual lived to 8 years in Connecticut [13]. Maximum size: 84 cm [1]. Habitat: At 

maturity, ascends natal rivers and lake systems to spawn [11]. 

Substrate—Coarse sand to large rubble for spawning [10]. 

Physical/chemical—Temperature: At least 2–8 °C in freshwater during spawning [10]. Less than 1–15 °C, 

mainly 2.5–9 °C at sea [14]. Salinity: Fresh to marine waters, depending on life stage [10].

Behavior

Diel—Fry emerge from substrate and make initial downstream migration at night. Smolt migrations to sea tend 

to be at night [10]. 

Seasonal—Smolts tend to migrate out of lakes when waters warm in spring and summer [11]. Utilization of 

offshore waters as well as migrations in U.S. Chukchi and Beaufort Seas are unknown.

Reproductive—Spawning occurs in freshwater, most often in shallow water of lakes [10]. All adults die after 

spawning [11]. 

Schooling—Fry and freshwater smolts form schools, as do at least younger individuals at sea. The degree of 

schooling at sea, particularly of older individuals, is not well understood [10]. 

Feeding—Juveniles and adults are carnivorous planktivores. Spawning adults cease feeding [11].

Benthic and reproductive distribution of Sockeye Salmon (Oncorhynchus nerka). 

3-3-7g_SockeyeSalmon

Some fish to 90 m

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Oncorhynchus nerka

Sockeye Salmon

Benthic distribution

Open ocean No reproductive distribution in seawater Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning, eggs, alevins and fry 

all occur in freshwater

Data from outside U.S. Beaufort-Chukchi Seas

Juveniles and pre-spawning adults

Smolts in nearshore

estuaries
Sockeye Salmon 155

Populations or Stocks

There have been no studies. 

Reproduction

Mode—Gonochoristic, oviparous, and semelparous with external fertilization [15].

Spawning season—Late July through January, typically from midsummer through late autumn [10]. 

Fecundity—Highly variable, 2,000–5,000 eggs on average [10].

Food and Feeding

Food items—Primarily zooplankton (for example, euphausiids, hyperiid amphipods, copepods, pteropods, and 

crustacean larvae), as well as small fishes and squids. Diets vary with area and year [16, 17]. 

Trophic level—3.4 [18].

Biological Interactions

Predators—Fishes, sea birds, and marine mammals [10]. 

Competitors—Unknown, but likely to include Pacific Herring, Arctic Cod, Saffron Cod, and Dolly Varden.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (K=0.37–0.58; t

m=2–4; tmax=7; Fecundity=300) [18].

Traditional and Cultural Importance 

Only rarely taken in subsistence fisheries in U.S. Chukchi Sea [19].

Commercial Fisheries

Currently, Sockeye Salmon are not commercially harvested in the U.S. Chukchi or Beaufort Seas.

Potential Effects of Climate Change

Warming temperatures will likely allow this species to become more abundant in the U.S. Chukchi and Beaufort 

Seas. Whether this species will be able to reproduce in the waters draining into these seas is unknown.
156 Alaska Arctic Marine Fish Ecology Catalog

Areas for Future Research [B]

Sockeye Salmon are well studied in Alaska, but are not of great abundance in the American high Arctic. Specific 

information needs include: (1) preferred habitat, including depth ranges for juveniles and adults; (2) spawning 

season; (3) seasonal and ontogenetic movements; (4) population studies; (5) prey; and (6) predators. If range 

expansions occur, surveys for suitable spawning and rearing habitats and evidence of successful spawning should 

be undertaken. 

References Cited 

Burgner, R.L., 1991, Life history of sockeye salmon (Oncorhynchus nerka), in Groot, C., and Margolis, L., eds., Pacific salmon 

life histories: University of British Columbia Press, p. 3–117. [10]

Craig, P.C., and Haldorson, L.J., 1986, Pacific salmon in the North American Arctic: Arctic, v. 39, no. 1, p. 2–7. [7]

Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates in 

west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments Division, 

ELMR Report no. 8, 327 p. [11]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

3. Stephenson, S.A., 2006, A review of the occurrence of Pacific Salmon (Oncorhynchus spp.) in the Canadian Western Arctic: 

Arctic, v. 59, no. 1, p. 37–46.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

6. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

7. Craig, P.C., and Haldorson, L.J., 1986, Pacific salmon in the North American Arctic: Arctic, v. 39, no. 1, p. 2–7. 

8. Ogura, M., and Ishida, Y., 1995, Homing behavior and vertical movements of four species of Pacific salmon (Oncorhynchus

spp.) in the central Bering Sea: Canadian Journal of Fisheries and Aquatic Sciences, v. 52, no. 3, p. 532–540.

9. Walker, R.V., Sviridov, V.V., Urawa, S., and Azumaya, T., 2007, Spatio-temporal variation in vertical distributions of Pacific 

salmon in the ocean: North Pacific Anadromous Fish Commission, Bulletin no. 4, p. 193–201.

10. Burgner, R.L., 1991, Life history of sockeye salmon (Oncorhynchus nerka), in Groot, C., and Margolis, L., eds., Pacific 

salmon life histories: University of British Columbia Press, p. 3–117.

11. Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates 

in west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments 

Division, ELMR Report no. 8, 327 p.
Sockeye Salmon 157

12. Beamish, R.J., McFarlane, G.A., and King, J.R., 2005, Migratory patterns of pelagic fishes and possible linkages between 

open ocean and coastal ecosystems off the Pacific coast of North America: Deep-Sea Research II, v. 52, p. 739–755.

13. Galligan, J.P., 1959, An 8-year-old sockeye salmon from Connecticut: Transactions of the American Fisheries Society, v. 88, 

p. 150–151.

14. Brodeur, R.D., and Livingston, P.A., 1988, Food habits and diet overlap of various Eastern Bering Sea fishes: National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS F/NWC-127, 76 p.

15. Moyle, P.B., 2002, Inland fishes of California (2nd ed.): Berkeley, University of California Press, 517 p.

16. Pearcy, W.G., Brodeur, R.D., Shenker, J.M., Smoker, W.W., and Endo, Y., 1988, Food habits of Pacific salmon and steelhead 

trout, midwater trawl catches and oceanographic conditions in the Gulf of Alaska, 1980–1985: University of Tokyo, Bulletin 

of the Ocean Research Institute, v. 26, no. 2, p. 29–78.

17. Brodeur, R.D., 1990, A synthesis of the food habits and feeding ecology of salmonids in marine waters of the North Pacific: 

Seattle, University of Washington, Fisheries Research Institute, FRI-UW-9016, 38 p.

18. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

19. Menard, J., 2008, 2008 Kotzebue Sound salmon fisheries management plan: Alaska Department of Fish and Game, 

Regional Information Report No. 3A08-02.
158 Alaska Arctic Marine Fish Ecology Catalog

Chinook Salmon (Oncorhynchus tshawytscha) 

(Walbaum, 1792)

Family Salmonidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: Iñupiat—Iqalugruaq [1].

Ecological Role: With its apparent increase in abundance in the 

U.S. Chukchi and Beaufort Seas [1], this species is likely to be of 

increasing ecological importance. Chinook Salmon are occasionally 

captured in recreational fisheries at Point Barrow.

Physical Description/Attributes: At sea, metallic green to blueblack back, silver flanks, and white belly, small black spots on 

both lobes of the caudal fin and on back and the gums at the base 

of the teeth of the lower jaw are black. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 207) [2]. Swim bladder: Present [3]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: U.S. Beaufort and Chukchi Seas [4, 5]. Worldwide, from Northern Japan to Canadian Beaufort Sea at Coppermine River 

[2, 4] and to Bahia de Sebastian Vizcaino, central Baja California, Mexico [6].

Relative Abundance: Common to at least the Barrow region [1]. Rare eastward to the Coppermine River, Canada [2, 4]. 

Currently, the northernmost spawning stock known is in Kotzebue Sound, Alaska [10].

Chinook Salmon (Oncorhynchus tshawytscha), southeastern 

Gulf of Alaska, 2003. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

.

Geographic distribution of Chinook Salmon (Oncorhynchus tshawytscha), at sea within Arctic Outer 

Continental Shelf Planning Areas [7] based on review of published literature and specimens from historical 

and recent collections [8, 9].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Chinook Salmon

Oncorhynchus tshawytscha

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Chinook Salmon 159

Depth Range: Epipelagic, coastal and offshore, from surface to 344 m [11]. Bering Sea fish tend to remain 30–50 m below 

the surface, making occasional forays into somewhat shallower waters during the day [12], although those farther south tend to 

remain in deeper waters [13].

Habitats and Life History

Anadromous. 

Eggs—Size: 6.0–8.5 mm [14]. Size at hatching: About 20–24 mm [14, 15]. Time to hatching: 30–160 days, 

warmer temperatures speed hatching time [16]. Habitat: Benthic, buried in gravel in streams and rivers [10]. 

Alevins (larvae)—Size range: 20–35 mm [14]. Habitat: Benthic, buried in gravel in streams and rivers [10]. Stay 

under gravel until yolk sac absorbed (4–6 weeks) [17]. 

Fry—Size at emergence: About 33–37 mm [15]. Length of time as fry: Several weeks to more than 1 year [10]. 

Habitat: Epipelagic in streams, rivers, and estuaries [10]. 

Smolt (juveniles) and ocean phase—Age and size at smoltification: Several weeks to at least 18 months and 

at least 35 mm to perhaps 160 mm [17]. Habitat: Epipelagic [10, 14, 18]. Smolts that quickly leave fresh water 

reside in estuaries for at least a few months, although length of residence varies annually and with location [10]. 

Two life history types: Ocean-type enters the ocean in their first year and migrates less. Stream-type spends first 

year in fresh water, enters the ocean in their second year, and tends to migrate farther [18]. Generally, ocean-type 

does not migrate out of nearshore waters when at sea, whereas stream-type disperses much broader and farther 

from shore [10]. 

Adults—Age at first maturity: Highly variable, depending on stock and ranges from 1 year for males to 7–8 

years for females [10]. Mean ages per stock range from 3–6 years [10]. Maximum age: 8 years [10]. Maximum 

size: 160 cm [2]. Habitat: At maturity, ascends rivers and streams to spawn [10]. 

Substrate—Gravel for spawning [10]. Mud, sand, gravel, and over eelgrass for juveniles in estuaries [14]. 

Physical/chemical—Temperature: At sea, 2–16.9 °C [19, 20], may prefer 7–12 °C [13, 20]. Salinity: Fresh to 

marine waters, depending on life stage [10].

Behavior

Diel—Fry emerge from gravel and travel downstream primarily at night [10]. In at least some areas, fry settle to 

the bottom at dusk and move into shallow waters at night. Juveniles entering estuaries do so at night [17]. At sea, 

appears to move toward surface waters in evening and returns to depth in morning [21]. 

Seasonal—Depending on geographical location, Chinook Salmon may return to natal rivers during almost any 

month of the year [10]. In the U.S. Chukchi and Beaufort Seas, utilization of ocean waters as well as migrations 

are unknown. 

Reproductive—Spawning occurs in freshwater streams and rivers. Females prepare a gravel nest and lay their 

eggs, which are fertilized by a male [10]. All adults die after spawning except for a few “jack” males [14]. 

Schooling—Fry form schools [22]. 

Feeding—Juveniles and adults are carnivorous, opportunistic feeders [10, 14].

Benthic and reproductive distribution of Chinook Salmon (Oncorhynchus tshawytscha). 

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Some fish to 344 m

Oncorhynchus tshawytscha

Chinook Salmon

Benthic distribution

Data from outside U.S. Beaufort-Chukchi Seas 500

400

300

200

100

0

Open ocean No reproductive distribution in seawater Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning, eggs, alevins

and fry all occur in

freshwater

Juveniles and pre-spawning adults

Smolts in nearshore

estuaries
160 Alaska Arctic Marine Fish Ecology Catalog

Populations or Stocks

There have been no studies in the U.S. Chukchi and Beaufort Seas.

Reproduction

Mode—Gonochoristic, oviparous, and semelparous with external fertilization [23].

Spawning season—May–January, depending on run, fish in the more northerly parts of the range tend to spawn 

earlier in the season [10]. 

Fecundity—Highly variable, less than 2,000 to more than 17,000 eggs [10].

Food and Feeding

Food items—At sea, primarily fishes, although zooplankton (for example, euphausiids, amphipods, copepods, 

and pteropods) also is consumed [24]. 

Trophic level—4.4 (standard error 0.76) [25].

Biological Interactions

Predators—Fishes, sea birds, and marine mammals [26]. 

Competitors—Likely other Pacific Salmon, as well as gadid species.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (t

m=4; tmax=9; Fecundity=4,000) [25].

Traditional and Cultural Importance

Historically, Chinook Salmon were only rarely taken in the subsistence fisheries of the Chukchi and Beaufort 

Seas [27, 28]. However, recent catches in the Point Barrow area suggest an increased abundance in recent 

years [1].

Commercial Fisheries

Currently, Chinook Salmon are not commercially harvested in the U.S. Chukchi and Beaufort Seas.

Potential Effects of Climate Change

Warming temperatures likely will allow this species to become more abundant in the U.S. Chukchi and Beaufort 

Seas. Whether this species will be able to successfully reproduce due to thermal requirements for incubation, 

phenology of seaward migration, and quality of nearshore habitats, is unknown.
Chinook Salmon 161

Areas for Future Research [B]

Chinook Salmon are well studied in areas where their populations are abundant in the State of Alaska. Little 

information is available about this species from the U.S. Chukchi and Beaufort Seas. Specific information needs 

include: (1) preferred habitat, including depth ranges for juveniles and adults; (2) spawning season; (3) seasonal 

and ontogenetic movements; (4) population studies; (5) prey; and (6) predators. Surveys to identify suitable 

habitats for spawning and rearing of young are needed. Coastal monitoring of freshwater drainages should 

include searches for evidence of Chinook Salmon spawning and successful reproduction.

References Cited

Beauchamp, D.A., Shepard, M.F., and Pauley, G.B., 1983, Species profiles—Life histories and environmental requirements of 

coastal fishes and invertebrates (Pacific Northwest)—Chinook Salmon: U.S. Fish and Wildlife Service, Biological Services 

Program FWS/OBS 82 (11.6), 15 p. [17]

Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates in 

west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments Division, 

ELMR Report no. 8, 327 p. [14]

Healey, M.C., 1991, Life history of Chinook salmon (Oncorhynchus tshawytscha), in Groot, C., and Margolis, L., eds., Pacific 

salmon life histories: Vancouver, British Columbia, University of British Columbia Press, p. 313–393. [10]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Stephenson, S.A., 2006, A review of the occurrence of Pacific Salmon (Oncorhynchus spp.) in the Canadian Western Arctic: 

Arctic, v. 59, no. 1, p. 37–46. [4]

Bibliography

1. Carothers, C., 2010, Subsistence use and knowledge of Beaufort salmon populations: 2010 Annual Report, Cooperative 

Agreement M09AC15378.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

4. Stephenson, S.A., 2006, A review of the occurrence of Pacific Salmon (Oncorhynchus spp.) in the Canadian Western Arctic: 

Arctic, v. 59, no. 1, p. 37–46.

5. Craig, P.C., and Haldorson, L.J., 1986, Pacific salmon in the North American Arctic: Arctic, v. 39, no. 1, p. 2–7. 

6. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

7. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

8. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

9. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.
162 Alaska Arctic Marine Fish Ecology Catalog

10. Healey, M.C., 1991, Life history of Chinook salmon (Oncorhynchus tshawytscha), in Groot, C., and Margolis, L., eds., 

Pacific salmon life histories: Vancouver, British Columbia, University of British Columbia Press, p. 313–393.

11. Walker, R.V., Sviridov, V.V., Urawa, S., and Azumaya, T., 2007, Spatio-temporal variation in vertical distributions of Pacific 

salmon in the ocean: North Pacific Anadromous Fish Commission, Bulletin no. 4, p. 193–201.

12. Ogura, M., and Ishida, Y., 1995, Homing behavior and vertical movements of four species of Pacific salmon (Oncorhynchus

spp.) in the central Bering Sea: Canadian Journal of Fisheries and Aquatic Sciences, v. 52, no. 3, p. 532–540.

13. Hinke, J.T., Foley, D.G., Wilson, C., and Watters, G.M., 2005, Persistent habitat use of Chinook salmon Oncorhynchus 

tshawytscha in the coastal ocean: Marine Ecology Progress Series, v. 304, p. 207–220.

14. Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates 

in west coast estuaries, Volume II—Species life history summaries: NOAA/NOS Strategic Environmental Assessments 

Division, ELMR Report no. 8, 327 p.

15. Beacham, T.D., and Murray, C.B., 1990, Temperature, egg size, and development of embryos and alevins of five species of 

Pacific salmon—A comparative analysis: Transactions of the American Fisheries Society, v. 119, no. 6, p. 927–945.

16. Alderdice, D.F., and Velsen, F.P.J., 1978, Relation between temperature and incubation time for eggs of Chinook salmon 

(Oncorhynchus tshawytscha): Journal of the Fisheries Research Board of Canada, v. 35, no. 1, p. 69–75.

17. Beauchamp, D.A., Shepard, M.F., and Pauley, G.B., 1983, Species profiles—Life histories and environmental requirements 

of coastal fishes and invertebrates (Pacific Northwest)—Chinook Salmon: U.S. Fish and Wildlife Service, Biological 

Services Program FWS/OBS 82 (11.6), 15 p.

18. Beamish, R.J., McFarlane, G.A., and King, J.R., 2005, Migratory patterns of pelagic fishes and possible linkages between 

open ocean and coastal ecosystems off the Pacific coast of North America: Deep-Sea Research II, v. 52, p. 739–755.

19. Squire, J.L.J., 1987, Relation of sea surface temperature changes during the 1983 El Niño to the geographical distribution 

of some important recreational pelagic species and their catch temperature parameters in the northeast Pacific: Marine 

Fisheries Review, v. 49, no. 2, p. 44–57.

20. Brodeur, R.D., 1988, Zoogeography and trophic ecology of the dominant epipelagic fishes in the northern North Pacific: 

University of Tokyo, Bulletin of the Ocean Research Institute, v. 26, no. 2, p. 1–27.

21. Candy, J.R., and Quinn, T.P., 1999, Behavior of adult Chinook salmon (Oncorhynchus tshawtyscha) in British Columbia 

coastal waters determined from ultrasonic telemetry: Canadian Journal of Zoology, v. 77, no. 7, p. 1,161–1,169.

22. Reimers, P.E., 1968, Social behavior among juvenile fall Chinook salmon: Journal of the Fisheries Research Board of 

Canada, v. 25, no. 9, p. 2,005–2,008.

23. Moyle, P.B., 2002, Inland fishes of California (2nd ed.): Berkeley, University of California Press, 517 p.

24. Brodeur, R.D., 1990, A synthesis of the food habits and feeding ecology of salmonids in marine waters of the North Pacific: 

Seattle, University of Washington, Fisheries Research Institute, FRI-UW-9016, 38 p.

25. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

26. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

27. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

28. Menard, J., 2008, 2008 Kotzebue Sound salmon fisheries management plan: Alaska Department of Fish and Game, 

Regional Information Report No. 3A08-02.
Dolly Varden 163

Dolly Varden (Salvelinus malma)

(Walbaum, 1792)

Family Salmonidae 

Note: Two subspecies are recognized by some taxonomists: 

Salvelinus malma malma, distributed from the Mackenzie River to 

the Alaska Peninsula, and S. malma lordi, living along and 

southward of the Alaska Peninsula [1, 2].

Colloquial Name: Iñupiat—Iqalukpik [3].

Ecological Role: Likely of considerable ecological importance as 

fish predator in nearshore areas of the U.S. Chukchi and Beaufort 

Seas. Dolly Varden are widely distributed and common throughout 

the region.

Physical Description/Attributes: Olive green to dark blue or brown on back and sides, with profuse yellow, orange, or red 

spots on side; largest spots usually smaller than pupil of eye; pectoral, pelvic, and anal fins with white leading edge and black 

or red line behind; spawning males orange to red, ventrally [4]. Coloration is highly variable and depends on race, age, and 

reproductive state. Extensive descriptions are given in Fishes of Alaska (Mecklenburg and others, 2002, p. 200) [4], McCart 

and others (1972) [5], and Armstrong and Morrow (1980) [1]. Swim bladder: Present [6]. Antifreeze glycoproteins in blood 

serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas. Elsewhere in Alaska, found in all coastal waters in Bering Sea, south to Aleutian 

Islands and southern Gulf of Alaska. Worldwide, south to northern Washington and Korean Peninsula and Japan, northward to 

the Chukchi Peninsula, Russia, and east in Arctic to Mackenzie River, Canada, [4, 7].

Dolly Varden (Salvelinus malma). Photograph by Alfred L. 

DeCicco, Alaska Department of Fish and Game. 
164 Alaska Arctic Marine Fish Ecology Catalog

Relative Abundance: Common near shore in U.S. Chukchi and Beaufort Seas during summer, except along northeastern 

Chukchi Sea coast (for example Point Lay) [11, 12]. Elsewhere, common in Alaska throughout its range [13]. 

Geographic distribution of Dolly Varden (Salvelinus malma), at sea within Arctic Outer Continental Shelf 

Planning Areas [8] based on review of published literature and specimens from historical and recent 

collections [9, 10]. 

tac14-5222_fig3-3-9m_Salvelinus_malma

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Dolly Varden

Salvelinus malma

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Dolly Varden 165

Depth Range: Epipelagic, mostly nearshore but occasionally well offshore [1]. Primarily stays in top 3–4 m of water column, 

with occasional excursions down to 8 m [14, 15]. 

Benthic and reproductive distribution of Dolly Varden (Salvelinus malma). 

3-3-9gDollyVarden

Open ocean Shore

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

Adults and juveniles

Some fish to 8 m

Salvelinus malma

Dolly Varden

Benthic distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Data from U.S. Beaufort-Chukchi Seas

Open ocean No reproductive distribution in seawater Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning, eggs and fry

occur in freshwater

Habitats and Life History

Note: The systematics of Dolly Varden are complex and the species exhibits a wide range of behaviors including 

amphidromous stocks, stocks with freshwater resident males, but ocean-going females (termed “facultative 

anadromous” in literature, but referred to here as “amphidromous”) [16], and various stocks where both sexes 

permanently reside in streams, springs, or lakes [1, 17]. Information in this account is on amphidromous S. 

malma malma living in U.S. Chukchi and Beaufort Sea drainages. Biological aspects of landlocked stocks and 

of more southerly subspecies are available in [1, 13, 18]. In U.S. Chukchi and Beaufort Sea drainages, most 

individuals are amphidromous, although some males and dwarf residents never leave freshwater [1]. In Beaufort 

Sea drainages, winter and spring are spent in those few areas of rivers kept ice-free by freshwater springs [1]. 

Eggs—Size: 3.2–6.0 mm [1, 19, 20]. Time to hatching: 7–9 months [1, 5]. Length at hatching: 8–12 mm [5, 19, 

21]. Habitat: Benthic, buried under gravel in freshwater streams and rivers [1]. 

Alevins (larvae)—Size: From 8–27 mm [5, 19]. Time to emergence: 60–70 days post-hatching [22]. Habitat: 

Benthic, in freshwater rivers and streams, under gravel until emergence (yolk-sac absorbed) [22].

Fry and parr—Size at emergence: 20–27 mm [5, 19]. Length of time as fry and parr: 1 to 5 years, mostly at 3–5 

years [1, 19, 20, 23]. Habitat: Freshwater, in streams and rivers. 

Smolt and Adult—Age and size at smoltification: 1 to 5 years, mostly at 3–5 years [1, 19, 20, 23], and 16–39 cm

[1]. Habitat: Nearshore waters during summer and early autumn and freshwater during winter and spring [1]. 

Adults—Age and size at first maturity: 4–10 years, mostly at 6–8 years [5, 19, 23, 24] and 32–47 cm [20, 

24–26]. Nonmigratory or “residual” males mature at 2 years [5]. Maximum age: At least 18 years [16], although 

fish over 10 years are relatively uncommon [1, 20, 27–29]. Anadromous fish may live slightly longer than resident 

individuals and grow larger [5, 30]. Maximum size: 100 cm TL [4]. Habitat: Nearshore, shallow waters in ocean. 

In freshwater on spawning grounds, main channels of rivers, usually in strong currents or in the presence of 

springs [1].

Substrate—In marine conditions, mostly in mid-depths and surface waters. Spawning occurs in freshwater over 

gravel [1]. 

Physical/chemical—Temperature: -1–14 °C [16, 31]. Salinity: Freshwater to 32 parts per thousand [16, 31].
166 Alaska Arctic Marine Fish Ecology Catalog

Behavior

Diel—Unknown. 

Seasonal—On North Slope watersheds, before their first seaward migration, parr often overwinter in areas 

kept free of ice by perennial springs [5]. Some offspring of amphidromous Dolly Varden do not migrate to sea, 

but remain and mature in freshwaters. These diminutive “resident,” “residual,” or “accessory” fish are mostly 

males [5, 16]. Seaward migrations occur with ice break-up between May and July, with larger individuals 

leaving over-wintering grounds first. Fish enter marine waters in June and July [19, 31, 32]. Most juveniles 

have a variable freshwater rearing period of 1–3 years before first migrating to sea. Smaller fish entering marine 

waters form small schools [23, 31] and remain near mouths of natal rivers, whereas larger fish disperse widely 

along the coast, sometimes traveling as much as 300 km (Beaufort Sea) [1, 16, 30, 33, 34]. Some fish from 

Chukchi Sea drainages make even longer marine excursions. At the extreme, two fish tagged on overwintering 

grounds in the Wulik River (southeastern Chukchi Sea) were recovered well upstream in the Anadyr River of 

Siberia with one of the recoveries as much 1,690 km from the tagging site. It is likely that these recoveries were 

of Anadyr stock that had overwintered in the Wulik River [32]. Once in coastal waters many Dolly Varden inhabit 

shallow nearshore depths [14, 31, 33], although some move away from the coast, perhaps to feed under pack ice 

or when brackish waters extend off shore [31, 33, 35]. All Beaufort and Chukchi Sea Dolly Varden overwinter 

in freshwater and must return to rivers and streams during summer or early autumn [23]. In Beaufort Sea 

watersheds, both spawning and overwintering always take place in tributaries where perennial springs prevent 

freezing throughout the water column [21, 36], although in rivers flowing to the Chukchi Sea the lower parts of 

the waterways also may be partially ice free [32]. In Beaufort Sea, most fish overwinter in their natal drainages 

located in the mountain streams and rivers flowing from the Brooks Range [37]. In the southeastern Chukchi Sea, 

fish do not show overwintering fidelity to natal waterways and stocks from various rivers share ice-free zones [1, 

32, 37]. During return migrations into fresh waters, the timing and behavior of Dolly Varden are quite complex 

and vary with location and with the state of maturity of individual fish [32, 37]. 

Reproductive—Spawning occurs in freshwater streams and rivers. Females prepare a gravel nest and lay their 

eggs, which are fertilized by a male [1]. Spawning periodicity is variable. Some individuals spawn annually, 

whereas many spawn every other year, and apparently there are a few that may reproduce only every third 

year [16, 19, 20, 23, 32]. In Beaufort Sea drainages, fish spawn in many of the “mountain” waterways located 

between the Colville and Mackenzie Rivers. In the southeastern Chukchi Sea, fish spawn in a number of rivers 

including the Kivalina, Noatak, and Wulik Rivers [32]. It is likely that spawning does not occur in Beaufort Sea 

coastal plain rivers that lie west of the Colville River because these waterways lack the perennial springs needed 

to keep waters from freezing solid [36, 38]. The vast majority of Dolly Varden seem to return to natal rivers and 

streams to spawn [32, 36]. 

Schooling—Juveniles form small schools upon entering marine waters [23, 31]. 

Feeding—Opportunistic feeders on various epibenthic and water column organisms including fishes [28]. 

Majority of annual growth is obtained while feeding at sea [22]. Fish migrating upstream in rivers during 

spawning runs feed only occasionally [20, 39].

Populations or Stocks

Genetic studies have shown that there are multiple populations in drainages throughout their range, that these 

populations are centered around the various spawning and overwintering areas, and that there may be more 

than one population occupying tributaries of a single river system [2, 32, 40, 41]. Aerial surveys are used to 

monitor abundance in North Slope drainages. A combination of mark-recapture and genetic studies examined the 

feasibility of estimating population size and stock composition at overwintering grounds on the Ivishak River. 

DIDSON sonar has been used to study abundance patterns in the Hulahula River. The largest overwintering 

populations of Dolly Varden occur in Ivishak, Kongakut and Anaktuvuk Rivers. Other large overwintering 

populations occur in the Hulahula, Canning, and Sagavanirktok Rivers.

Reproduction

Mode—Gonochoristic, oviparous, and iteroparous with external fertilization [1]. 

Spawning season—August–December, peaking in September and October [5, 19, 20, 23, 27, 32]. 

Fecundity—1,500–7,000 eggs [1].
Dolly Varden 167

Food and Feeding

Food items—Epibenthic and water column organisms. Crustaceans (for example, amphipods, mysids, and 

isopods), fishes (for example, Dolly Varden, Fourhorn Sculpin, Arctic Cod, Least Cisco, Arctic Cisco, Arctic 

Lamprey, and snailfish), and insects tend to dominate the diet; fish eggs and polychaetes also are consumed. 

Smaller fish target insects and crustaceans, whereas larger individuals are more apt to feed on fishes [19, 24, 

31, 42]. 

Trophic level—4.5 [43].

Biological Interactions

Predators—Beluga whales, ringed seals, polar bears, and other Dolly Varden [39, 44–46]. 

Competitors—Likely other nearshore fishes, including salmonids, whitefishes, sculpins, and Arctic Cod.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (t

m=3–5) [43].

Traditional and Cultural Importance

A very important subsistence species along much of the Alaskan Beaufort and Chukchi Seas, although catches 

in the northeastern Chukchi Sea are small. Depending on location, large numbers are taken during the summer 

in coastal waters and at inland sites in autumn and winter [12, 24, 47–49]. Also an important recreational 

species [50].

Commercial Fisheries

Currently, Dolly Varden are not commercially harvested.

Potential Effects of Climate Change

Unknown. Of particular relevance is the effect of climate change on the freshwater springs of rivers on the North 

Slope. If these springs dry up in the winter, the winter river refuges afforded Dolly Varden will not exist.

Areas for Future Research [A]

Future research should address the need for reliable estimates of population abundance from major overwintering 

habitats and genetic samples collected. Critical spawning and overwintering habitat should be identified and 

mapped. Subsistence harvests should be monitored, and the stock contribution in those harvests determined. 

Population parameters at key freshwater and coastal monitoring sites should be routinely monitored.
168 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Armstrong, R.H., and Morrow, J.E., 1980, The Dolly Varden charr, Salvelinus malma, in Balon, E.K., ed., Charrs, salmonid 

fishes of the genus Salvelinus: The Hague, Netherlands, Dr. W. Junk B.V. Publishers, p. 99–140. [1]

Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, Simpson 

Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649. [31]

DeCicco, A.L., 1997, Movements of postsmolt anadromous Dolly Varden in northwestern Alaska: American Fisheries Society 

Symposium no. 19, p. 175–183. [32]

McCart, P.J., Craig, P.C., and Bain, H., 1972, Report of fisheries investigations in the Sagavanirtok River and neighboring 

drainages: Bellevue, Washington, Aquatic Environments Ltd., Report to Alyeska Pipeline Service Company, 165 p. [5]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [4]

Yoshihara, H.T., 1973, Annual progress report for monitoring and evaluation of Arctic waters with emphasis on the North Slope: 

Alaska Department of Fish Game, Division of Sport Fisheries, 49 p. [19]

Bibliography

1. Armstrong, R.H., and Morrow, J.E., 1980, The Dolly Varden charr, Salvelinus malma, in Balon, E.K., ed., Charrs, salmonid 

fishes of the genus Salvelinus: The Hague, Netherlands, Dr. W. Junk B.V. Publishers, p. 99–140.

2. Crane, P., DeCicco, F., Spearman, B., and Wenburg, J., 2005, Genetic diversity of Dolly Varden populations in Norton and 

Kotzebue Sound: U.S. Fish and Wildlife Service, Alaska Fisheries Technical Report Number 80, 28 p.

3. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

4. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

5. McCart, P.J., Craig, P.C., and Bain, H., 1972, Report of fisheries investigations in the Sagavanirtok River and neighboring 

drainages: Bellevue, Washington, Aquatic Environments Ltd., Report to Alyeska Pipeline Service Company, 165 p.

6. Jordan, D.S., and Evermann, B.W., 1896, The fishes of North and Middle America—A descriptive catalogue of the species 

of fish-like vertebrates found in the waters of North America, North of the Isthmus of Panama, part 1: Bulletin of the United 

States National Museum, no. 47, 1,240 p.

7. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

8. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

9. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

10. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

11. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.
Dolly Varden 169

12. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

13. Blackett, R.F., 1968, Spawning behavior, fecundity, and early life history of anadromous Dolly Varden, Salvelinus malma

(Walbaum) in southeastern Alaska: Alaska Department of Fish and Game, Research Report Number 6, 85 p.

14. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

15. Jarvela, L.E., and Thorsteinson, L.K., 1997, Movements and temperature occupancy of sonically tracked Dolly Varden and 

Arctic ciscoes in Camden Bay, Alaska: American Fisheries Society Symposium, no. 19, p. 165–174.

16. Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.

17. Reist, J.D., Johnson, J.D., and Carmichael, T.J., 1997, Variation and specific identity of char from northwestern Arctic 

Canada and Alaska: American Fisheries Society Symposium no. 19, p. 250–261.

18. Blackburn, J.E., and Jackson, P.B., 1982, Seasonal composition and abundance of juvenile and adult marine finfish and crab 

species in the nearshore zone of Kodiak Island’s eastside during April 1978 through March 1979: Outer Continental Shelf 

Environmental Assessment Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 377–570.

19. Yoshihara, H.T., 1973, Annual progress report for monitoring and evaluation of Arctic waters with emphasis on the North 

Slope: Alaska Department of Fish Game, Division of Sport Fisheries, 49 p.

20. Craig, P.C., 1976, Fisheries investigations in the Shaviovik River drainage, Alaska, with emphasis on Arctic char in the 

Kavik River, Calgary, Alberta: Aquatic Environments Limited, 36 p.

21. Craig, P.C., and McCart, P.J., 1974, Fall spawning and overwintering areas of fish populations along routes of proposed 

pipeline between Prudhoe Bay and the Mackenzie Delta 1972–1973, in McCart, P.J., ed., Life histories of anadromous and 

freshwater fish in the Western Arctic: Arctic Gas Biological Report Series, v. 20, p. 229.

22. Committee on the Status of Endangered Wildlife in Canada, 2010, COSEWIC assessment and status report on the Dolly 

Varden Salvelinus malma malma (western Arctic populations) in Canada: Ottawa, Canada, Committee on the Status of 

Endangered Wildlife in Canada, 65 p.

23. Craig, P.C., and McCart, P.J., 1975, Fish utilization of nearshore coastal waters between the Colville and Mackenzie Rivers 

with an emphasis on anadromous species, in Craig, P.C., ed., Fisheries Investigations in a coastal region of the Beaufort 

Sea: Calgary, Alberta, Canadian Arctic Gas Study Ltd. and Alaskan Arctic Gas Study Co., Biological Report Series 34, 

p. 172–219.

24. Furniss, R.A., 1975, Inventory and cataloging of Arctic area waters: Alaska Department of Fish and Game, Federal Aid in 

Fish Restoration, Annual Report of Performance, Project F-9-7, Study G-1-1, v. 16, 47 p.

25. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

26. Underwood, T.J., Millard, M.J., and Thorpe, L.A., 1996, Relative abundance, length frequency, age, and maturity of Dolly 

Varden in nearshore waters of the Arctic National Wildlife Refuge, Alaska: Transactions of the American Fisheries Society, 

v. 125, no. 5, p. 719–728.

27. Kendel, R.E., Johnston, R.A.C., Kozak, M.D., and Lobsiger, U., 1974, Movements, distribution, populations and food 

habits of fish in the western coastal Beaufort Sea: Department of the Environment, Canada, Interim Report of Beaufort Sea 

Project, Study B1, 64 p.

28. Bendock, T.N., 1977, Beaufort Sea estuarine fishery study: Alaska Department of Fish and Game Annual Report, Contract 

#03-5-022-69, p. 670–729.
170 Alaska Arctic Marine Fish Ecology Catalog

29. Underwood, T.J., Gordon, J.A., Millard, M.J., Thorpe, L.A., and Osborne, B.M., 1995, Characteristics of selected fish 

populations of Arctic National Wildlife Refuge coastal waters, final report, 1988–91: Fairbanks, Alaska, U.S. Fish and 

Wildlife Service, Fairbanks Fishery Resource Office, Alaska Fisheries Technical Report, no. 28, 590 p.

30. Craig, P.C., and McCart, P.J., 1976, Fish use of nearshore coastal waters in the western Arctic—Emphasis on anadromous 

species, in Hood, D.W., and Burrell, D.C., eds., Assessment of the Arctic marine environment: Fairbanks, University of 

Alaska, Institute of Marine Sciences, p. 361–387.

31. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

32. DeCicco, A.L., 1997, Movements of postsmolt anadromous Dolly Varden in northwestern Alaska: American Fisheries 

Society Symposium no. 19, p. 175–183.

33. Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., and Colonell, J.M., 1996, The 1995 fish and oceanography 

study in Mikkelsen Bay, Alaska: Anchorage, Alaska, Prepared by LGL Alaska Research Associates and Woodward-Clyde 

Consultant for BP Exploration (Alaska) Inc., 102 p. plus apps.

34. Wiswar, D.W., and Frugé, D.J., 2006, Fisheries investigations in western Camden Bay, Arctic National Wildlife Refuge, 

Alaska, 1987: Alaska Fisheries Data Series, U.S. Fish and Wildlife Service, 2006-10, 49 p.

35. Thorsteinson, L.K., Jarvela, L.E., and Hale, D.A., 1990, Arctic fish habitat use investigations—Nearshore studies in the 

Alaskan Beaufort Sea, summer 1988: Final Report, Alaska Office, Ocean Assessments Division, National Oceanic and 

Atmospheric Administration, Research Unit 682, 125 p.

36. Burns, J.J., 1990, Proceedings of a technical workshop on fishes utilized in subsistence fisheries in National Petroleum 

Reserve, Alaska—Barrow, Alaska, October 26–28, 1988: Report to North Shore Bureau, Department of Wildlife 

Management, 94 p.

37. Crane, P., Viavant, T., and Wenburg, J., 2005, Overwintering patterns of Dolly Varden Salvelinus malma in the 

Sagavanirktok River in the Alaska North Slope inferred using mixed-stock analysis: U.S. Fish and Wildlife Service, Alaska 

Fisheries Technical Report, no. 84, 25 p.

38. Gallaway, B.J., and Fechhelm, R.G., 1997, Anadromous and amphidromous fishes, in Truett, J.C., and Johnson, S.R., eds., 

The natural history of an Arctic oil field: San Diego, Academic Press, p. 349–369.

39. Griffiths, W.B., Craig, P., Walder, G., and Mann, G., 1975, Fisheries investigations in a coastal region of the Beaufort Sea 

(Nunaluk Lagoon, Yukon), in Craig, P.C., ed., Fisheries investigations in a coastal region of the Beaufort Sea: Arctic Gas, 

Biological Report Series, v. 34, p. 1–129.

40. Everett, R.J., Wilmot, R.L., and Krueger, C.C., 1997, Population genetic structure of Dolly Varden from Beaufort Sea 

drainages of Northern Alaska and Canada: American Fisheries Society Symposium, no. 19, p. 240–249.

41. Everett, R.J., Wilmot, R.L., and Krueger, C.C., 1998, Population genetic structure of Dolly Varden char from Beaufort Sea 

drainages of northern Alaska and Canada: U.S. Fish and Wildlife Service, Alaska Fisheries Technical Report, no. 43, 

p. 1–16.

42. Fechhelm, R.G., Bryan, J.D., Griffiths, W.B., and Martin, L.R., 1997, Summer growth patterns of northern Dolly Varden 

(Salvelinus malma) smolts from the Prudhoe Bay region of Alaska: Canadian Journal of Fisheries and Aquatic Sciences, 

v. 54, no. 5, p. 1,103–1,110.

43. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

44. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.

45. Lowry, L.F., Frost, K.J., and Seaman, G.A., 1986, Investigations of belukha whales in coastal waters of western and 

northern Alaska: Outer Continental Shelf Environmental Program Unit 612, Final Report, p. 359–392.
Dolly Varden 171

46. Dyck, M.G., and Romberg, S., 2007, Observations of a wild polar bear (Urus maritimus) successfully fishing Arctic charr 

(Salvelinus alpinus) and fourhorn sculpin (Myoxocephalus quadricornis): Polar Biology, v. 30, p. 1,625–1,628.

47. Craig, P.C., 1989b, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers University 

of Alaska, no. 24, p. 131–152.

48. Wiswar, D.W., West, R.L., and Winkleman, W.N., 1995, Fisheries investigation in Oruktalik Lagoon, Arctic National 

Wildlife Refuge, Alaska, 1986: U.S. Fish and Wildlife Service, Alaska Fisheries Technical Report, no. 30, 38 p. 

49. Pedersen, S., and Linn, A., Jr., 2005, Kaktovik 2000–2002 subsistence fishery harvest assessment: U.S. Fish and Wildlife 

Service, Fisheries Resource Monitoring Program, Final Report for FIS Study, Study No. 01-101.

50. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.
172 Alaska Arctic Marine Fish Ecology Catalog

Inconnu and Glacier Lanternfish

Inconnu (Stenodus leucichthys) 

(Güldenstadt, 1772)

Family Salmonidae 

Colloquial Name: Iñupiat—Siiġruaq [1]. Most often called 

sheefish in Alaska.

Ecological Role: Rarely enters marine waters and thus is not of 

ecological importance in the U.S. Chukchi and Beaufort Seas. 

Common in coastal lagoons in Kotzebue Sound, this fish is a major 

predator of pelagic species, such as Pacific Herring and possibly 

juvenile salmon. Inconnu is an important subsistence species in western Alaska including the southeastern Chukchi Sea.

Physical Description/Attributes: Body not much compressed; and colored green, blue, or brown dorsally and silvery white 

ventrally. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 181) [2]. Swim bladder: 

Present [3]. Antifreeze glycoproteins in blood serum: Unknown.

Range: Northward to Kobuk River [2] and probably Noatak River drainages of U.S. Chukchi Sea [4]. Elsewhere in Alaska, this 

fish has been found as far south as Kuskokwim River [2]. Worldwide, Inconnu are found from Firth to Anderson Rivers, Canada 

[5], and in Caspian, Siberian, and White Sea drainages, south to Kamchatka, Russia [2]. Reported in nearshore semi-saline 

waters of Canadian Beaufort Sea to at least as far west as Herschel Island and Nunaluk Lagoon, Yukon Territory; most numerous 

just west of Mackenzie River, Northwest Territory [6].

Inconnu (Stenodus leucichthys). Photograph by R.J. Brown, 

U.S. Fish and Wildlife Service.
Inconnu 173

Geographic distribution of Inconnu (Stenodus leucichthys), in brackish nearshore and delta waters within 

Arctic Outer Continental Shelf planning areas [7] based on review of published literature and specimens from 

historical and recent collections [2, 8].

tac14-5222_fig3-4-1m_Stenodus_leucichthys

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Inconnu

Stenodus leucichthys

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Relative Abundance: Common in the Kobuk River. Despite earlier reports of scattered fish in the Meade and Colville Rivers, 

there have been no recent reports of fish from any North Slope drainage [2].
174 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Shallow waters in rivers and brackish, near-shore coastal systems [9].

Habitats and Life History

Amphidromous.

Eggs—Size: 2.5–2.7 mm [5, 10–13]. Time to hatching: 6–9 months. Habitat: Benthic, buried in gravel in 

freshwater rivers [5, 10, 14]. 

Larvae—Size at hatching: 11.0–11.3 mm [15]. Size at juvenile transformation: 7.0 cm [15]. Days to juvenile 

transformation: Unknown. Habitat: Newly hatched larvae are carried down river to nursery and overwintering 

areas in river deltas, estuaries, or lower reaches of watersheds [11]. 

Juveniles—Age and size: 0–7 years and about 70 cm average [5, 10, 14]. Habitat: Fresh and brackish water [5, 

10, 14]. 

Adults—Age and size at first maturity: Males mature at 4–11 years (70–85 cm) [10, 11, 14] and females at 6–14 

years (75–90 cm) [10, 14]; most fish mature at 8–12 years [16]. On average, males mature when younger and 

smaller than females [5]. Maximum age: At least 41 years [17]. Older studies using scales to age fish rather than 

otoliths underestimated fish ages beginning at about 10 years [18]. Females grow larger than males, live longer, 

and tend to reach maturity later [9–11, 14]. Growth rates, life spans, and size and age at maturity vary between 

watersheds and between populations within watersheds [11, 19]. For instance, fish in Kobuk and Selawik Rivers 

live longer and grow larger than those in Yukon and Kuskokwim systems; however, Kobuk and Selawik fish 

grow slower [14]. Maximum size: 140 cm [2]. Habitat: Large and slow moving rivers and estuaries. Entire life is 

spent within or adjacent to their home rivers [5, 11, 16]. 

Substrate—Coarse gravel and cobble mixed with sand for spawning [10, 11, 16]. 

Physical/chemical—Temperature: Spawning occurs at least between 0–7 °C [5, 13]. Salinity: Primarily 

freshwater, but some in brackish waters to about 20 parts per thousand [5, 20].

Benthic and reproductive distribution of Inconnu (Stenodus leucichthys).

3-4-1g_Inconnu

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Stenodus leucichthys

Inconnu

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Anadromous.

Spawning, eggs and larvae 

all occur in freshwater

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Adults and juveniles 

Depths reported as shallow and brackish Depths reported as shallow and brackish
Inconnu 175

Behavior

Diel—Unknown. 

Seasonal—Many watersheds contain a relatively small resident freshwater population that migrate within a river 

system and a larger amphidromous population that may or may not enter estuarine waters in a given year [5, 10]. 

For example, Mackenzie River sea-going inconnu spend their first 1–2 years in freshwater and then tend to move 

annually into brackish and more marine coastal waters. Among these stocks, some regularly migrated to sea 

throughout their lives, whereas others made only occasional estuarine migrations or sea migrations followed by 

extended periods in freshwater or in estuaries [9]. In some rivers, resident and amphidromous forms do not share 

feeding, overwintering, or spawning grounds [20, 21]. Overwintering areas vary with watershed. Fish in Kobuk 

and Selawik Rivers overwinter in brackish waters of Hotham Inlet and Selawik Lake. Fish in the lower Yukon 

and Kuskokwim Rivers spend winter in those rivers’ deltas [11], whereas Mackenzie River fish winter both in the 

delta and nearshore coastal waters of Canadian Beaufort Sea [9]. Slightly before or at about ice break-up time, 

adults and some juveniles begin to leave overwintering grounds [14, 22], although in some areas (for example, 

Selawik Lake) juveniles remain on nursery grounds throughout the year [10]. Individuals that will spawn in 

autumn migrate with other fish to feeding grounds during summer, but do not feed, instead they continue on to 

spawning grounds [5, 14]. Feeding (non-spawning) fish migrate back to overwintering grounds during August 

and September [5, 10, 16]. 

Reproductive—Spawn in their natal rivers. Spawning migrations may be long; as much as 1,800 km on 

the Mackenzie River [23], 1,500 km on the Yukon River [4], and 2,400 km on Siberian rivers [5]. Arrive on 

spawning grounds as early as 1–2 months before spawning in early autumn [5, 14, 24]. In the Selawik and 

Kobuk Rivers, spawning occurs in late afternoon and evening between at least 1500 and 1800 hours (local), 

perhaps peaking between 1545 and 2200 hours (local) [5, 10, 14]. Spawning sites are in relatively small and 

restricted areas, although these may change with time [5]. Inconnu are broadcast spawners in shallow and fast 

moving waters over coarse gravel and cobble mixed with sand [10, 11, 16]. Females release eggs at the surface 

and males release sperm just below the surface and rarely come to the surface [5, 11]. A female emits eggs in a 

series of discrete spawning spurts, returning to the bottom between each episode [5]. Eggs fall to the riverbed and 

are slightly adhesive to gravel and cobble [11]. Eggs are deposited in autumn and reportedly hatch around time 

of ice break-up in early spring [12, 16]. Some populations spawn annually [17]. However, in other populations, 

most individuals do not spawn annually; however, males are more likely to spawn in sequential years [16, 25, 

26]. Although many migrate downstream immediately after spawning (arriving in October), others remain on 

spawning grounds for some length of time (as late as January) [5, 10, 16]. 

Schooling—Forms schools [5, 10, 14]. 

Feeding—Migrating juveniles and non-spawning adults travel to feeding areas. In western Alaska, foraging 

areas tend to be in lower reaches of rivers, upstream of overwintering grounds [22], but also include the brackish 

waters of Kotzebue Sound (for Kobuk and Selawik River fish) [5] and the Beaufort Sea (for Mackenzie River 

fish) [20]. Does not feed during spawning migrations (about 1–4 months) [14].

Populations or Stocks

Two distinct spawning locations have been identified, one in the upper Kobuk River and one on the refuge in 

the upper Selawik River. In cooperation with the Native Village of Kotzebue and Alaska Department of Fish and 

Game, genetic mixed stock analysis is ongoing to understand the proportion of the winter inconnu harvest that 

comes from each of the two spawning populations.

Reproduction

Mode—Gonochoristic, oviparous, iteroparous with external fertilization [5, 10, 11]. 

Spawning season—Autumn, primarily September and October [9, 16, 27]. 

Fecundity—26,000–455,000 eggs [5, 10–12, 17].

Food and Feeding

Food items—Plankton and insects for juveniles. For fish 2 years and older, food is primarily fishes (for example, 

whitefishes, Arctic Lamprey, Pacific Herring, and salmon) and secondarily on small invertebrates such as 

isopods, mysids, and insects [5, 6, 11]. 

Trophic level—4.15 (standard error 0.75) [28].
176 Alaska Arctic Marine Fish Ecology Catalog

Biological Interactions

Predators—Unknown, although grayling, whitefish, and char consume newly spawned eggs [10]. 

Competitors—Likely omnivores such as various whitefish species, char, and grayling.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (K=0.05–0.10; t

m=9–12; tmax=22; Fecundity=80,000) [28].

Traditional and Cultural Importance 

Inconnu are an important subsistence species for humans and dogs. In many watersheds, they are usually taken 

around the time of spawning, but in the Selawik-Kobuk river system a large fishery operates during winter in the 

Hotham Inlet area [4, 9, 10]. Many fish are taken by gill nets; however, seines and hook and line also account for 

large numbers. The fish are eaten fresh, dried, or aged and frozen, and the fat-riddled large intestines are boiled 

for the oil [27]. 

Commercial Fisheries

Currently, inconnu are not commercially harvested. A small barter and trade fishery exists in the Kotzebue area 

and in Great Slave Lake [9, 25] and Inconnu are popular with recreational anglers, particularly on the Kobuk 

River [5].

Potential Effects of Climate Change

Unknown. Generally, Durand and others (2011) [60] predict that, at least for anadromous fishes in subarctic 

rivers, shifts in biology will be effected by spring ice break-up and resultant peak flows and surrounding 

permafrost processes: both of which affect the supply of nutrients and (or) sediment to the watershed of climate 

change on spring break-up intensity. Climate change and its effects on the spawning recruitment of inconnu in 

the Selawik River are being studied in cooperative research between U.S. Fish and Wildlife Service and U.S. 

Geological Survey.

Areas for Future Research [A]

Research needs include: movements and migrations, behavior of larval and juveniles in response to 

environmental variables, and enumeration of predator-prey relationships in coastal waters. Catch and subsistence 

use patterns should continue to be monitored.

References Cited

Alt, K.T., 1969, Taxonomy and ecology of the inconnu, Stenodus leucichthys nelma, in Alaska: Anchorage, University of Alaska, 

Biological Papers of the University of Alaska, no. 12, 63 p. [10]

Alt, K.T., 1987, Review of sheefish (Stenodus leucichthys) studies in Alaska: Juneau, Alaska Department of Fish and Game, 

Division of Sport Fish, Fishery Manuscript no. 3., 69 p. [5]

Alt, K.T., 1988, Biology and management of inconnu (Stenodus leucichthys) in Alaska: Finnish Fisheries Research, v. 9, p. 

127–132. [14]

Alt, K.T., 2008, Alaska wildlife notebook series—Sheefish: Alaska Department of Fish and Game. [11]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]
Inconnu 177

Bibliography

1. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Moyle, P.B., and Cech, J.J., Jr., 1996, Fishes—An introduction to ichthyology: Upper Saddle River, New Jersey, PrenticeHall, 590 p.

4. Savereide, J.W., 2002, Under-ice gillnet harvest of sheefish in Hotham Inlet in 2000–2001—Fisheries data series: U.S. Fish 

and Wildlife Service, Office of Subsistence Management, Fisheries Resource Monitoring Program, Final Report (Study No. 

00-020), and Alaska Department of Fish and Game, Division of Sport Fish, Fairbanks, Alaska, No. 02-04, 22 p.

5. Alt, K.T., 1987, Review of sheefish (Stenodus leucichthys) studies in Alaska: Juneau, Alaska Department of Fish and Game, 

Division of Sport Fish, Fishery Manuscript no. 3., 69 p.

6. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.

7. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

8. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

9. Howland, K.L., Tonn, W.M., Babaluk, J.A., and Tallman, R.F., 2001, Identification of freshwater and anadromous inconnu 

in the Mackenzie River system by analysis of otolith strontium: Transactions of the American Fisheries Society, v. 130, 

no. 5, p. 725–741.

10. Alt, K.T., 1969, Taxonomy and ecology of the inconnu, Stenodus leucichthys nelma, in Alaska: Anchorage, University of 

Alaska, Biological Papers of the University of Alaska, no. 12, 63 p.

11. Alt, K.T., 2008, Alaska wildlife notebook series—Sheefish: Alaska Department of Fish and Game.

12. Chereshnev, I., Shestakov, A.V., Yusupov, R.R., Shtundyuk, Y.V., and Slugin, I.V., 2000, Biology of nelma Stenodus 

leucichthys nelma (Coregonidae) from the Anadyr basin (the northeast of Russia): Journal of Ichthyology, v. 40, no. 6, 

p. 457–470.

13. Howland, K.L., Tonn, W.M., and Tallman, R.F., 2002, The influence of genetic and environmental factors on egg 

development and juvenile growth in two life history forms of inconnu (Stenodus leucichthys), in Todd, T., and Fleischer, 

G., eds., Biology and management of coregonid fishes, 1999—Proceedings of the seventh international symposium on the 

biology and management of coregonid fishes held in Ann Arbor, Michigan, U.S.A.: Schweizerbart, Stuttgart, Germany, 

p. 253–264.

14. Alt, K.T., 1988, Biology and management of inconnu (Stenodus leucichthys) in Alaska: Finnish Fisheries Research, v. 9, 

p. 127–132.

15. Sturm, E.A., 1994, Description and identification of larval and juvenile Stenodus leucichthys nelma (Güldenstadt) from 

central Alaska: Copeia, v. 1994, no. 2, p. 472–484.

16. Hander, R.F., Brown, R.J., and Underwood, T.J., 2008, Comparison of inconnu spawning abundance estimates in the 

Selawik River, 1995, 2004, and 2005, Selawik National Wildlife Refuge: U.S. Fish and Wildlife Service, Alaska Fisheries 

Technical Report No. 99.

17. Brown, R.J., Brown, C., Braem, N.M., Carter, W.K., III, Legere, N., and Slayton, L., 2012, Whitefish biology, distribution, 

and fisheries in the Yukon and Kuskokwim river drainages in Alaska—A synthesis of available information: U.S. Fish and 

Wildlife Service, Alaska Fisheries Data Series Number 2012-4, 316 p.
178 Alaska Arctic Marine Fish Ecology Catalog

18. Howland, K.L., Gendron, M., Tonn, W.M., and Tallman, R.F., 2004, Age determination of a long-lived coregonids from the 

Canadian North—Comparison of otoliths, fin rays and scales in inconnu (Stenodus leucichthys): Annales Zoologici Fennici, 

v. 41, p. 205–214.

19. Alt, K.T., 1973b, Age and growth of the inconnu (Stenodus leucichthys) in Alaska: Journal of the Fisheries Research Board 

of Canada, v. 30, no. 1, p. 457–459.

20. Howland, K.L., Tonn, W.M., and Goss, G., 2001, Contrasts in the hypo-osmoregulatory abilities of a freshwater and an 

anadromous population of inconnu: Journal of Fish Biology, v. 59, no. 4, p. 916–927.

21. Howland, K.L., Tallman, R.F., and Tonn, W.M., 2000, Migration patterns of freshwater and anadromous inconnu in the 

Mackenzie River system: Transactions of the American Fisheries Society, v. 129, no. 1, p. 41–59.

22. Alt, K.T., 1977, Inconnu, Stenodus leucichthys, migration studies in Alaska 1961–74: Journal of the Fisheries Research 

Board of Canada, v. 34, no. 1, p. 129–133.

23. Stephenson, S.A., Burrows, J.A., and Babaluk, J.A., 2005, Long-distance migrations by inconnu (Stenodus leucichthys) in 

the Mackenzie River system: Arctic, v. 58, no. 1, p. 21–25.

24. Underwood, T.J., 2000, Abundance, length composition, and migration of spawning inconnu in the Selawik River, Alaska: 

North American Journal of Fisheries Management, v. 20, no. 2, p. 386–393.

25. Alt, K.T., and Taube, T.T., 2002, Abundance and composition of inconnu spawners from the Kobuk River, Alaska: Archiv 

für Hydrobiologie, Special Issues Advances in Limnology, v. 57, p. 639–645.

26. Taube, T.T., 1997, Abundance and composition of sheefish in the Kobuk River, 1996: Anchorage, Alaska Department of 

Fish and Game, Fishery Manuscript No. 97-1, 38 p.

27. Georgette, S., and Loon, H., 1990, Subsistence and sport fishing of sheefish on the upper Kobuk River, Alaska: Kotzebue, 

Alaska Department of Fish and Game, Division of Subsistence, Technical Paper No. 175.

28. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

29. Durand, J.R., Lusardi, R.A., Nover, D.M., Suddeth, R.J., Carmona-Catot, G., Connell-Buck, C.R., and others, 2011, 

Environmental heterogeneity and community structure of the Kobuk River, Alaska, in response to climate change: 

Ecosphere, v. 2, no. 4.
Glacier Lanternfish 179

Glacier Lanternfish (Benthosema glaciale)

(Reinhardt, 1837)

Family Myctophidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Rarely observed in the U.S. Chukchi Sea. The role 

of the species in regional food webs is minimal.

Physical Description/Attributes: Small, silvery fish with compressed body, blunt head, large eyes, and numerous round 

photophores in a specific pattern. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, 

p. 249) [1]. Swim bladder: Present [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea [1, 3]. Elsewhere, from Baffin Bay and northwest Greenland, east to Svalbard Islands, Norway; 

Barents Sea; and Kara Sea [3]. 

Relative Abundance: Rare, one record from U.S. Chukchi Sea near Point Barrow, Alaska [1, 3]. Elsewhere, common in Barents 

Sea [6].

Glacier Lanternfish (Benthosema glaciale). Photograph by 

Rudolf Svensen, http://www.uwp.no.

Geographic distribution of Glacier Lanternfish (Benthosema glaciale), within Arctic Outer Continental Shelf 

planning areas [4] based on review of published literature and specimens from historical and recent collections 

[1, 3, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Glacier Lanternfish

Benthosema glaciale

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
180 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Near surface to 225 m at night [1], mainly 30–90 m [7]; descending to 275–1,456 m during day [1, 5], mainly 

350–450 m [7].

Habitats and Life History

Eggs—Size: 0.75–0.80 mm [8].Time to hatching: Unknown. Habitat: Pelagic [9]. 

Larvae—Size at hatching: Unknown. 5 mm or less [8]. Size at juvenile transformation: 11–15 mm [8]. Days to 

juvenile transformation: Unknown. Habitat: Pelagic [9]. 

Juveniles—Age and size: Age unknown. 1.1–5.0 cm [6, 8]. Habitat: Epipelagic to mesopelagic [1]. 

Adults—Age and size at first maturity: 2–3 years and 4.5–5.0 cm [6]. Maximum age: 8 years in Barents Sea 

[6]. Maximum size: As long as 10.3 cm (reported as both TL and SL) [10], usually less than 7.0 cm [1]. Habitat: 

Epipelagic to mesopelagic, typically offshore [1]. 

Substrate—Unknown. 

Physical/chemical—Temperature: Common between 4 and 16 °C in northwest Atlantic Ocean. Has been 

captured at temperatures of -0.1–21 °C [5]. Salinity: Marine [9].

Behavior

Diel—Mesopelagic by day, epipelagic by night [1]. 

Seasonal—Unknown. 

Reproductive—Spawns pelagically [6]. 

Schooling—Forms schools [6]. 

Feeding—Filter feeder [6]. Feeds year-round, but activity is most intensive in spring and summer [11].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous, separate sexes [9]. 

Spawning season—June–July in Barents Sea [6]. Early spring off Nova Scotia, Canada [8]. 

Fecundity—750–800 eggs [6].

Benthic and reproductive distribution of Glacier Lanternfish (Benthosema glaciale). 

3-4-2g_GlacialLanternfish

Open ocean Shore

Depth, in meters

1,250

1,000

750

500

250

0

1,250

1,000

750

500

250

0

Night distribution

No data for this species

Day distribution

Adults and juveniles

Some fish 

to 1,250 m

Benthosema glaciale

Glacier Lanternfish

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive Distribution

Spawning, eggs, and larvae

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Depth ranges of spawning,

eggs and larvae is uncertain. 

Potential locations are based 

on depth ranges of adults

Data from outside 

U.S. Beaufort-Chukchi Seas
Glacier Lanternfish 181

Food and Feeding

Food items—Mainly copepods and euphausiids [6, 7, 11]. 

Trophic level—2.99 (standard error 0.29) [12].

Biological Interactions

Predators—Leach’s Storm-Petrels off Newfoundland, Canada [13]. For lanternfish in general, predators are 

squids, larger fishes, and marine mammals [1]. 

Competitors—As one of the few mesopelagic species in the U.S. Chukchi Sea, Glacier Lanternfish probably 

have few fish competitors, especially at depth. Arctic Cod and Ice Cod co-occur with Glacier Lanternfish and 

may compete for zooplankton prey.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (K=0.20–0.45; t

m=2–3; tmax=8; fecundity=700) [12].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Glacier Lanternfish are not currently harvested commercially.

Potential Effects of Climate Change

Unknown. 

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators.

References Cited

Gjøsaeter, J., 1981, Growth, production and reproduction of the Myctophid fish Benthosema glaciale from western Norway and 

adjacent seas: Fiskeridirektoratets skrifter, Serie Havundersøkelse, v. 17, no. 3, p. 79–108. [8]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]
182 Alaska Arctic Marine Fish Ecology Catalog

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Sameoto, D.D., 1988, Feeding of lantern fish Benthosema glaciale off the Nova Scotia shelf: Marine Ecology Progress Series, v. 

44, p. 113–129. [7]

Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, A., 

Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 2011, 

Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p. [6]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Salvanes, A.G.V., and Kristoffersen, J.B., 2001, Mesopelagic fishes, in Steels, J., and others, eds., Encyclopedia of ocean 

sciences: Academic Press, p. 1,711–1,717.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

7. Sameoto, D.D., 1988, Feeding of lantern fish Benthosema glaciale off the Nova Scotia shelf: Marine Ecology Progress 

Series, v. 44, p. 113–129.

8. Gjøsaeter, J., 1981, Growth, production and reproduction of the Myctophid fish Benthosema glaciale from western Norway 

and adjacent seas: Fiskeridirektoratets skrifter, Serie Havundersøkelse, v. 17, no. 3, p. 79–108.

9. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

10. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

11. Gjøsaeter, J., 1973, The food of the myctophid fish, Benthosema glaciale (Reinhardt), from western Norway: Sarsia, v. 52, 

no. 1, p. 53–58.

12. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

13. Hedd, A., and Montevecchi, W.A., 2006, Diet and trophic position of Leach’s storm-petrel Oceanodroma leucorhoa during 

breeding and moult, inferred from stable isotope analysis of feathers: Marine Ecology Progress Series, v. 322, p. 291–301.
Ice Cod 183

Ice Cod to Pacific Cod

Ice Cod (Arctogadus glacialis) 

(Peters, 1872)

Family Gadidae 

Note on taxonomy: Evidence from morphology and molecular 

genetics demonstrates that Arctogadus borisovi (Dryagin, 1932) is 

a junior synonym of A. glacialis [1]. Data on fish originally 

identified as A. borisovi are included here. Commmonly referred to 

as Polar Cod in North America.

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: The ecological role of the species in marine 

ecosystems of the U.S. Chukchi and Beaufort Seas is not as significant as Polar and Saffron Cod. 

Physical Description/Attributes: An olive brown to bluish gray cod with darker fins and head. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 291–292) [2]. Swim bladder: Present; no otophysic 

connection [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Beaufort [2] and Chukchi Sea [3, 4]. Worldwide, circumpolar, northward to at least 81°41’N; Arctic Canada south 

to southern tip of Greenland, east through Barents Sea to East Siberian Sea and Chukchi Sea [2–4].

Ice Cod (Arctogadus glacialis) 221 mm, Chukchi Borderland, 

2009. Photograph by C.W. Mecklenburg, Point Stephens 

Research.
184 Alaska Arctic Marine Fish Ecology Catalog

Geographic distribution within Arctic Outer Continental Shelf planning areas [5] of Ice Cod (Arctogadus 

glacialis) based on review of published literature and specimens from historical and recent collections [3, 4]. 

tac14-5222_fig3-5-1m_Arctogadus_glacialis

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Ice Cod

Arctogadus glacialis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Relative Abundance: Rare in U.S. Beaufort Sea (two specimens captured north of Point Barrow) [2] and Chukchi Sea (one 

specimen found on beach at Wainwright) [4].Abundant to at least as far eastward to deep waters off Tuktoyaktuk Peninsula and 

off Capes Bathurst and Parry, Canada [6–8].
Ice Cod 185

Depth Range: 5–930 m, on continental shelf and upper slope [1, 2]. Highest abundance is found off Europe at 300–400 m [9]. 

In northeast Greenland fjords, abundant at 120–575 m [10]. Eggs and larvae are pelagic [11] but specific depths unknown. 

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Size: Unknown. Habitat: Pelagic [11]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [11]. 

Juveniles—Age and size: Unknown. Habitat: Cryopelagic and benthic [9]. 

Adults—Age and size at first maturity: Unknown. Females 25–26 cm long (TL) have been found with ripening 

gonads [12]. Maximum age: At least 11 years [13]. Maximum size: 60 cm TL [2] and 1.2 kg [12]. Habitat: 

Nearshore to well offshore [4, 8, 14]. Cryopelagic and benthic [5], throughout the water column (including near 

the seafloor) as well as under ice and within ice cracks [10, 15, 16]. 

Substrate—Unknown. 

Physical/chemical—Temperature: -1.7 to about 4 °C [10], may prefer temperatures of about 1 °C or less [1, 10]. 

Salinity: Marine, estuarine, and occasionally fresh waters from near the coast to well offshore [8, 14]. 

Ice dependence—Although characterized as an ice-associate, also found well away from ice, sometimes in large 

numbers [10, 17].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Forms schools [2]. 

Feeding—Opportunistic pelagic feeder [9].

Populations or Stocks

There have been no studies.

Reproduction mode

Mode—Oviparous [11]. 

Spawning season—Ripe fish were observed in October and during the summer in the European Arctic [9, 10]. 

Fecundity—Unknown.

Benthic and reproductive distribution of Ice Cod (Arctogadus glacialis).

3-5-1_IceCod

Open ocean Shore

Depth, in meters 1000

800

600

400

200

0

1000

800

600

400

200

0

Adults and juveniles

Some fish to 825 m

Arctogadus glacialis

Ice Cod

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Spawning

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Depth range of spawning is uncertain. 

Potential location is based on depth 

range of adults

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Eggs and larvae
186 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Crustaceans (for example, mysids, copepods, and amphipods), fishes, and polychaetes comprise 

much of the diet of this species. Fishes assume a greater part of the diet in larger cod [15, 18, 19]. 

Trophic level—3.82 (standard error 0.61) [20].

Biological Interactions

Predators—Commonly, bearded seals and narwhals in the Canadian Arctic [21, 22]. 

Competitors—Unknown. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [20]. 

Traditional and Cultural Importance

None reported. Form only a small part of the subsistence fisheries in the Canadian Arctic [8]. Commercially 

fished for fishmeal and oil in Norway, Greenland and northern Siberia [23].

Commercial Fisheries

Currently, Ice Cod are not commercially harvested.

Potential Effects of Climate Change

Unknown.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Although information should improve with 

increased sampling, the role of this species in the gadid assemblage and how this might change with global 

warming is of research interest. Spawning areas and other important habitats remain to be described.
Ice Cod 187

References Cited

Aschan, M., Karamushko, O.V., Byrkjedal, I., Wienerroither, R., Borkin, I.V., and Christiansen, J.S., 2009, Records of the 

gadoid fish Arctogadus glacialis (Peters, 1874) in the European Arctic: Polar Biology, v. 32, no. 7, p. 963–970. [9]

Christiansen, J.S., ed., 2003, TUNU-1 Expedition—The fish fauna of the NE Greenland fjord systems—Technical report: 

Tromsø, Norway, University of Tromsø, Norwegian College of Fishery Science, Institute of Aquatic Resources, 33 p. [10]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p. [8]

Bibliography

1. Jordan, A.D., Møller, P.R., and Nielsen, J.G., 2003, Revision of the Arctic cod genus Arctogadus: Journal of Fish Biology, v. 

62, no. 6, p. 1,339–1,352.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Byers, S.C., and Kashino, R.K., 1980, Survey of fish populations in Kugmallit Bay and Tuktoyaktuk Harbour, Northwest 

Territories: Dobrocky Seatech Limited Ocean Services for Dome Petroleum Limited, Calgary, Alberta, 20 p. 

7. Arctic Laboratories Limited, 1987, Beaufort Ocean dumpsite characterization: Prepared for Environment Protection, 

Conservation and Protection, Environment Canada, Yellowknife, Northwest Territories, by Arctic Laboratories Limited and 

LGL Limited, 135 p.

8. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

9. Aschan, M., Karamushko, O.V., Byrkjedal, I., Wienerroither, R., Borkin, I.V., and Christiansen, J.S., 2009, Records of the 

gadoid fish Arctogadus glacialis (Peters, 1874) in the European Arctic: Polar Biology, v. 32, no. 7, p. 963–970.

10. Christiansen, J.S., ed., 2003, TUNU-1 Expedition—The fish fauna of the NE Greenland fjord systems—Technical report: 

Tromsø, Norway, University of Tromsø, Norwegian College of Fishery Science, Institute of Aquatic Resources, 33 p.

11. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

12. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.
188 Alaska Arctic Marine Fish Ecology Catalog

13. Boulva, J., 1979, Comparison of the Arctic cod (Boreogadus saida), the polar cod (Arctogadus glacialis), and the toothed 

cod (A. borisovi): Quebec City, Quebec, Government of Canada, Fisheries and Oceans, Fisheries Sciences Division, 

CAFSAC Research Document 79/50, 12 p.

14. Berg, L.S., 1949, Freshwater fishes of the U.S.S.R. and adjacent countries, volume 3 (4th ed.): Moscow, Academy of 

Sciences of the U.S.S.R. Zoological Institute, 250 p. [Translated from Russian by Israel Program for Scientific Translations, 

Jerusalem, IPST Catalog No. 743.]

15. Walters, V., 1961, Winter abundance of Arctogadus glacialis in the polar basin: Copeia, no. 2, p. 236–237.

16. McAllister, D.E., 1975, Ecology of the marine fishes of Arctic Canada, in Proceedings of the Circumpolar Conference on 

Northern Ecology, September 15–18, 1975: Ottawa, National Research Council of Canada, p. II-49–II-65.

17. Borkin, I.V., and Mel’yantsev, R.V., 1984, New data on the distribution of polar cod, Arctogadus glacialis (Gadidae), in the 

Arctic region: Journal of Ichthyology, v. 24, no. 1, p. 101–103.

18. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

19. Joensen, J., 2008, Comparative feeding ecology of the sympatric cod fishes Arctogadus glacialis and Boreogadus saida in 

north-east Greenland evaluated from diet and stable isotope analyses: Tromsø, Norway, University of Tromsø, Master’s 

thesis. 54 p.

20. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

21. Finley, K.J., and Gibb, E.J., 1982, Summer diet of the narwhal (Monodon monoceros) in Pond Inlet, northern Baffin Island: 

Canadian Journal of Zoology, v. 60, no. 12, p. 3,353–3,363.

22. Finley, K.J., and Evans, C.R., 1983, Summer diet of the bearded seal (Erignathus barbatus) in the Canadian High Arctic: 

Arctic, v. 36, no. 1, p. 82–89.

23. Cohen, D.M., Inada, T., Iwamoto, T., and Scialabba, N., 1990, FAO species catalogue, volume 10—Gadiform fishes of the 

world (Order Gadiformes)—An annotated and illustrated catalogue of cods, hakes, grenadiers and other gadiform fishes 

known to date: Rome, Food and Agriculture Organization of the United Nations, FAO Fisheries Synopsis, no. 125, v. 10, 

442 p.
Arctic Cod 189

Arctic Cod (Boreogadus saida)

(Lepechin, 1774)

Family Gadidae

Colloquial Name: Iñupiat: Iqalugaq, Uugaq [1]. Tomcod [2].

Notes on Taxonomy: Boreogadus saida is referred to as both 

“Arctic Cod” and “Polar Cod” in North American scientific 

literature. The American Fisheries Society and Society of 

Ichthyologists and Herpetologists recommend using “Polar 

Cod” for Boreogadus saida to bring consistency with European 

conventions and this recommendation is gaining in acceptance 

in contemporary reporting and publications. We chose to use 

“Arctic Cod” for Boreogadus saida in this report to ensure consistency with the vast majority of Alaskan literature and to avoid 

confusion with Ice Cod, which has been referred to as Polar Cod by North American researchers. Anyone using literature that 

does not specify the scientific name must read carefully to decide which species is meant. However, A. glacialis is rare in Arctic 

Alaska waters, and most references to Polar Cod from that region that do not provide the scientific name will refer to B. saida.

Ecological Role: Arctic Cod play a vital role in anchoring Arctic food webs in the U.S. Chukchi and Beaufort Seas. The small 

fish is one of the main consumers of plankton that flourish around sea ice. Arctic Cod compose 92 percent of all fish in numbers 

and 80 percent in weight in a 2008 western Beaufort Sea fish survey [3], and this species comprises the forage base for a wide 

range of marine mammals and birds. A recent model predicted a mass loss of most Arctic Cod within 30 years because of rising 

temperatures and receding ice pack. Key interactions between sea ice, Arctic Cod biology, and marine ecosystem function must 

be better understood to identify possible effects of climate change and cumulative effects of human activities.

Physical Description and Attributes: Brownish back and sides with violet or yellowish sheen covered with tiny black dots, and 

silvery white lower sides and belly. Fins are dusky yellow or gray, and dorsal and caudal fins are edged in white. For specific 

diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 290) [4]. Swim bladder: present. Antifreeze 

glycoproteins in blood serum: Present [5].

Range: Throughout U.S. Chukchi and Beaufort Seas from very shallow, neritic waters to well offshore (although abundance 

hundreds of kilometers offshore is poorly known) [6–9]. A circumpolar species, documented in Chukchi Sea northward nearly 

to the North Pole at 88°26′N, 126°26′E. In continental shelf waters, west and south of U.S. Chukchi Sea from Siberian Arctic 

to Olyutorskiy Bay in western Bering Sea and to Bristol Bay in eastern Bering Sea; and east of U.S. Beaufort Sea continuous 

throughout the Canadian Beaufort Sea [10, 11].

Arctic Cod (Boreogadus saida) 174 mm TL, Chukchi Sea, 2004. 

Photograph by B.A. Sheiko and C.W. Mecklenburg, Russian 

Academy of Sciences and Point Stephens Research.
190 Alaska Arctic Marine Fish Ecology Catalog

Relative Abundance: Very abundant in U.S. Chukchi and Beaufort Seas [9, 12–15].

tac14-5222_fig3-5-2m_Boreogadus_saida

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Cod

Boreogadus saida

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution within Arctic Outer Continental Shelf Planning Areas [16] of Arctic Cod (Boreogadus 

saida) based on review of published literature and specimens from historical and recent collections [11, 17]. 
Arctic Cod 191

Depth Range: Larvae: Maximum depth unknown. Surface to at least 20 m [12]. Juveniles: Maximum depth unknown. Near 

surface to 75 m [9, 18]. Older juveniles and adults: Maximum depth could be 930 m (the species’ maximum recorded depth) but 

such depths have not been sampled in the U.S. Chukchi and Beaufort Seas. Found throughout the water column, abundant from 

surface waters to at least 400 m [9, 12, 19] and deeper (500 to 1,000 m in the United States Beaufort Sea, Kathleen Wedemeryer, 

Bureau of Ocean Energy Management, Alaska OCS Region, oral commun., October 13, 2015). Elsewhere, Arctic Cod are found 

from barely subtidal waters to depths of 930 m [20]. Spawning: Shallow nearshore waters and under nearshore ice [21, 22].

Benthic and reproductive distribution of Arctic Cod (Boreogadus saida).

Boreogadus saida

Arctic Cod

Benthic distribution

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Spawning

Some spawning occurs in 

very shallow nearshore water

and under nearshore ice .

 Data from outside

U.S. Beaufort-Chukchi Seas

Reproductive distribution

Eggs and larvae Juveniles in study area

Depths reported common 

in study area

Some fish to 930 m

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

Adults and

older juveniles

Overall benthic range.

Specific depth range of

either juveniles or adults

is unknown 

 Maximum depth data from

outside U.S. Beaufort-Chukchi Seas

Habitats and Life History

Although Arctic Cod are assumed to both spawn and live under ice in Arctic Alaskan waters, virtually no surveys 

have been conducted to determine the importance of this habitat to Arctic Cod in the Alaskan Arctic. Throughout 

its range, Arctic Cod occupy a remarkably wide breadth of habitats. They occupy all parts of the water column 

in estuaries and off river mouths and in shallow subtidal waters, and are found many hundreds of kilometers off 

the coast [6, 21, 23, 24]. These fish are often associated with ice although they have been captured during spring 

and summer in the northern Bering Sea at least 300 km (186 mi) away from the nearest floe or pack ice and are 

abundant along the U.S. Chukchi and Beaufort Seas in seasonally ice-free areas [6, 25]. However, they can be 

found in very large numbers under ice and are often seen in cracks, crevices, and in melt-water ponds on the ice 

[20, 26, 27]. 

Eggs—Size: 1.5–1.9 mm [28, 29]. Time to hatching: Unknown. Elsewhere, between 26–90 days; highly variable, 

apparently dependent on water temperature [30, 31]. Habitat: Planktonic. Location unknown.

Larvae—Size at hatching: 6 mm [32] Size at juvenile transformation: 27–50 mm [30, 33]. Days to juvenile 

transformation: Unknown. In the Russian Arctic, the larval stage lasts about 2 months [30]. Habitat: Pelagic. 

Location poorly understood. Elsewhere, found under ice [34]. 

Juveniles—Habitat: Poorly known. Some pelagic juveniles are found in near-surface waters away from ice [18, 

35]. Some young-of-the-year recruit from the plankton directly into inshore habitat, but it is likely that others 

remain well offshore (to at least 175 km off Prudhoe Bay) [6, 18] in near-surface waters [6, 32]. Large numbers 

of juveniles were found in shallow U.S. Beaufort Sea lagoons [36]. Elsewhere, juveniles are common under ice 

[20, 26, 27, 37]. However, in the Alaskan Arctic, they also are very abundant in the summer well away from 

ice [6, 8, 38]. In other areas, in offshore waters and in the absence of ice, smaller fish tend to inhabit shallower 

depths in the water column than do larger individuals [32, 39, 40]. 

Adults—Age and size at first maturity: A few fish, possibly only males, are mature at 1year and around 100 mm 

FL. Most fish mature at –3 years (120 mm FL and larger) and males may mature about 1 year earlier than 

females [6, 19]. Maximum age: In study area, 7–8 years [15]. Maximum size: 46 cm TL [41]. Habitat: Adults are 

common under ice [20, 26, 27, 37]. However, in the Alaskan Arctic, they also are very abundant in the summer 

well away from ice [6, 8, 38].
192 Alaska Arctic Marine Fish Ecology Catalog

Physical/Chemical—Temperature: Preferred temperatures poorly known, but probably about –1.5–5 °C [12, 

13, 42], although sometimes they are abundant in waters as warm as 10 °C [8, 43]. Documented between -2.0 

and 13.5 °C [43, 44]. Elsewhere, to -2.1 °C [45]. In Bering Sea, mainly -2.0 to -0.7 °C [45]. In the Canadian 

High Arctic, larvae are reported to develop only at temperatures less than 3.0 °C [46] and in the Russian Arctic, 

less than 5 °C [30]. Juveniles may favor warmer waters than adults [8, 39]. Salinity: Documented from 0 to 32.6 

practical salinity units (psu) [10, 43, 44] and elsewhere to 34.9 psu [17]. All life stages live in marine waters. 

Although occasionally abundant in brackish waters [43], low abundance in such areas as the Mackenzie River 

estuary may reflect general avoidance of very low salinity waters [47].

Ice Dependence—Although it is clear that many Arctic Cod live under ice, it is unclear whether these fish are ice 

associated or ice dependent [27, 48]. The hypothesis that Arctic Cod are ice dependent is derived from a number 

of inconclusive observations. Examples include:

4. Known to spawn along the ice edge [49] and under ice [23, 50]. However, whether spawning is limited to ice or a near-ice 

area is unknown. For instance, Arctic Cod eggs have been taken in May around the ice-free Pribilof Islands, Alaska [51]. 

In addition, surveys of possible spawning in ice-free areas have not been conducted during the overwinter, for instance, the 

Bering Sea.

5. Juveniles and adults are very abundant under ice [52], but can be extremely abundant during summer in relatively warm 

water and ice-free conditions [9, 12, 32, 44]. 

6. The relationship between ice densities, water temperatures, and fish growth and survival is uncertain. In the Greenland Sea, 

larvae living in low-ice, relatively warm waters survived better than those in thicker ice and colder (<0 °C) temperatures, 

implying that the lengthening of the ice-free season may result in improved recruitment and larger populations in Arctic 

Cod in the short term [50]. This has also been noted among polynyas in Arctic Canada [48]). In addition, juvenile and 

adult Arctic Cod in the northeast Chukchi Sea grew fastest in a warmer-water year [15]. However, this position has 

been challenged by some authors based on the hypothesis that Arctic sea warming will reduce sea ice habitat and allow 

sub-Arctic or temperate taxa to replace this species [50]. 

Behavior

Diel—Unknown. Elsewhere, from January to April, post-spawning adults in the eastern Beaufort Sea rise in the 

water column at night [52]. 

Seasonal—Poorly understood [6]. Current information suggests that throughout their range, fish move into 

nearshore waters in the summer. Precise time and intensity varies between locations and between years at the 

same location [6, 13, 53, 54]. In the U.S. Beaufort Sea, relatively scarce in shallow waters immediately after 

ice-out, but move into shallow waters as the season progresses [6, 7, 53]. However, ringed seal diet studies in the 

nearshore of Arctic Alaska [55], and other fish surveys [44, 56], imply that at least some Arctic Cod overwinter in 

nearshore waters under ice. In nearshore waters, schools can move quickly through an area [44, 57] or reside in 

the same location for weeks [58]. Proportion of population migrating into shallow waters is unknown; many fish 

might not migrate into the shallowest waters [19, 38]. In summer, Arctic Cod are by far the most abundant fish in 

Alaska Arctic nearshore waters [14, 15]. For example, estimates of summer cod abundances in Simpson Lagoon 

have been as high as 12–27 million fish [44].

Data regarding the environmental parameters driving inshore migrations are often contradictory. For instance, 

in the U.S. Chukchi Sea, catches increased when water temperatures rose and salinities decreased [38], but was 

the opposite in the Sagavanirktok River Delta of the U.S. Beaufort Sea [59]. In Prudhoe Bay of the U.S. Beaufort 

Sea, highest densities were noted in frontal areas bordering low salinity and high temperature surface waters, 

and high salinity and low temperature bottom waters, perhaps an area of high productivity [60]. Another U.S. 

Chukchi Sea study found no environmental-parameter-associated abundance; authors hypothesized that food 

availability might underlay fish movements [15].

Thus far, the most complete study of winter behavior was in Franklin Bay, eastern Canadian Beaufort Sea. It was 

documented that after spawning during the early winter (perhaps over deep waters in the Amundsen Gulf; (D. 

Benoit, Université Laval, 2010 ), very large numbers of fish either migrated, or were passively carried, into 180 

m or deeper depths and did not feed during this time. Migration out of these waters began with an abrupt upward 

migration and coincided with phytoplankton blooms and the onset of feeding [52]. Similar work in the nearby 

Amundsen Gulf demonstrates a similar pattern [61].

Reproductive—Poorly known. In the Arctic in general, spawning occurs near the bottom along the ice edge 

[49] and under ice [23], whether limited to under-ice areas is unknown. For instance, eggs have been found in 
Arctic Cod 193

May around the ice-free Pribilof Islands [51]. The winter (spawning-season) behavior throughout the Arctic is 

very poorly known. In the U.S. Chukchi and Beaufort Seas, at least some fish spend winters under nearshore ice 

(presumably spawning) [38, 44, 56, 62]. However, whether the bulk of the population overwinters and spawns 

in shallow waters is unknown. For instance, spawned-out cod were reported both near the coast and 175 km off 

Prudhoe Bay [6]. In the autumn and winter, large, spawning-oriented migrations occurred in the Russian Arctic 

and spawning in the Barents Sea may have occurred from near shore to hundreds of kilometers off the coast [49]. 

Schooling—Schooling behavior under ice has not been studied. In ice-free areas, very large schools are formed, 

often millions of individuals [6, 44]. This species often schools by size class [8]. The amount of cohesion of fish 

schools in Arctic Alaska is unknown, although in the Canadian Arctic some schools stay together for at least 1–2 

months in summer [24, 58]. 

Feeding—Prior to spawning, a few nearly ripe fish in Simpson Lagoon were still feeding [44], whereas in the 

Canadian Arctic, feeding ceases for several months beforehand [52].

Population or Stocks

Initial research underway in U.S. Chukchi and Beaufort Seas and Arctic Ocean. Microsatellite markers imply 

some population structure among Chukchi Sea, Canadian and Siberian Arctic Cod, and potentially low 

differentiation between those from Hudson Bay and the eastern Beaufort Sea [63]. The genetics research 

indicates very little structuring across the United States Beaufort Sea with respect to the three dichotomies of 

east/west; coastal/slope; and riverine/marine water influence (Kathleen Wedemeyer, Bureau of Ocean Energy 

Management, oral commun,, October 13, 2015). A single circum-artctic population with only minor differences 

is currently hypothesized. 

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external. 

Spawning season— Poorly understood. Over the species’ entire geographic range, spawning occurs under 

ice floes from November to at least April, perhaps peaking in January and February in the Beaufort Sea [6, 15, 

31, 49]. The capture of spawned-out adults in May, 175 km off Prudhoe Bay, implies that some spawning may 

continue well into the spring [6]. The capture of newly hatched larvae in July in the northeastern Chukchi Sea

[33] and as late as July and August in Tuktoyaktuk Harbor, Northwest Territories, and near Baffin Bay [64, 65], 

also may imply late spawning. However, eggs fertilized in February will often remain as larvae into July [30]. 

Fecundity—Unknown. Elsewhere, 9,000–33,251 eggs, apparently in one batch [28, 29]. Females may not spawn 

every year [6]. However, in a laboratory study, several females spawned in two successive years, implying that 

some fish are capable of spawning more than once in their lives and in sequential years [46].

Food

Food items—Larvae: Unknown. In Hudson Bay, pelagic larvae under ice feed on nauplii and eggs of those 

copepods feeding on under-ice phytoplankton [34]. Juveniles and Adults: Diets vary with fish size and location, 

although epibenthic or pelagic crustaceans (for example, mysids, isopods, copepods, gammarid and hyperiid 

amphipods, and shrimps), as well as larval fish, polychaetes, chaetognaths, and small fishes (such as other Arctic 

Cod), are important. Fish living under ice often target ice-associated crustaceans such as amphipods [6, 57, 

 66-68]. Feed primarily on copepods and amphipods in northern Bering Sea [69]. Diets also may vary with 

 season. In Simpson Lagoon of the Alaskan Beaufort Sea, mysids, amphipods, isopods were most important during

 the summer and mysids dominated during the winter [44]. 

 Trophic level—3.6 [70].

Biological Interactions

Predators—Arctic Cod are an extremely important prey for a wide range of predators and are possibly the 

most important forage fish in the U.S. Chukchi and Beaufort seas. They are consumed by at least 8 fish species, 

17 bird species, and 3 marine mammal species [19, 38, 67, 71–75]. The major predators of Arctic Cod in the 

Alaska Beaufort Sea are considered to be, in order of importance, Arctic Cod (cannibalism), ringed seals, Beluga 

whales, and seabirds (particularly Black-legged Kittiwake, Thick-billed Murre, Ivory Gull, Black Guillemot, 

Glaucous Gull, loons, Ross’ Gull, Arctic Tern, and Sabine’s Gull) [67]. Almost all studies were conducted during 

summer months, although it has been shown that in the Alaskan High Arctic they form a major part of the diet 

of ringed seals (particularly important to pups) throughout the year [55, 67] and bearded seals in the U.S. 

Chukchi Sea from at least November through June [71]. Arctic Cod appear to be particularly susceptible to beach 

strandings, caused by predators or storms [6, 58]. 
194 Alaska Arctic Marine Fish Ecology Catalog

Competitors—Likely competitors are other schooling midwater feeders, particularly Walleye Pollock, but also 

Dolly Varden, whitefish species, Capelin, and Pacific Sand Lance. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (K=0.22; t

m=2-5; Fecundity =30,000) [76].

Traditional and Cultural Importance

In the past, this was a fairly important human subsistence species in the Alaskan High Arctic. For instance, 

it was reported that Arctic Cod were heavily fished through the ice off Barrow in the winter [77]. Over time 

and today, Arctic Cod are of only limited importance as food fish [1, 78–80]. Arctic Cod appear to be of more 

importance in Canadian subsistence fisheries of the Barrow Strait and Hudson Bay where they are widely caught 

and consumed [58, 81]. 

Commercial Fisheries

Arctic Cod are not commercially harvested in the U.S. Chukchi and Beaufort Seas. The commercial fishery for 

Arctic Cod is small and limited to Russian vessels fishing primarily in the northwest Russian Arctic [82].

Potential Effects of Climate Change

Climate change may influence the numbers of Arctic Cod through a number of mechanisms. (1) Assuming that 

this species is in some way ice-dependent, a poleward shift in distribution would be expected with retreating 

ice. (2) There is some evidence that survivorship of Arctic Cod larvae increases with earlier ice break-up, 

more frequent winter polynyas, a warmer (ocean) surface layer, and increased river discharge [83], all possible 

effects of warming conditions. (3) Arctic Cod coming under increased competition for resources from some 

northward-migrating species would be expected, particularly from Saffron Cod and possibly Walleye Pollock. 

(4) Greater periods of ice-free conditions likely will alter predation patterns, but in ways that are not yet 

predictable. Receding ice may increase predation. In Resolute Bay, North West Territories, fish under heavy 

ice cover were less aggregated than when the bay was relatively ice-free. When ice drifted into the bay, fish 

would move under it [84]. In Hudson Bay, a sharp decrease in the abundance of Arctic Cod coincided with an 

approximately 50 percent decrease in summer ice cover [85]. However, the millions of Arctic Cod inhabiting the 

ice-free Simpson Lagoon in the summer do not appear to suffer heavy predation [44]. Reduced ice pack can be 

argued to cause an increase or decrease predation depending on predator. For instance, reduced ice pack would 

decrease resting habitat for seals, while making Arctic Cod perhaps more available to cetaceans or seabirds 

[48]. (5) Food availability and growth rates will change, although the direction and intensity of this change are 

unknown. As an example, fish in the northeast Chukchi Sea grew fastest in warmer water years [15] and larvae 

residing in the low-ice, relatively warm waters in Greenland survived better than those under thicker ice [50]. 

(6) Effects on Arctic Cod predators are unknown, but may be substantial. For instance, retreating pack ice near 

Point Barrow led to reduced Arctic Cod availability for Black Guillemots and subsequent reductions in nestling 

growth and brood size [86]. (7) Effects of predation by Arctic Cod on prey are unknown but the species midlevel role in transferring energy from low to high trophic levels is hypothesized to be significant. Local effects 

of Arctic Cod predation on prey concentrations also may be significant. For instance, feeding by large schools of 

adult Arctic Cod in the Canadian Arctic may be sufficiently intense as to cause localized depletion of zooplankton 

[24]. The effects of possible changes in Arctic Cod distribution and abundance, in association with climate 

warming, may have profound, cascading effects on the Arctic marine ecosystems. The effects of increasing 

ocean acidification on Arctic Cod food webs dynamics and developmental biology are of concern. The protocols 

for capturing, transporting, breeding, and rearing larvae through adult stages in the laboratory have been tested 

and described [87, 88] making empirical studies of thermal sensitivity to warming using an Arctic Cod model 

possible. New information is available describing the thermal limits of cardiac function on Arctic Cod [89, 90], 

effects of warming and ocean acidification on metabolism and performance on Arctic Cod and Atlantic Cod 

(Gadus morhua) [91], and temperature-dependent growth and swimming behaviors of Arctic Cod, Saffron Cod, 

Walleye Pollock, and Pacific Cod [92]. In general, the results suggest optimal food conversion for juvenile Arctic 
Arctic Cod 195

Cod in Cold waters (0 °C), near optimal growth at 5 °C, and diminished growth and condition with increasing 

temperatures above this (detrimental effects above 16 °C). Differential acute effects of warming on larvae and 

adults further suggest the potential role thermal limitations of younger-aged cod may have on the Arctic Cod 

distribution in coastal waters.

Areas for Future Research [A]

Considering the species central role in Arctic marine ecosystems dynamics, relatively little focused research 

attention has been given to Arctic Cod in Arctic Alaska. In particular, the role of sea ice in the species’ life cycle, 

though speculated, is not well understood. The location and timing of spawning locations and presence of stock 

structures are unknown. Information is needed regarding population movements and behaviors, particularly 

during winter months, and with respect to the relative important habitats in slope, shelf, and nearshore, and 

deeper areas of the Canada Basin. The latter need is of particular importance because a recent model predicted 

a mass extinction of most Arctic Cod within 30 years [93]. However, the model appears to be at least partially 

based on the assumption that there are no Arctic Cod well offshore of northern Alaska, although no surveys have 

been conducted there. The use of Autonomous Underwater Vehicles to investigate Arctic Cod ecology should be 

explored. There needs to be new and continued empirical research to determine the seasonal effects of changing 

temperatures, ocean acidification, and ice coverage on the reproduction ecology and population growth and 

condition of Arctic Cod. An additional experimental priority is for toxicological research on the potential effects 

of spilled, dispersed, and weathered oil on Arctic Cod under Arctic conditions. Accurate assessments of species 

interactions and effects of human developments and climate changes will require that the population dynamics of 

the species are understood and that abundance patterns and population parameters are monitored over time.

References Cited

Benoit, D., Simard, Y., and Fortier, L., 2008, Hydroacoustic detection of large winter aggregations or Arctic cod (Boreogadus 

saida) at depth in ice-covered Franklin Bay (Beaufort Sea): Journal of Geophysical Research, v. 113, no. CO6S90, 9 p. [52]

Craig, P.C., Griffiths, W.B., Haldorson, L., and McElderry, H., 1982, Ecological studies of Arctic cod (Boreogadus saida) in 

Beaufort Sea coastal waters—Alaska: Canadian Journal of Fisheries and Aquatic Sciences, v. 39, no. 3, p. 395–406. [6]

Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, Simpson 

Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649. [44]

Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., and Colonell, J.M., 1996, The 1995 fish and oceanography study in 

Mikkelsen Bay, Alaska: Anchorage, Alaska, Prepared by LGL Alaska Research Associates and Woodward-Clyde Consultant 

for BP Exploration (Alaska) Inc., 102 p. plus apps. [14]

Fortier, L., Sirois, P., Michaud, J., and Barber, D., 2006, Survival of Arctic cod larvae (Boreogadus saida) in relation to sea ice 

and temperature in the Northeast Water Polynya (Greenland Sea): Canadian Journal of Fisheries and Aquatic Sciences, v. 63, 

no. 7, p. 1,608–1,616. [50]

Frost, K.J., and Lowry, L.F., 1984, Trophic relationships of vertebrate consumers in the Alaskan Beaufort Sea, in Barnes, P.W., 

Schell, D.M., and Reimnitz, E., eds., The Alaska Beaufort Sea—Ecosystems and environments: San Diego, Academic Press, 

p. 382–401. [67]

Gillispie, J.G., Smith, R.L., Barbour, E., and Barber, W.E., 1997, Distribution, abundance, and growth of Arctic cod in the 

Northeastern Chukchi Sea, in Reynolds, J.B., ed., Symposium 19, Proceedings of the Fish Ecology in Arctic North America 

Symposium, Fairbanks, Alaska, May 19–21, 1992: Fairbanks, Alaska, American Fisheries Society, p. 81–89. [15]

Jarvela, L.E., and Thorsteinson, L.K., 1999, The epipelagic fish community of Beaufort Sea coastal waters, Alaska: Arctic, v. 52, 

no. 1, p. 80–94. [8]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [4]

Quast, J.C., 1974, Density distribution of juvenile Arctic cod, Boreogadus saida, in the eastern Chukchi Sea in the fall of 1970: 

Fishery Bulletin, v. 72, no. 4, p. 1,094–1,105. [32]
196 Alaska Arctic Marine Fish Ecology Catalog

Bibliography

1. Craig, P.C., 1989b, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers University 

of Alaska, no. 24, p. 131–152.

2. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

3. Logerwell, E.A., and Rand, K.M., 2010, Beaufort Sea marine fish monitoring 2008—Pilot survey and test of hypotheses: 

Seattle, Washington, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Alaska 

Fisheries Science Center, Final Report, BOEMRE 2010-048, 262 p.

4. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

5. Power, G., 1997, A review of fish ecology in Arctic North America: American Fisheries Society Symposium, no. 19, 

p. 13–39.

6. Craig, P.C., Griffiths, W.B., Haldorson, L., and McElderry, H., 1982, Ecological studies of Arctic cod (Boreogadus saida) in 

Beaufort Sea coastal waters—Alaska: Canadian Journal of Fisheries and Aquatic Sciences, v. 39, no. 3, p. 395–406. 

7. Wiswar, D.W., West, R.L., and Winkleman, W.N., 1995, Fisheries investigation in Oruktalik Lagoon, Arctic National 

Wildlife Refuge, Alaska, 1986: U.S. Fish and Wildlife Service, Alaska Fisheries Technical Report, no. 30, 38 p.

8. Jarvela, L.E., and Thorsteinson, L.K., 1999, The epipelagic fish community of Beaufort Sea coastal waters, Alaska: Arctic, 

v. 52, no. 1, p. 80–94.

9. Parker-Stetter, S., Horne, J., Logerwell, L., Rand, K., and Weingartner, T. 2010, Assessment of Arctic cod and young-of-theyear fish distributions in the Beaufort Sea [abs.]: Alaska Marine Science Symposium, Book of Abstracts, p. 24.

10. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. 

11. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

12. Crawford, R.E., 2010, Occurrence of Arctic cod shoals along the Chukchi and Beaufort continental slope regions [abs.]: 

Alaska Marine Science Symposium, Book of Abstracts, p. 23.

13. Thedinga, J.F., Johnson, S.W., and Neff, A.D., 2010, Nearshore fish assemblages in the Chukchi Sea near Barrow, Alaska 

[abs.]: Alaska Marine Science Symposium, Book of Abstracts, p. 24.

14. Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., and Colonell, J.M., 1996, The 1995 fish and oceanography 

study in Mikkelsen Bay, Alaska: Anchorage, Alaska, Prepared by LGL Alaska Research Associates and Woodward-Clyde 

Consultant for BP Exploration (Alaska) Inc., 102 p. plus apps.

15. Gillispie, J.G., Smith, R.L., Barbour, E., and Barber, W.E., 1997, Distribution, abundance, and growth of Arctic cod in the 

northeastern Chukchi Sea, in Reynolds, J.B., ed., Symposium 19, Proceedings of the Fish Ecology in Arctic North America 

Symposium, Fairbanks, Alaska, May 19–21, 1992: Fairbanks, Alaska, American Fisheries Society, p. 81–89.

16. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

17. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at http://

caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

18. Parker-Stetter, S.L., Horne, J.K., and Weingartner, T.J., 2011, Distribution of polar cod and age-0 fish in the U.S. Beaufort 

Sea: Polar Biology, v. 34, no. 10, p. 1,543–1,557.
Arctic Cod 197

19. Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western 

Beaufort seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA 

Technical Report NMFS-SSRF-764, 22 p.

20. Jensen, A.S., 1948, A contribution to the ichthyofauna of Greenland: Spoila Zoologica Musei Hauniensis, v. 9, p. 27–57. 

21. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations (1972), p. 161–224.

22. Cohen, D.M., Inada, T., Iwamoto, T., and Scialabba, N., 1990, FAO species catalogue, volume 10—Gadiform fishes of the 

world (Order Gadiformes)—An annotated and illustrated catalogue of cods, hakes, grenadiers and other gadiform fishes 

known to date: Rome, Food and Agriculture Organization of the United Nations, FAO Fisheries Synopsis, no. 125, v. 10, 

442 p.

23. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

24. Hop, H., Welch, H.E., and Crawford, R.E., 1997, Population structure and feeding ecology of Arctic cod schools in the 

Canadian High Arctic: American Fisheries Society Symposium 19, p. 13–39.

25. Lowry, L.F., and Frost, K.J., 1981, Distribution, growth, and foods of Arctic cod (Boreogadus saida) in the Bering, 

Chukchi, and Beaufort Seas: The Canadian Field-Naturalist, v. 92, p. 186–191.

26. Bradstreet, M.S.W., 1980, Thick-billed murres and black guillemots in the Barrow Strait area, N.W.T., during spring—Diets 

and food availability along ice edges: Journal of Zoology, v. 58, no. 11, p. 2,120–2,140. 

27. Gradinger, R.R., and Bluhm, B.A., 2004, In-situ observations on the distribution and behavior of amphipods and Arctic cod 

(Boreogadus saida) under the sea ice of the High Arctic Canada Basin: Polar Biology, v. 27, no. 10, p. 595–603.

28. Dunn, J.R., and Matarese, A.C., 1987, A review of the early life history of northeast Pacific gadoid fishes: Fisheries 

Research, v. 5, nos. 2–3, p. 163–184.

29. Hop, H., Graham, M., and Treadeau, V.L., 1995, Spawning energetics of Arctic cod (Boreogadus saida) in relation to 

seasonal development of the ovary and plasma sex steroid levels: Canadian Journal of Fisheries and Aquatic Sciences, v. 52, 

no. 3, p. 541–550.

30. Rass, T.S., 1968, Spawning and development of polar cod: Rapports et Procčs-Verbeaux des Réunions du Conseil 

International pour l’Exploration de la Mer, v. 158, p. 135–137.

31. Aronovich, T.M., Doroshev, S.I., Spectorova, L.V., and Makhotin, V.M., 1975, Egg incubation and larval rearing of navaga 

(Eleginus navaga Pall.), Arctic Cod (Boreogadus saida lepechin) and Arctic flounder (Liopsetta glacialis Pall.) in the 

laboratory: Aquaculture, v. 6, no. 3, p. 233–242.

32. Quast, J.C., 1974, Density distribution of juvenile Arctic cod, Boreogadus saida, in the eastern Chukchi Sea in the fall of 

1970: Fishery Bulletin, v. 72, no. 4, p. 1,094–1,105.

33. Wyllie-Echeverria, T., Barber, W.E., and Wyllie-Echeverria, S., 1997, Waters masses and transport of age-0 Arctic cod and 

age-0 Bering flounder into the northeastern Chukchi Sea, in Reynolds, J.B., ed., Symposium 19, Proceedings of the “Fish 

Ecology in Arctic North America Symposium,” held at Fairbanks, Alaska, May 19–21, 1992: American Fisheries Society, 

p. 60–67.

34. Ponton, D., and Fortier, L., 1992, Vertical distribution and foraging of marine fish larvae under the ice cover of southeastern 

Hudson Bay: Marine Ecology Progress Series, v. 81, p. 215–227. 

35. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, nos. 1–2, p. 57–70.

36. Dunton, K.H., Schonberg, S.V., and Cooper, L.W., 2012, Food web structure of the Alaskan nearshore shelf and estuarine 

lagoons of the Beaufort Sea: Estuaries and Coasts, v. 35, no. 2, p. 416–435.
198 Alaska Arctic Marine Fish Ecology Catalog

37. Bradstreet, M.S.W., 1980, Thick-billed murres and black guillemots in the Barrow Strait area, N.W.T., during spring—Diets 

and food availability along ice edges: Journal of Zoology, v. 58, no. 11, p. 2,120–2,140.

38. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

39. Hognestad, P.T., 1968, Observations on polar cod in the Barents Sea: Rapports et procès-verbaux des reunions, Conseil 

permanent international pour l’exploration de la mer., v. 158, p. 126–130.

40. Falk-Petersen, I.-B., Frivoll, V., Gulliksen, B., and Haug, T., 1986, Occurrence and size/age relations of polar cod, 

Boreogadus saida (Lepechin), in Spitsbergen coastal waters: Sarsia, v. 71, no. 3–4, p. 235–245.

41. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

42. Logerwell, E., Rand, K., and Weingartner, T.J., 2011, Oceanographic characteristics of the habitat of benthic fish and 

invertebrates in the Beaufort Sea: Polar Biology, v. 34, p. 1,783–1,796.

43. Cannon, T.C., Glass, D.R., and Prewitt, C.M., 1991, Habitat use patterns of juvenile Arctic cod in the coastal Beaufort Sea 

near Prudhoe Bay, Alaska, in Benner, C.S., and Middleton, R.W., eds., Symposium 11, Proceedings from the 119th Annual 

Meeting of the American Fisheries Society, Anchorage, Alaska, 1989: American Fisheries Society, p. 157–162.

44. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

45. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010. 

46. Graham, M., and Hop, H., 1995, Aspects of reproduction and larval biology of Arctic cod (Boreogadus saida): Arctic, v. 48, 

no. 2, p. 130–135.

47. Bradstreet, M.S.W., Sekerak, A.D., Finley, K.J., Griffiths, W.B., Evans, C.R., Fabijan, M.F., and Stallard, H.E., 1986, 

Aspects of the biology of Arctic cod (Boreogadus saida) and its importance in Arctic marine food chains: Canada 

Department of Fisheries and Oceans, Canadian Technical Report of Fisheries and Aquatic Sciences, iss. 1491, 193 p.

48. Thanassekos, S., Virginia Institute of Marine Science, written commun., 2010. 

49. Ponomarenko, V.P., 1968, Some data on the distribution and migrations of polar cod in the seas of the Soviet Arctic: 

Rapports et Procès-Verbeaux des Réunions du Conseil International pour l’Exploration de la Mer, v. 158, p. 131–135.

50. Fortier, L., Sirois, P., Michaud, J., and Barber, D., 2006, Survival of Arctic cod larvae (Boreogadus saida) in relation to sea 

ice and temperature in the Northeast Water Polynya (Greenland Sea): Canadian Journal of Fisheries and Aquatic Sciences, 

v. 63, no. 7, p. 1,608–1,616.

51. Doyle, M.J., University of Washington, written commun., 2010. 

52. Benoit, D., Simard, Y., and Fortier, L., 2008, Hydroacoustic detection of large winter aggregations or Arctic cod 

(Boreogadus saida) at depth in ice-covered Franklin Bay (Beaufort Sea): Journal of Geophysical Research, v. 113, 

no. CO6S90, 9 p.

53. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

54. Wiswar, D.W., and Frugé, D.J., 2006, Fisheries investigations in western Camden Bay, Arctic National Wildlife Refuge, 

Alaska, 1987: Alaska Fisheries Data Series, U.S. Fish and Wildlife Service, 2006-10, 49 p.

55. Lowry, L.F., Frost, K.J., and Burns, J.J., 1980, Variability in the diet of ringed seals, Phoca hispida, in Alaska: Canadian 

Journal of Fisheries and Aquatic Sciences, v. 37, no. 12, p. 2,254–2,261.

56. Schmidt, D.R., Griffiths, W.B., and Martin, L.R., 1987, Importance of anadromous fish overwintering habitat in the 

Sagavanirktok River Delta, Alaska: Anchorage, Alaska, Report by Ecological Research Associates for Standard Alaska 

Production Company and North Slope Borough, 71 p.
Arctic Cod 199

57. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

58. Welch, H.E., Crawford, R.E., and Hop, H., 1993, Occurrence of Arctic cod (Boreogadus saida) schools and their 

vulnerability to predation in the Canadian High Arctic: Arctic, v. 46, no. 4, p. 331–339. 

59. Griffiths, W.B., 1983, Fish, in Truett, J.C., ed, Final reports of principal investigators—Volume 24—Environmental 

characterization and biological use of lagoons in the eastern Beaufort Sea: Anchorage, Alaska, National Oceanic and 

Atmospheric Administration, Office of Oceanography and Marine Assessment, Ocean Assessments Division, p. 325–366.

60. Moulton, L.L., and Tarbox, K.E., 1987, Analysis of Arctic cod movements in the Beaufort Sea nearshore region, 1978–79: 

Arctic, v. 40, no. 1, p. 43–49.

61. Geoffroy, M., Robert, D., Darnis, G., and Fortier, L., 2011, The aggregation of polar cod (Boreogadus saida) in the deep 

Atlantic layer of ice-covered Amundsen Gulf (Beaufort Sea) in winter: Polar Biology, v. 34, no. 12, p. 1,959–1,971.

62. Thorsteinson, L.K., Jarvela, L.E., and Hale, D.A., 1990, Arctic fish habitat use investigations—Nearshore studies in the 

Alaskan Beaufort Sea, summer 1988: Final Report, Alaska Office, Ocean Assessments Division, National Oceanic and 

Atmospheric Administration, Research Unit 682, 125 p.

63. Bouchard, C., Madsen, M.L., Fevolden, S.-E., and L. Fortier, 2008, Population structure in polar cod (Boreogadus saida)—

First results from a circumpolar study using microsatellites: Quebec City, Quebec, Arctic Change 2008, Conference 

Programme and Abstracts, p. 187–188.

64. Sekerak, A.D., Stallard, N., and Griffiths, W.B., 1992, Distribution of fish and fish harvests in the nearshore Beaufort Sea 

and Mackenzie Delta during ice-covered periods, October–June: Environmental Studies Research Funds Report, LGS Ltd. 

No. 117, 524 p.

65. Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbour, N.W.T.: Canadian Technical 

Report of Fisheries and Aquatic Sciences, no. 1218, 21 p.

66. Frost, K.J., and Lowry, L.F., 1981, Foods and trophic relationships of cetaceans in the Bering Sea, in Hood, D.W., and 

Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and Resources: National Oceanic and Atmospheric 

Administration, p. 825–836.

67. Frost, K.J., and Lowry, L.F., 1984, Trophic relationships of vertebrate consumers in the Alaskan Beaufort Sea, in Barnes, 

P.W., Schell, D.M., and Reimnitz, E., eds., The Alaska Beaufort Sea—Ecosystems and environments: San Diego, Academic 

Press, p. 382–401.

68. Coyle, K.O., Gillispie, J.A., Smith, R.L., and Barber, W.E. 1997, Food habits of four demersal Chukchi Sea fishes, in

Reynolds, J.B., ed., Symposium 19, Proceedings of the Fish Ecology in Arctic North America Symposium: American 

Fisheries Society, Fairbanks, Alaska, May 19–21, 1992, p. 310–318.

69. Cui, X., Grebmeier, J.M., and Cooper, L.W., 2012, Feeding ecology of dominant groundfish in the northern Bering Sea: 

Polar Biology, v. 35, no. 9, p. 1,407–1,419.

70. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320. 

71. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.

72. Jones, M.L., and Den Beste, J., 1977, Tuft Point and adjacent coastal areas fisheries projects: Calgary, Alberta, Canada, 

Aquatic Environments, Ltd., 152 p.

73. Frost, K.J., and Lowry, L.F., 1981, Trophic importance of some marine gadids in northern Alaska and their body-otolith size 

relationships: Fishery Bulletin, v. 79, no. 1, p. 187–192.

74. Lowry, L.F., Frost, K.J., and Seaman, G.A., 1986, Investigations of belukha whales in coastal waters of western and 

northern Alaska: Outer Continental Shelf Environmental Program Unit 612, Final Report, p. 359–392.
200 Alaska Arctic Marine Fish Ecology Catalog

75. Bond, W.A., and Erickson, R.N., 1989, Summer studies of the nearshore fish community at Phillips Bay, Beaufort Sea coast, 

Yukon: Winnepeg, Manitoba, Canadian Technical Report of Fisheries and Aquatic Sciences, Central and Arctic Region, 

Department of Fisheries and Oceans, no. 1676, 102 p.

76. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

77. Murdoch, J., 1884, Fish and fishing at Point Barrow, Arctic Alaska: Transactions of the American Fisheries Society, v. 13, 

no. 1, p. 111–115.

78. Saario, D.J., and Kessel, B., 1966, Human ecological investigations at Kivalina, in Wilimovsky, N.J., and Wolfe, J.N., eds., 

Environment of the Cape Thompson region, Alaska: Oak Ridge, Tennessee, U.S. Atomic Energy Commission, Division of 

Technical Information, p. 969–1,039.

79. Nelson, R.K., 1969, Hunters of the Northern Ice: Chicago, University of Chicago Press, 429 p.

80. Bendock, T.N., 1977, Beaufort Sea estuarine fishery study: Alaska Department of Fish and Game Annual Report, Contract 

#03-5-022-69, p. 670–729.

81. Wein, E.E., Freeman, M.M.R., and Makus, J.C., 1996, Use of and preference for traditional foods among the Belcher Island 

Inuit: Arctic, v. 49, no. 3, p. 256–264.

82. Food and Agriculture Organization of the United Nations, 2011, Boreogadus saida (Lepechin, 1774): Fisheries and 

Aquaculture Department, Food and Agriculture Organization of the United Nations, Species Fact Sheet, Web site, accessed 

May 18, 2014, at http://www.fao.org/fishery/species/2233/en.

83. Bouchard, C., and Fortier, L., 2011, Circum-Arctic comparison of the hatching season of polar cod Boreogadus saida—A 

test of the freshwater winter refuge hypothesis: Progress in Oceanography, v. 90, nos. 1–4, p. 105–116.

84. Crawford, R.E., and Jorgenson, J.K., 1993, Schooling behaviour of Arctic cod, Boreogadus saida, in relation to drifting 

pack ice: Environmental Biology of Fishes, v. 36, no. 4, p. 345–357. 

85. Gaston, A.J., Woo, K., and Hipfner, J.M., 2003, Trends in forage fish populations in northern Hudson Bay since 1981, as 

determined from the diet of nestling thick-billed murres Uria lomvia: Arctic, v. 56, no. 3, p. 227–233.

86. Divoky, G., 2008, Annual and seasonal variation in nearshore fish availability associated with the record Arctic pack ice 

minimum of 2007: Quebec City, Quebec, Arctic Change 2008, Conference Programme and Abstracts, p. 69–70.

87. Kunz, K.L., Frickenhaus, S., Hardenberg, S., Johansen, T., Leo, E., Pörtner, H.O., Schmidt, M., Windisch, H.S., Knust, 

R., and Mark, F.C., 2016, New encounters in Arctic waters—A comparison of metabolism and performance of polar cod 

(Boreogadus saida) and Atlantic cod (Gadus morhua) under ocean acidification and warming: Polar Biology, no. 39, 

p. 1,137–1,153.

88. Kent, D., Drost, H.E., Fisher, J., Oyama, T., and Farrell, A.P., 2015, Laboratory rearing of wild Arctic cod Boreogadus saida

from egg to adulthood: Journal of Fish Biology, doi:10.1111/jfb.12893.

89. Drost, H.E., Carmack, E.C., and Farrell, A.P., 2014, Upper thermal limits of cardiac function for Arctic cod Boreogadus 

saida, a key food web fish species in the Arctic Ocean: Journal of Fish Biology, doi:10.1111/jfb.12397.

90. Drost, H.E., Fisher, J., Randall, F., Kent, D., Carmack, E.C., and Farrell, A.P., 2015, Upper thermal limits of the hearts of 

Arctic cod Boreogadus saida: Journal of Fish Biology, doi:10.1111/jfb.12807.

91. Drost, H.E., Lo, M., Carmack, E., and Farrell, A.P., 2016, Acclimation potential of Arctic Cod (Boreogadus saida Lepechin) 

in the rapidly warming Arctic Ocean—Advanced publication: Journal of Experimental Biology, doi:10.1242/jeb.140194.

92. Laurel, B.J., Spencer, M., Iseri, P., and Copeman, L.A., 2015, Temperature-dependent growth and behavior of juvenile 

Arctic cod (Boreogadus saida) and co-occurring North Pacific gadids: Polar Biology, no. 39, p. 1,127–1,135.

93. Cheung, W.W.L., Lam, V.W.Y., and Pauly, D., 2008, Dynamic bioclimate envelope model to predict climate-induced 

changes in distribution of marine fishes and invertebrates, in Cheung, W.W.L., Lam, V.W.Y., and Pauly, D., eds., Modeling 

present and climate-shifted distribution of marine fishes and invertebrates: Vancouver, University of British Columbia, 

Fisheries Centre Research Reports, v. 16, no. 3, p. 5–50.
Saffron Cod 201

Saffron Cod (Eleginus gracilis) 

(Tilesius, 1810) 

Family Gadidae 

Colloquial Name: Iñupiat: Uugak [1]. This species and the Arctic 

Cod are called Tomcod (not to be confused with Microgadus 

proximus). 

Ecological Role: This is a species of major ecological importance, 

particularly in the Chukchi Sea. Saffron Cod are believed to be a 

major competitor of Arctic Cod and changes in sea ice associated 

with warming may give the species a competitive advantage.

Physical Description/Attributes: Mottled brown to gray-green body washed with yellow. Ventral areas are white to yellow, 

pectoral fins are yellow, and margins of dorsal and anal fins are white. For specific diagnostic characteristics, see Fishes of 

Alaska (Mecklenburg and others, 2002, p. 293) [2]. Swim bladder: Present [2]. Antifreeze glycoproteins in blood serum: 

Present [3].

Range: U.S. Chukchi and Beaufort Seas [4]. Elsewhere in Alaska, from Bering Sea and Gulf of Alaska to Sitka, southeastern 

Alaska. Worldwide, from Sea of Japan and Sea of Okhotsk to East Siberian Sea and eastward in Arctic to Melville Sound, 

Bathurst Inlet, Nunavut [4]. 

Relative Abundance: Patchily abundant in U.S. Chukchi and Beaufort Seas [7–10] eastward to Cambridge Bay, Nunavut [6]. 

Most abundant species during summer in northern Bering and southern U.S. Chukchi Seas [12]. Abundant from Sea of Japan 

and Sea of Okhotsk to eastern Bering Sea [13–15] and central Gulf of Alaska [16]. Appears to be increasing in abundance in 

Prince William Sound [17].

Saffron Cod (Eleginus gracilis), 233 mm, Chukchi Sea, 2007. 

Photograph by C.W. Mecklenburg, Point Stephens Research.

Geographic distribution of Saffron Cod (Eleginus gracilis) within Arctic Outer Continental Shelf Planning Areas 

[5] based on review of published literature and specimens from historical and recent collections [2, 4, 6]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Saffron Cod

Eleginus gracilis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
202 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Shallow, nearshore to 200 m, typically less than 50 m [18–20]. Four pelagic larvae taken in U.S. Chukchi Sea 

between 18 and 36 m and in Bering Sea from 0–162 m. Largest catches were in less than 60 m [21]. One pelagic juvenile was 

taken in U.S. Chukchi Sea in midwaters between 45 m and surface [22]. Older juveniles recruit to very shallow near-shore 

waters [23–26]. However, off Hokkaido, Japan, and the Kuril Islands, Russia, juveniles are abundant to depths of at least 200 m 

[27]. Spawning occurs in shallow waters [23, 28, 29] to at least 32 m in the western Pacific Ocean [27].

Habitats and Life History

Eggs—Size: 0.8–1.7 mm [27, 30]. Time to hatching: 28–49 days [30]. Habitat: Demersal, non-adhesive [27, 

30–32]. 

Larvae—Size at hatching: 3.5–3.9 mm SL [31]. Size at juvenile transformation: 24–27 mm SL [21]. Days to 

juvenile transformation: Unknown. Habitat: Pelagic [17, 21].

Juveniles—Age and size: 21–35 cm FL [21, 31]. Habitat: Early juveniles are pelagic, becoming more benthic as 

fish mature [21]. In Gulf of Alaska, closely associated with nearshore eelgrass beds [16, 17].

Adults—Age and size at first maturity: Very little research has been conducted. Generally, maximum age, 

growth rates, and age at first maturity vary with location. Off Hokkaido, Japan, few mature as early as one 

year and all are mature by 2 years [33]. In Siberian Chukchi Sea, fish mature at 4–5 years. Overall, fish mature 

at 21.0–35.0 cm FL [31]. Maximum age: 19 years in Canadian Beaufort Sea, [34]. Maximum ages are highly 

variable among geographic locations [7, 27, 35]. In Siberian Chukchi Sea, fish live to 15 years. Maximum life 

spans steeply decline to the south. Fish living in Peter the Great Bay, in Sea of Japan, only reach about 8 years of 

age [27]. Maximum size: 55 cm TL [2] and possibly to 63 cm TL [6]. Females are slightly heavier at length than 

males. Habitat: Benthic and midwater [2, 36–39]. Shallow, nearshore and, at least around Kodiak Island and Sea 

of Japan, often associated with eelgrass [16, 40].

Substrate—Soft and hard sea floors [41]. Sandy-stone or gravel bottoms for spawning [32]. 

Physical/chemical—Temperature: Unknown. Elsewhere, between -1.7 and 11.7 °C in southeastern Bering 

Sea [42]. In Amundsen Gulf, a large mortality event occurred when fish encountered 18.0 °C waters flowing 

out of Coppermine River [11]. Spawn between -1.8 and 1.8 °C [27, 32, 34, 43]. Eggs remain viable at water 

temperatures of somewhat greater than -3.8–8.0 °C [43]. Salinity: Primarily marine and brackish waters [36–39], 

although described as entering both rivers and lakes [43], and not ascending upstream of river mouths [32]. In 

Russia, spawning occurred only at 27 parts per thousand or more [32]. Temperatures higher than 1.2 °C and 

salinities less than 21 parts per thousand are reportedly unfavorable for egg and larval survival [27, 32, 44].

Benthic and reproductive distribution of Saffron Cod (Eleginus gracilis).

Some fish 

to 200 m

Open ocean Shore

Depth, in meters

250

200

150

100

50

0

250

200

150

100

50

0

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Eleginus gracilis

Saffron Cod

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Larvae and 

younger juveniles

Overall depth data from outside U.S. Beaufort-Chukchi Seas Spawning data from outside U.S. Beaufort-Chukchi Seas

Depths of larvae in U.S. 

Beaufort-Chukchi Seas Adults and older

 juveniles 

Depths documented in U.S. 

Beaufort-Chukchi Seas
Saffron Cod 203

Behavior

Diel—Unknown. 

Seasonal—Juveniles recruit to very shallow near-shore waters in summer [23–26]. Movements of juvenile and 

adults are not well known. Generally, Saffron Cod have relatively circumscribed movements, with a limited 

winter inshore and summer offshore migration [29]. However, large numbers move into shallow waters of the 

Yukon Territory, Canada and southeastern Beaufort Sea in early summer [28] and fish in northern Bering Sea 

may move northwards into U.S. Chukchi Sea in summer [45]. Alongshore movements may be quite limited. One 

fish tagged in the Arctic National Wildlife Refuge only moved 30 km in 3 years [46]. Locations of overwintering 

grounds are not well known. Some fish overwinter in nearshore estuarine and marine waters and these 

aggregations may be limited to specific geographic areas [34, 47–49]. 

Reproductive—Spawning behavior and locations are poorly understood. Throughout their geographic range, 

some spawning occurs in shallow waters [23, 28, 29]. In Russia, fish spawned in areas with strong tidal currents 

and sandy-stone or gravel bottoms [32]. 

Schooling—Schools, sometimes in high densities [40]. 

Feeding—Feed throughout the year at least in U.S. Chukchi Sea and northern Bering Sea [23]. Juveniles in 

White Sea feed during day and night. Fish fed among rockweed patches during day and over sand at night. They 

also fed in the water column on the flood tide [50].

Populations or Stocks

Initial investigations on genetic diversity and stock structure are underway at the University of Alaska Fairbanks.

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external. 

Spawning season—Winter in U.S. Chukchi Sea [23]. Mainly, December–February throughout Alaska [31]. 

Elsewhere, from December to at least May [29, 37] and off Kamchatka Peninsula, Russia, perhaps as late as 

June [43]. 

Fecundity—4,900–690,000 eggs, varies with location [27].

Food and Feeding

Food items—Fishes (for example, Arctic Cod, Capelin, Fourhorn Sculpin, and Saffron Cod) and crustaceans 

(for example, amphipods, isopods, mysids, and shrimps) often are very important, and priapulids, polychaetes, 

clams, insects, pteropods, and plant material also are consumed [11, 23, 35, 36, 51–53]. Juveniles prey on 

zooplankton [35]. Larger fish prey on a wide range of benthic and epibenthic organisms. During spawning 

season, adults reportedly feed heavily on Saffron Cod eggs [19]. 

Trophic level—4.1 [54]. 

Biological Interactions

Predators: Very important prey for ringed seals from at least Nome, Alaska (during at least mid-summer to 

December) to the U.S. Chukchi Sea (throughout the year) [55, 56]. Important summer food for belugas to at 

least as far north as Wainwright [57, 58]. Other predators include Arctic Lamprey and Fourhorn Sculpin [20, 

23]. Additional predators that have been reported include Great, Plain, and Thorny Sculpins, Pacific Cod, Pacific 

Halibut, Arctic Smelt, Saffron Cod, Black-legged Kittiwake, Common and Thick-billed Murres, bearded and 

ribbon seals, Steller sea lion, harbor porpoise, Beluga, Fin, Humpback, Ninke, and Sperm whales [59–67].

Competitors: Likely co-occurring gadids including Arctic Cod, Ice Cod, Pacific Cod (including ogac), and 

Walleye Pollock [7, 21, 68]. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (t

m=2–3; Fecundity=4,900) [69]. 
204 Alaska Arctic Marine Fish Ecology Catalog

Traditional and Cultural Importance

Commonly taken in subsistence fisheries in both the U.S. Chukchi and Beaufort Seas and in the Bering Sea, 

usually through the ice by both hook and line and gill nets [1, 24, 37, 70]. Historically, this was an extremely 

important species to the Inuits residing along the Bering Sea of Alaska where fish were commonly taken during 

spring as soon as the ice melted from the nearshore, but were particularly important in November, when the pack 

ice returned. Large numbers of Saffron Cod were utilized by the inhabitants of Norton Sound. They are used as 

food for both man and dog [71].

Commercial Fisheries

Currently, Saffron Cod are not commercially harvested.

Potential Effects of Climate Change

Uncertain. Reproducing in Arctic as well as Boreal waters [4], this is a somewhat eurythermic species, apparently 

able to function within a relatively wide temperature range. Assuming that such coldwater-adapted competitors 

as Arctic Cod are negatively effected, the reduced competition could be beneficial to the Saffron Cod population. 

This is supported by new experimental studies that indicate juvenile Saffron Cod growth rate responded 

positively to increasing temperatures ranging from 0 to 16 °C and above [72].

Areas for Future Research [A]

Little is known about the ecology of this species. It is an important forage fish, subsistence resource, and 

competitor of Arctic Cod. Information about seasonal habitats and life history and stocks structure of the 

populations is needed. Initial laboratory and modeling studies suggest the competitive capacity of Saffron Cod 

with respect to Arctic Cod and other gadids. Additional studies are needed to evaluate the effects of temperature 

and other population limiting factors, including competition, on this species.

References Cited

Dunn, J.R., and Matarese, A.C., 1987, A review of the early life history of northeast Pacific gadoid fishes: Fisheries Research, 

v. 5, nos. 2–3, p. 163–184. [31]

Dunn, J.R., and Vinter, B.M., 1984, Development of larvae of the saffron cod, Eleginus gracilis, with comments on the 

identification of gadid larvae in Pacific and Arctic waters contiguous to Canada and Alaska: Canadian Journal of Fisheries and 

Aquatic Sciences, v. 41, no. 2, p. 304–318. [21]

Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p. [23]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Safronov, S.N., 1986, Peculiarities of reproduction and principles of change in the fecundity of Pacific navaga, Eleginus gracilis

(Gadidae): Journal of Ichthyology, v. 26, no. 5, p. 59–68. [27] 

Bibliography

1. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. 
Saffron Cod 205

3. Power, G., 1997, A review of fish ecology in Arctic North America: American Fisheries Society Symposium, no. 19, 

p. 13–39.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arctic-marine-fishes.

7. Wolotira, R.J., Jr., Sample, T.M., and Morin, M., Jr., 1977, Demersal fish and shellfish resources of Norton Sound, the 

southeastern Chukchi Sea, and adjacent waters in the baseline year 1976: Seattle, Washington, Northwest and Alaska 

Fisheries Center, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Processed Report, 

69 p.

8. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

9. Bond, W.A., and Erickson, R.N., 1993, Fisheries investigations in coastal waters of Liverpool Bay, Northwest Territories: 

Winnipeg, Manitoba, Canada Department of Fisheries and Oceans, Central and Arctic Region, Canadian Manuscript Report 

of Fisheries and Aquatic Sciences No. 2204, 59 p.

10. Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., and Colonell, J.M., 1996, The 1995 fish and oceanography 

study in Mikkelsen Bay, Alaska: Anchorage, Alaska, Prepared by LGL Alaska Research Associates and Woodward-Clyde 

Consultant for BP Exploration (Alaska) Inc., 102 p. plus apps.

11. Ellis, D.V., 1962, Observations on the distribution and ecology of some Arctic fish: Arctic, v. 15, no. 3, p. 179–189.

12. Lowry, L.F., and Frost, K.J., 1981, Distribution, growth, and foods of Arctic cod (Boreogadus saida) in the Bering, 

Chukchi, and Beaufort Seas: The Canadian Field-Naturalist, v. 92, p. 186–191.

13. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

14. Kolpakov, N.V., 2005, Diversity and seasonal dynamics of ichthyocenosis of the Circumlittoral of Russkaya Bight 

(Northern Primor’s): Journal of Ichthyology, v. 45, no. 9, p. 744–753.

15. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, p. 

226–237.

16. Laurel, B.J., Stoner, A.W., Ryer, C.H., Hurst, T.P., and Abookire, A.A., 2007, Comparative habitat associations in juvenile 

Pacific cod and other gadids using seines, baited cameras and laboratory techniques: Journal of Experimental Marine 

Biology and Ecology, v. 351, nos. 1–2, p. 42–55.

17. Johnson, S.W., Thedinga, J.F., and Neff, A.D., 2009, Invasion by saffron cod Eleginus gracilis into nearshore habitats of 

Prince William Sound, Alaska, USA: Marine Ecology Progress Series, v. 389, p. 203–212.

18. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

19. Chuchukalo, V.I., Radchenko, V.I., Nadtochii, V.A., Koblikov, V.N. and Slabinskii, A.M., 1999, Feeding and some features 

of ecology of Gadidae of the western Kamchatka shelf in summer 1996: Journal of Ichthyology, v. 39, no. 4, p. 309–321.
206 Alaska Arctic Marine Fish Ecology Catalog

20. Majewski, A.R., Sareault, J.E., and Reist, J., 2006, Fish catch data from offshore sites in the Mackenzie River estuary and 

Beaufort Sea during the open water season, August 2004 aboard the CCGS Nahidik: Winnipeg, Manitoba, Fisheries and 

Oceans Canada, no. 2771, 42 p.

21. Dunn, J.R., and Vinter, B.M., 1984, Development of larvae of the saffron cod, Eleginus gracilis, with comments on the 

identification of gadid larvae in Pacific and Arctic waters contiguous to Canada and Alaska: Canadian Journal of Fisheries 

and Aquatic Sciences, v. 41, no. 2, p. 304–318.

22. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, nos. 1–2, p. 57–70.

23. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

24. Barton, L.H., 1978, Finfish resource surveys in Norton Sound and Kotzebue Sound: Alaska Department of Fish and Game, 

Commercial Fisheries Division, p. 75–313.

25. Wiswar, D.W., and Frugé, D.J., 2006, Fisheries investigations in western Camden Bay, Arctic National Wildlife Refuge, 

Alaska, 1987: Alaska Fisheries Data Series, U.S. Fish and Wildlife Service, 2006-10, 49 p.

26. Laurel, B.J., Ryer, C.H., Knoth, B., and Stoner, A.W., 2008, Temporal and ontogenetic shifts in habitat use by juvenile 

Pacific cod (Gadus macrocephalus): Journal of Experimental Marine Biology and Ecology, v. 377, no. 1, p. 28–35.

27. Safronov, S.N., 1986, Peculiarities of reproduction and principles of change in the fecundity of Pacific navaga, Eleginus 

gracilis (Gadidae): Journal of Ichthyology, v. 26, no. 5, p. 59–68.

28. Bond, W.A., and Erickson, R.N., 1989, Summer studies of the nearshore fish community at Phillips Bay, Beaufort Sea 

coast, Yukon: Winnepeg, Manitoba, Canadian Technical Report of Fisheries and Aquatic Sciences, Central and Arctic 

Region, Department of Fisheries and Oceans, no. 1676, 102 p.

29. Andriashev, A.P., 1964, Fishes of the northern seas of the U.S.S.R.: Jerusalem, Israel Program for Scientific Translations, 

617 p.

30. Aronovich, T.M., Doroshev, S.I., Spectorova, L.V., and Makhotin, V.M., 1975, Egg incubation and larval rearing of navaga 

(Eleginus navaga Pall.), Arctic Cod (Boreogadus saida lepechin) and Arctic flounder (Liopsetta glacialis Pall.) in the 

laboratory: Aquaculture, v. 6, no. 3, p. 233–242.

31. Dunn, J.R., and Matarese, A.C., 1987, A review of the early life history of northeast Pacific gadoid fishes: Fisheries 

Research, v. 5, nos. 2–3, p. 163–184.

32. Berg, L.S., 1949, Freshwater fishes of the U.S.S.R. and adjacent countries, volume 3 (4th ed.): Moscow, Academy of 

Sciences of the U.S.S.R. Zoological Institute, 250 p. [Translated from Russian by Israel Program for Scientific Translations, 

Jerusalem, IPST Catalog No. 743.]

33. Chen, A.-L., and Sakurai, Y., 1993, Age and growth of saffron cod (Eleginus gracilis): Scientific Reports of Hokkaido 

Fisheries Experimental Station, v. 42, p. 251–264.

34. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.

35. Anukhina, A.M., 1968, Population dynamics of White Sea navaga: Rapports et Procčs-Verbeaux des Réunions du Conseil 

International pour l’Exploration de la Mer, v. 158, p. 138–142.

36. Bond, W.A., and Erickson, R.N., 1987, Fishery data from Phillips Bay, Yukon, 1985: Winnipeg, Manitoba, Canadian Data 

Report of Fisheries and Aquatic Sciences, Central and Artic Region, Department of Fisheries and Oceans, no. 635, 47 p.

37. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

38. Dragoo, D.E., 2006, Seabird, fish, marine mammals and oceanography coordinated investigations (SMMOCI) near Bluff, 

Norton Sound, Alaska, July 2002: U.S. Fish and Wildlife Service Report AMNWR 06/03, 35 p.
Saffron Cod 207

39. Walters, V., 1955, Fishes of western Arctic America and eastern Arctic Siberia: Bulletin of the American Museum of Natural 

History, v. 106, p. 259–368.

40. Gomelyuk, V.E., and Shchetkov, S.Y., 1992, Distribution of fish in coastal biotopes of Peter the Great Bay, Sea of Japan, in 

the summer period: Biologiya Morya, nos. 3–4, p. 26–32.

41. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

42. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

43. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific, Part V: Soviet Fisheries Investigation in the Northeastern Pacific, p. 161–224. 

[Russian translation by Israel Program for Scientific Translations, Jerusalem (1972).]

44. Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbour, N.W.T.: Canadian Technical 

Report of Fisheries and Aquatic Sciences, no. 1218, 21 p.

45. Lowry, L.F., Frost, K.J., and Burns, J.J., 1980, Variability in the diet of ringed seals, Phoca hispida, in Alaska: Canadian 

Journal of Fisheries and Aquatic Sciences, v. 37, no. 12, p. 2,254–2,261.

46. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

47. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

48. Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.

49. Sekerak, A.D., Stallard, N., and Griffiths, W.B., 1992, Distribution of fish and fish harvests in the nearshore Beaufort Sea 

and Mackenzie Delta during ice-covered periods, October–June: Environmental Studies Research Funds Report, LGS Ltd. 

No. 117, 524 p.

50. Girsa, I.I., 1986, Effect of ecological conditions on changes in the behavior of yearling navaga, Eleginus navaga (Gadidae) 

during summer in the coastal waters of the White Sea: Journal of Ichthyology, v. 26, no. 2, p. 140–143.

51. Jones, M.L., and Den Beste, J., 1977, Tuft Point and adjacent coastal areas fisheries projects: Calgary, Alberta, Canada, 

Aquatic Environments, Ltd., 152 p.

52. Lacho, G., 1991, Stomach content analyses of fishes from Tuktoyaktuk Harbour, N.W.T., 1981: Winnipeg, Manitoba, 

Canadian Data Report of Fisheries and Aquatic Sciences, Central and Arctic Region, Department of Fisheries and Oceans, 

no. 853, 15 p.

53. Coyle, K.O., Gillispie, J.A., Smith, R.L., and Barber, W.E. 1997, Food habits of four demersal Chukchi Sea fishes, in

Reynolds, J.B., ed., Symposium 19, Proceedings of the Fish Ecology in Arctic North America Symposium: American 

Fisheries Society, Fairbanks, Alaska, May 19–21, 1992, p. 310–318.

54. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.

55. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.

56. Frost, K.J., and Lowry, L.F., 1981, Foods and trophic relationships of cetaceans in the Bering Sea, in Hood, D.W., and 

Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and Resources: National Oceanic and Atmospheric 

Administration, p. 825–836.

57. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.
208 Alaska Arctic Marine Fish Ecology Catalog

58. Lowry, L.F., Frost, K.J., and Seaman, G.A., 1986, Investigations of belukha whales in coastal waters of western and 

northern Alaska: Outer Continental Shelf Environmental Program Unit 612, Final Report, p. 359–392.

59. Kawakami, T., 1980, A review of sperm whale food: Scientific reports of the Whales Research Institute, no. 32, p. 199–218.

60. Kawamura, A., 1980, A review of food of balaenopterid whales: Scientific Reports of the Whales Research Institute, v. 32, 

p. 115–197.

61. Pitcher, K.W., 1981, Prey of the Steller sea lion, Eumetopias jubatus, in the Gulf of Alaska: Fishery Bulletin, v. 79, no. 3, 

p. 467–472.

62. Kajimura, H., and Loughlin, T.R., 1988, Marine mammals in the oceanic food web of the eastern subarctic Pacific: 

University of Tokyo, Bulletin of the Ocean Research Institute, v. 26, no. 2, p. 187–233.

63. Lowry, L.F., and Frost, K.J., 1981, Feeding and trophic relationships of phocid seals and walruses in the eastern Bering Sea, 

in Hood, D.W., and Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and resources: National Oceanic and 

Atmospheric Administration, p. 813–824.

64. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.

65. Tokranov, A.M., and Vinnikov, A.B., 1991, Diet of the Pacific cod, Gadus morhua macrocephalus, and its position in the 

food chain in Kamchatkan coastal waters: Journal of Ichthyology, v. 31, no. 5, p. 84–98.

66. Karpenko, V.I., and Vasilets, P.M., 1996, Biology of smelt (Osmeridae) in the Korf-Karagin coastal area of the southwestern 

Bering Sea, in Mathisen, O.A., and Coyle, K.O., eds., Ecology of the Bering Sea—A review of Russian literature: 

Fairbanks, Alaska, University of Alaska, Alaska Sea Grant Program, AK-SG-96-01, p. 217–235.

67. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-158, 249 p.

68. Quast, J.C., 1974, Density distribution of juvenile Arctic cod, Boreogadus saida, in the eastern Chukchi Sea in the fall of 

1970: Fishery Bulletin, v. 72, no. 4, p. 1,094–1,105.

69. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

70. Moulton, L.L., Field, L.J., and Kovalsky, R., 1991, Predictability in the catch of Arctic cisco in the Colville River, Alaska: 

American Fisheries Society Symposium no. 11, p. 145–156.

71. Turner, L.M., 1886, Contributions to the natural history of Alaska—Arctic series of publications, no. 2, Washington: 

Washington D.C., U.S. Government Printing Office, 226 p.

72. Laurel, B.J., Spencer, M., Iseri, P., and Copeman, L.A., 2015, Temperature-dependent growth and behavior of juvenile 

Arctic cod (Boreogadus saida) and co-occurring North Pacific gadids: Polar Biology, no. 39, p. 1,127–1,135.
Walleye Pollock 209

Walleye Pollock (Gadus chalcogrammus) 

Pallas, 1814 

Family Gadidae 

Scientific name: Previously called Theragra chalcogramma

(Pallas, 1814), this species was recently returned to its original 

genus Gadus on the basis of morphological and molecular 

evidence [1].

Colloquial Name: None within U.S. Chukchi and Beaufort Seas. 

Ecological Role: Current information on the distribution and abundance of this species suggests it could be of low to moderate 

ecological importance in some parts of the Chukchi Sea and offshore waters north of Barrow, Alaska. This is a key species in 

ecosystem dynamics of the Gulf of Alaska, Prince William Sound, and Bering Sea [2]. 

Physical Description/Attributes: Olive green to brown with dark mottling and blotches on back, and interrupted dark brassy 

olive stripes on upper sides. Fins are brown, dusky gray, or black. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 295) [3]. Swim bladder: Present; no otophysic connection [3]. Antifreeze glycoproteins in 

blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [4–6]. Elsewhere, through Bering Sea, Aleutian Islands and Gulf of Alaska south to 

Carmel, central California and west to Seas of Okhotsk and Japan [3]. Also found in Barents Sea off Norway, where it used to be 

called Theragra finnmarchica [1].

Walleye Pollock (Gadus chalcogrammus) juvenile, 141 mm, 

Bering Strait, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.
210 Alaska Arctic Marine Fish Ecology Catalog

Relative Abundance: Common in U.S. Chukchi and rare in U.S. Beaufort Sea [1, 4, 5, 8, 9]. Elsewhere, abundant in Sea of 

Japan, northern Kuril Islands, Kamchatka Peninsula, Russia, and throughout Bering Sea southward to southeastern Alaska and 

Puget Sound [10–15]. 

tac14-5222_fig3-5-4m_Gadus_chalcogrammus

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Walleye Pollock

Gadus chalcogrammus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Walleye Pollock (Gadus chalcogrammus) within Arctic Outer Continental Shelf 

Planning Areas [7] based on review of published literature and specimens from historical and recent 

collections [1, 3, 8].
Walleye Pollock 211

Depth Range: Entire water column, from surf zone to 1,200 m, commonly at 400 m or less, though large numbers have been 

taken at 800–1,000 m in Bering Sea [13, 16, 17]. Spawning occurs from 46 to 700 m, most commonly between 100 and 250 m 

on deeper continental shelf and upper continental slope [18–20]. Pelagic eggs are from 0 to 400 m, typically less than 200 m in 

Gulf of Alaska and eastern Bering Sea [21–23]. Pelagic larvae are from 0 to 153 m, typically 60 m or less in Gulf of Alaska and 

eastern Bering Sea [22, 24–26].

Open ocean Shore

Depth, in meters

1,500

1,200

900

600

300

0

1,000

800

600

400

200

0

Some fish to 1,200 m

Larvae

Eggs

Gadus chalcogrammus

Walleye Pollock

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Data from outside 

U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Spawning

Benthic and reproductive distribution Walleye Pollock (Gadus chalcogrammus).

Habitats and Life History

Eggs—Size: 1.2–1.8 mm [27, 28]. Time to hatching: 25.5 days at 2 °C and 14 days at 5 °C [29]. Habitat: Pelagic, 

in deep water, rising to shallower water as they develop [3, 22, 30]. 

Larvae—Size at hatching: 3.0–4.0 mm SL [28]. Size at juvenile transformation: About 2.5–4.0 cm SL [20, 

29]. Days to juvenile transformation: 60 days [29]. Habitat: Epipelagic, over continental shelf and slope [3, 27, 

28, 31]. 

Juveniles—Age and size: 2 months to 2–7 years [29] and 2.5 cm SL to 20–48 cm FL [32, 33]. Habitat: Semibenthic, in nearshore waters [3, 34, 35] and then migrate somewhat deeper as they mature [34–36]. Taken among 

eelgrass and kelp [37]. 

Adults—Age and size at first maturity: In eastern Bering Sea, a few males matured at 20 cm FL (2 years), 50 

percent were mature at 31 cm (3 years), and 100 percent were mature at 48 cm (about 7 years) [32]. For females, 

size and age at maturity varied somewhat with location and year. On average, a few matured at 25 cm FL, 

50 percent at 37.4 cm (4 years), and virtually all by 56 cm [33]. In the Gulf of Alaska, 50 percent of females 

matured at 42 cm FL and 5 years [38]. Older females are perhaps somewhat larger at age than are males. 

Maximum age: 33 years [20]. Maximum size: 91 cm TL [3]. Habitat: Semi-demersal to pelagic [3]; commonly 

associated with outer shelf and slope but also uses a wide variety of habitats including nearshore eelgrass and 

kelp beds, large estuaries (such as the Puget Sound), coastal embayments, and open ocean basins (such as the 

Aleutian Basin of Bering Sea) [30, 37]. 

Substrate—Sand, gravel, mud, silt, and bedrock [20, 37]. 

Physical/chemical—Temperature: -1.8–12 °C; rare in waters less than 0 °C [6, 30, 39, 40]. Salinity: Marine 

[20]. Found at 31.3–33.5 ppt in U.S. Chukchi Sea [6].
212 Alaska Arctic Marine Fish Ecology Catalog

Behavior

Diel—They make limited day-night vertical migrations, moving into shallow waters at dusk and night, then 

deeper during day [25, 41, 42]. Juveniles aggregate near sea floor during day then disperse and move shallower 

at night [30, 35, 43]. Juveniles often associated with tentacles of medusae during day [43, 44]. Some adults 

migrate into near-surface waters at night [42]. 

Seasonal—Young-of-the-year recruit to nearshore waters from early summer through autumn [34, 35]. Make 

seasonal inshore-offshore migrations, overwintering in deep part of their depth range [45]. Strong year classes 

have been linked to warm water years when juveniles are transported offshore and away from cannibalistic adults 

[46]. 

Reproductive—Spawning occurs in a number of discrete locations in Strait of Georgia, Gulf of Alaska, Bering 

Sea, and in western Pacific Ocean off Asia [30]. Fish may return to their natal sites to spawn. Females are batch 

spawners [27, 47], spawning at least 14 times in a season [20]. Spawning of an individual female probably takes 

less than 1 month [18]. 

Schooling—Forms large schools [34–36]. 

Feeding—Juveniles and adults are mainly nocturnal feeders [48] whereas most feeding of larvae occurs during 

the day [25].

Populations or Stocks

There is evidence for semi-discrete populations in the Gulf of Alaska and Bering Sea, although the degree of 

genetic isolation of these stocks is unclear. As many as 12 stocks in waters between Japan and southeastern 

Alaska have been postulated [30, 38, 49, 50].

Reproduction

Mode—Oviparous [20]. 

Spawning season—Over all their range, some spawning may occur throughout the year [51]. However, most 

spawning takes place in winter and spring, varying somewhat with location. For instance, in Gulf of Alaska, fish 

spawn around Shumagin Island, Alaska from about 15 February to 1 March, 15 March to 1 April in Shelikov 

Strait [38] and mostly April to Mid-May in southeastern Bering Sea [30]. 

Fecundity—58,000–1,400,000 non-adhesive eggs per season, in batches [27, 47, 52]. 

Food and Feeding

Food items—Larvae: Copepod naupli, larval copepods and small euphausiids [29]. Juveniles: Mainly 

euphausiids [48] as well as copepods and other planktonic crustaceans [29]. Adults: A wide array of midwater 

and benthic organisms. Smaller pollock feed primarily on zooplankton (for example, euphausiids, copepods, and 

gammarid amphipods). Among larger fish, copepods and euphausiids are often very important, as are a number 

of fish species (for example, capelin, eulachon, and lanternfishes) and shrimps. Other frequently eaten organisms 

include mysids, crabs, polychaetes, and cephalopods and crustacean larvae [53–56]. 

Trophic level—3.7 [57] 

Biological Interactions

Predators—Walleye Pollock are extremely important prey for many fishes, seabirds, and mammals. A literature 

search discloses that at least 42 species of fishes, 18 species of seabirds, 7 species of pinnipeds, 9 species of 

cetaceans, and river otters prey on pollock. In Gulf of Alaska, pollock are very important prey to Arrowtooth 

Flounder, Pacific Cod, Pacific Halibut, and Steller sea lion [38]. In some years, juvenile pollock are a major part 

of the diet of older pollock [30]. 

Competitors—Walleye Pollock, an ecologically generalist species, compete with a very wide range of other fish 

species [30].

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years [58].
Walleye Pollock 213

Traditional and Cultural Importance 

None in study area.

Commercial Fisheries

In the United States, Walleye Pollock are not commercially harvested north of the Bering Sea. Walleye Pollock 

was a major food fish in southeastern Alaska and the Gulf of Alaska [59]. Commercial catches by foreign fleets 

began in the early 1950s and increased substantially with the advent of at-sea processing of fish for surimi. 

Currently, the average Alaskan harvest of pollock is 1.1 million metric tons with processed catches destined for 

U.S. and export markets [30, 38, 60].

Potential Effects of Climate Change

It is hypothesized that this species will become more abundant in the U.S. Chukchi and Beaufort Seas marine 

ecosystem changes resulting from climate change. Increased abundance will result in changes in food web 

dynamics such as competition with other gadid species, especially in the Chukchi Sea if benthic-pelagic energy 

flows become decoupled. New experimental results indicate that Walleye Pollock and Pacific Cod grow at 2–3 

times the rate of other Arctic gadids when exposed to increasing temperature regimes in the laboratory that are 

similar to field conditions in summer in the coastal Chukchi and Beaufort Seas. This suggests a potential competitive advantage 

for Walleye Pollock under warming conditions [61].

Areas for Future Research [A]

Field identifications of young pollock may be confused with other gadid species, especially Arctic Cod. A 

rapid diagnostic (genetic) identification tool is needed for field applications as these cods may occur in mixed 

assemblages. Pollock are not well adapted to cold-water environments and, as temperatures warm, monitoring 

programs should be sufficient to detect abrupt changes in abundance. Predator-prey relationships should be 

established to investigate competition with other gadid species. Important spawning and overwintering habitats 

require delineation. Improved information about the species physiological tolerances and growth rate in Arctic 

waters is needed to evaluate potential climate change effects.

References Cited

Bailey, K.M., Powers, D.M., Quattro, J.M., Villa, G., Nishimura, A., Traynor, J.J., and Walters, G., 1999, Population ecology and 

structural dynamics of Walleye Pollock (Theragra chalcogramma), in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the 

Bering Sea: Fairbanks, University of Alaska Sea Grant, AK-SG-99-03, p. 581–613. [30]

Carlson, H.R., 1995, Consistent yearly appearance of age-0 Walleye Pollock, Theragra chalcogramma, at a coastal site in 

southeastern Alaska, 1973–1994: Fishery Bulletin, v. 93, no. 2, p. 386–390. [35]

Dorn, M., Aydin, K., Barbeaux, S., Guttormsen, M., Megrey, B., Spalinger, K., and others, 2007, Gulf of Alaska walleye pollock, 

in The Plan Team for the Groundfish Fisheries of the Gulf of Alaska, compilers, Stock assessment and fishery evaluation 

report for the groundfish resources of the Gulf of Alaska: Anchorage, Alaska, North Pacific Fishery Management Council, 

p. 51–168. [38]

Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p. [20]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

North Pacific Fishery Management Council, 2010, Fishery management plan for groundfish of the Bering Sea and Aleutian 

Islands management area—Appendices—Fishery management plan for groundfish of the BSAI Management Area: North 

Pacific Fisheries Management Council, 233 p. [29]
214 Alaska Arctic Marine Fish Ecology Catalog

Bibliography

1. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

2. Bailey, K.M., and Ciannelli, L., 2007, Walleye pollock, in Spies, R.B., ed., Long-term ecological change in the northern 

Gulf of Alaska: Amsterdam, Elsevier, , p. 85–93.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

5. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

6. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

7. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

8. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

9. Logerwell, E.A., and Rand, K.M., 2010, Beaufort Sea marine fish monitoring 2008—Pilot survey and test of hypotheses: 

Seattle, Washington, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Alaska 

Fisheries Science Center, Final Report, BOEMRE 2010-048, 262 p.

10. Balykin, P.A., 1996, Dynamics and abundance of western Bering Sea walleye pollock, in Mathisen, O.A., and Coyle, K.O., 

eds., Ecology of the Bering Sea: Fairbanks, University of Alaska, Alaska Sea Grant College Program, p. 177–182.

11. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

12. Gustafson, R.G., Lenarz, W.H., McCain, B.B., Schmitt, C.C., Grant, W.S., Builder, T.L., and Methot, R.D., 2000, Status 

review of Pacific hake, Pacific cod, and walleye pollock from Puget Sound, Washington: Seattle, Washington, U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFSNWFSC-44. 

13. Orlov, A.M., 2005, Bottom trawl-caught fishes and some features of their vertical distribution in the Pacific waters off the 

north Kuril Islands and south-east Kamchatka, 1993–1999: Aqua, Journal of Ichthyology and Aquatic Biology of Fishes, 

v. 9, no. 4, p. 139–160.

14. Martin, M.H., and Clausen, D.M., 1995, Data report—1993 Gulf of Alaska Bottom Trawl Survey: U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum, NMFS-AFSC-59, 217 p.

15. Zenger, H.H., Jr., 2004, Data report—2002 Aleutian Islands bottom trawl survey: U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS-AFSC-143, 247 p.

16. Hoff, G.R., and Britt, L.L., 2003, The 2002 eastern Bering Sea upper continental slope survey of groundfish and 

invertebrate resources: National Oceanic and Atmospheric Administration, Technical Memorandum NMFS-AFSC-141, 

286 p.

17. Hoff, G.R., and Britt, L.L., 2005, Results of the 2004 Eastern Bering Sea upper continental slope survey of groundfish and 

invertebrate resources: National Oceanic and Atmospheric Administration, Technical Memorandum, 300 p.
Walleye Pollock 215

18. Hinckley, S., 1987, The reproductive biology of walleye pollock, Theragra chalcogramma, in the Bering Sea, with 

reference to spawning stock structure: Fishery Bulletin, v. 85, no. 3, p. 481–498.

19. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

20. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

21. Siefert, D.L., Incze, L.S., and Ortner, P.B., 1988, Vertical distribution of zooplankton, including ichthyoplankton, in 

Shelikof Strait, Alaska—Data from fisheries-oceanography coordinated investigations (FOCI) cruise in May 1986: National 

Marine Fisheries Service, NWAFC Processed Report, v. 88, no. 28, 232 p.

22. Kendall, J., Arthur W., Incze, L.S., Ortner, P.B., Cummings, S.R., and Brown, P.K., 1994, The vertical distribution of eggs 

and larvae of walleye pollock, Theragra chalcogramma, in Shelikof Strait, Gulf of Alaska: Fishery Bulletin, v. 92, no. 3, 

p. 540–554.

23. Duffy-Anderson, J.T., Busby, M.S., Mier, K.L., Deliyanides, C.M., and Stabeno, P.J., 2006, Spatial and temporal patterns in 

summer ichthyoplankton assemblages on the eastern Bering Sea shelf 1996–2000: Fisheries Oceanography, v. 15, no. 1, 

p. 80–94.

24. Nishiyama, R., Hirano, K., and Haryu, T., 1986, The early life history and feeding habits of larval walleye pollock Theragra 

chalcogramma (Pallas) in the southeast Bering Sea, in Symposium on biology, stock assessment, and management of 

pollock, Pacific cod and hake in the North Pacific region: Standing Committee on Biology and Research, Anchorage, 

Alaska, October 1983, International North Pacific Fisheries Commission Bulletin, Issue 45, p. 177–227.

25. Kendall, J., Arthur W., Clarke, M.E., Yoklavich, M.M., and Boehlert, G.W., 1987, Distribution, feeding, and growth of 

larval walleye pollock, Theragra chalcogramma, from Shelikof Strait, Gulf of Alaska: Fishery Bulletin, v. 85, no. 3, 

p. 499–521.

26. Siefert, D.L., Incze, L.S., and Ortner, P.B., 1990, Vertical distribution of zooplankton, including ichthyoplankton, in 

Shelikof Strait, Alaska—Data from fisheries-oceanography coordinated investigations (FOCI) cruise in May 1987: National 

Oceanic and Atmospheric Administration, National Marine Fisheries Service, Northwest and Alaska Fisheries Center 

(NWAFC) Processed Report v. 90, no. 5, 129 p.

27. Mason, J.C., 1985, The fecundity of the walleye pollock, Theragra chalcogramma (Pallas), spawning in Canadian waters: 

Journal of Fish Biology, v. 27, no. 3, p. 335–346.

28. Dunn, J.R., and Matarese, A.C., 1987, A review of the early life history of northeast Pacific gadoid fishes: Fisheries 

Research, v. 5, nos. 2–3, p. 163–184.

29. North Pacific Fishery Management Council, 2010, Fishery management plan for groundfish of the Bering Sea and Aleutian 

Islands management area—Appendices—Fishery management plan for groundfish of the BSAI Management Area: North 

Pacific Fisheries Management Council, 233 p.

30. Bailey, K.M., Powers, D.M., Quattro, J.M., Villa, G., Nishimura, A., Traynor, J.J., and Walters, G., 1999, Population 

ecology and structural dynamics of Walleye Pollock (Theragra chalcogramma), in Loughlin, T.R., and Ohtani, K., eds., 

Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, AK-SG-99-03, p. 581–613.

31. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.

32. Smith, G.B., 1981, The biology of walleye pollock, in Hood, D.W., and Calder, J.A., eds.: The eastern Bering Sea shelf—

Oceanography and resources, .Volume 1: Seattle, University of Washington Press, p. 527–551.

33. Stahl, J.P., and Kruse, G.H., 2008, Spatial and temporal variability in size at maturity of Walleye Pollock in the eastern 

Bering Sea: Transactions of the American Fisheries Society, v. 137, no. 5, p. 1,543–1,557.
216 Alaska Arctic Marine Fish Ecology Catalog

34. Walters, G.E., 1984, Ecological aspects of larval and juvenile Pacific cod (Gadus macrocephalus), walleye pollock 

(Theragra chalcogramma), and Pacific tomcod (Microgadus proximus) in Port Townsend, Washington: Seattle, University 

of Washington, Master’s thesis, 129 p.

35. Carlson, H.R., 1995, Consistent yearly appearance of age-0 Walleye Pollock, Theragra chalcogramma, at a coastal site in 

southeastern Alaska, 1973–1994: Fishery Bulletin, v. 93, no. 2, p. 386–390.

36. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

37. Johnson, S.W., Thedinga, J.F., and Lindeberg, M.R., 2012, Nearshore fish atlas of Alaska: National Oceanic and 

Atmospheric Administration Fisheries, accessed February 2012 at http://www.fakr.noaa.gov/habitat/fishatlas/.

38. Dorn, M., Aydin, K., Barbeaux, S., Guttormsen, M., Megrey, B., Spalinger, K., and others, 2007, Gulf of Alaska walleye 

pollock, in The Plan Team for the Groundfish Fisheries of the Gulf of Alaska, compilers, Stock assessment and fishery 

evaluation report for the groundfish resources of the Gulf of Alaska: Anchorage, Alaska, North Pacific Fishery Management 

Council, p. 51–168.

39. Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution in the Kronitsky Bay in relation to abiotic factors: Zoologicheskii 

Zhurnal, v. 63, p. 1,363–1,373.

40. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

41. Brodeur, R.D., and Rugen, W.C., 1994, Diel vertical distribution of ichthyoplankton in the northern Gulf of Alaska: Fishery 

Bulletin, v. 92, no. 2, p. 223–235.

42. Adams, C.F., Foy, R.J., Kelley, J.J., and Coyle, K.O., 2009, Seasonal changes in the diel vertical migration of walleye 

pollock (Theragra chalcogramma) in the northern Gulf of Alaska: Environmental Biology of Fishes, v. 86, no. 2, p. 

297–305.

43. Brodeur, R.D., 1998, In situ observations of the association between juvenile fishes and scyphomedusae in the Bering Sea: 

Marine Ecology Progress Series, v. 163, p. 11–20.

44. Van Hyning, J.M., and Cooney, R.T., 1974, Association of walleye pollock, Theragra chalcogramma, with the jellyfish, 

Cyanea: Copeia, v. 1974, no. 3, p. 791.

45. Kotwicki, S., Buckley, T.W., Honkalehto, T., and Walters, G.E., 2005, Variation in the distribution of walleye pollock 

(Theragra chalcogramma) with temperature and implications for seasonal migration: Fishery Bulletin, v. 103, no. 4, 

p. 574–587.

46. Wespestad, V.G., Fritz, L.W., Ingraham, W.J., and Megrey, B.A., 2000, On relationships between cannibalism, climate 

variability, physical transport, and recruitment success of Bering Sea walleye pollock (Theragra chalcogramma): ICES 

Journal of Marine Science, v. 57, no. 2, p. 272–278.

47. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

48. Wilson, M.T., Jump, C.M., and Duffy-Anderson, J.T., 2006, Comparative analysis of the feeding ecology of two pelagic 

forage fishes—Capelin Mallotus villosus and Walleye Pollock Theragra chalcogramma: Marine Ecology Progress Series, 

v. 317, p. 245–258.

49. Olsen, J.B., Merkouris, S.E., and Seeb, J.E., 2002, An examination of spatial and temporal genetic variation in walleye 

pollock (Theragra chalcogramma) using allozyme, mitochondrial DNA, and microsatellite data: Fishery Bulletin, v. 100, 

no. 4, p. 752–764.

50. Brykov, V.A., Polyakova, N.E., Priima, T.F., and Katugin, O.N., 2004, Mitochondrial DNA variation in northwestern Bering 

Sea walleye pollock, Theragra chalcogramma (Pallas): Environmental Biology of Fishes, v. 69, nos. 1–4, p. 167–175.

51. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific, Part V: Soviet Fisheries Investigation in the Northeastern Pacific, p. 161–224. 

[Russian translation by Israel Program for Scientific Translations, Jerusalem (1972).]
Walleye Pollock 217

52. Baird, T.A., and Olla, B.L., 1991, Social and reproductive behavior of a captive group of walleye pollock, Theragra 

chalcogramma: Environmental Biology of Fishes, v. 30, no. 3, p. 295–301.

53. Simenstad, C.A., Isakson, J.S., and Nakatani, R.E., 1977, Marine fish communities, in Merritt, M.L., and Fuller, R.G., eds., 

The environment of Amchitka Island, Alaska: National Technical Information Center, Energy Research and Development 

Administration TID-26712, p. 451–492.

54. Yang, M.-S., 1996, Diets of the important groundfishes in the Aleutian Islands in summer 1991: Seattle, Washington, U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, Technical Memorandum NMFS-AFSC-60, 

105 p.

55. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, Technical Memorandum NMFS-AFSC-112, 174 p.

56. Sturdevant, M.V., Brase, A.L., and Hulbert, L.B., 2001, Feeding habits, prey fields, and potential competition of young-ofthe-year walleye pollock (Theragra chalcogramma) and Pacific herring (Clupea pallasi) in Prince William Sound, Alaska, 

1994–1995: Fishery Bulletin, v. 99, no. 3, p. 482–501.

57. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320. 

58. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

59. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

60. Witherell, D., 2000, Groundfish of the Bering Sea and Aleutian Islands area—Species profiles 2001: Anchorage, Alaska, 

North Pacific Fishery Management Council, 15 p.

61. Laurel, B.J., Spencer, M., Iseri, P., and Copeman, L.A., 2015, Temperature-dependent growth and behavior of juvenile 

Arctic cod (Boreogadus saida) and co-occurring North Pacific gadids: Polar Biology, no. 39, p. 1,127–1,135.
218 Alaska Arctic Marine Fish Ecology Catalog

Pacific Cod (Gadus macrocephalus) 

Tilesius, 1810

Family Gadidae 

Note on taxonomy: Based on analyses of both morphology and 

mtDNA, the Greenland Cod, Gadus ogac (Richardson, 1836), is a 

subspecies of G. macrocephalus [1, 2]. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Of little known ecological importance in U.S. 

Chukchi and Beaufort Seas. Maybe of small seasonal importance in 

food webs in the Bering Strait and southeastern Chukchi Sea.

Physical Description/Attributes: Robust body, large head. Light 

gray-brown with brown to bright golden yellow spots on back and sides to olive-blackish with no distinct spots [3, 4]. For 

specific diagnostic characteristics see Fishes of Alaska (Mecklenburg and others, 2002, p. 296) [3]. Swim bladder: Present [3]. 

Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [1, 3]. Elsewhere in Alaska, throughout Bering Sea, Aleutian Islands and Gulf of 

Alaska. Worldwide, in Pacific Ocean south to southern California and to Yellow Sea off Manchuria, China; east across Canada 

to west Greenland and south to Gulf of St. Lawrence. Isolated population in White Sea [1, 3].

Relative Abundance: Uncommon in U.S. Chukchi and Beaufort Seas [1, 3]. Uncommon in northern Bering Sea [1]. Abundant 

throughout Bering Sea northward to Norton Sound and Gulf of Anadyr [1, 6, 7], and southward to Seas of Japan and Okhotsk [8, 

9] and Washington [10].

Pacific Cod (Gadus macrocephalus) 597 mm, western Gulf 

of Alaska, 2005. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Pacific Cod (Gadus macrocephalus) within Arctic Outer Continental Shelf Planning 

Areas [5] based on review of published literature and specimens from historical and recent collections [1, 3, 4].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Pacific Cod

Gadus macrocephalus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Pacific Cod 219

Depth Range: In water column, near surface to near bottom depth of 875 m [3], typically 50–300 m; sometimes in surf zone 

[11–13]. Spawning takes place at 40–265 m [14]. Fertilized eggs are benthic on continental shelf [10]. Newly hatched larvae are 

primarily in upper 45 m of water column (highest abundances at 15–30 m) [14], moving downward as they grow [15]. Juveniles 

are mainly at 60–150 m in Gulf of Alaska and eastern Bering Sea [15].

Habitats and Life History

Eggs—Size: 1.0–1.2 mm [16, 17]. Time to hatching: 8.5–28 days at 11–4.5 °C. Hatching is most successful at 

lower temperatures [16, 18]. Habitat: Benthic [10, 16]. 

Larvae—Size at hatching: 3.0–4.0 mm [16, 17]. Size at juvenile transformation: 2.5–3.5 cm FL [14, 19]. Days 

to juvenile transformation: Unknown. Yolk sac is absorbed in 10 days [14]. Habitat: Pelagic and neritic [17]. 

Juveniles—Age and size: 2.5 cm FL to 38–81 cm TL [13, 14, 19]. Habitat: Shallow nearshore waters at [14, 17, 

20], initially associated with algae and eelgrass but later in their first year some fish migrate into deeper water 

and over a wide range of habitats including plants, soft substrates, and mounds formed by sea cucumbers [10, 18, 

21–23]. 

Adults—Age and size at first maturity: In eastern Bering Sea, 50 percent of females were mature at 58.0 cm TL

and 4.9 years, whereas 50 percent of those in the Gulf of Alaska were mature at 50.3 cm TL and 4.4 years. A 

few females were mature at as small as 38 cm TL and a few were immature until about 81 cm TL [13]. Length at 

maturity is highly dependent on environmental factors and varies widely between areas and years. For example, 

off British Columbia. Canada, length at 50 percent maturity differed by almost 10 cm between samples taken 

in the mid-1970s and mid-1980s [24]. In Gulf of Alaska and eastern Sea of Okhotsk, females grow larger than 

males, although both sexes reach about the same maximum length in the eastern Bering Sea [13]. In Gulf of 

Alaska (although not in the eastern Bering Sea), male and female growth rates differ. Apparently, cod living in 

Alaskan waters grow more slowly but reach a larger size and live longer than those living off British Columbia 

and Washington [14]. Maximum age: 17 years [25], but rarely beyond 14 years [13]. Maximum size: 120 cm TL 

[3]. Habitat: Pelagic, both near the bottom and in the midwaters [14] over soft sea floors [10, 17, 22, 23]. 

Substrate—Cobble and rocky bottoms [10, 22, 23]. Coarse sand and cobble for spawning and eggs [14]. 

Physical/chemical—Temperature: -1.7–18 °C, mainly 0–10 °C [21, 26, 27]. Salinity: Eggs are in polyhaline to 

euhaline waters. Marine and estuarine [14].

Benthic and reproductive distribution of Pacific Cod (Gadus macrocephalus).

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

500

400

300

200

100

0

Juveniles

Some fish to 875 m

Gadus macrocephalus

Pacific Cod

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Larvae

Adults Spawning and eggs
220 Alaska Arctic Marine Fish Ecology Catalog

Behavior

Diel—Pelagic juveniles have been found in surface waters in association with medusae [28]. 

Seasonal—Make annual inshore and offshore movements linked to spawning and feeding. The timing and extent 

of annual migrations vary with location. Annual migrations in eastern Bering Sea, eastern Aleutian Islands, and 

Gulf of Alaska, occur when fish attempt to avoid temperature extremes that accompany the seasonal changes 

[29]. Fish move offshore during winter, as nearshore waters get very cold, and move inshore during summer [30]. 

Farther south on both sides of the Pacific Ocean (for example, Puget Sound, Korea, and Japan), migrations to 

deeper waters occur during summer months to avoid excessively heated coastal waters and return inshore for the 

winter [10]. Some fish move fairly long distances. Pacific Cod in the eastern Bering Sea, for instance, summer on 

the eastern Bering Sea shelf, then move southward and deeper to the Bering Sea in the vicinity of Unimak Pass 

and Unalaska Island, Alaska, and in the nearby Gulf of Alaska to spawn [14]. 

Reproductive—Single batch spawners, releasing all eggs in a few minutes [14]. Spawning depth depends on 

its depth-temperature profile. For instance, off Washington and southwest Vancouver Island, British Columbia, 

Pacific Cod spawn in more shallow waters than those in northern British Columbia [10]. 

Schooling—Forms schools [14]. 

Feeding—Juveniles and adults are carnivorous and feed at night [14]. Feeding increases during the summer and 

decreases in winter [31].

Populations or Stocks

Fish in Puget Sound and the Strait of Georgia may form several semi-isolated populations from fish on the outer 

coast [14, 32]. Based on analyses of morphology and mtDNA, Pacific Cod are most closely related to Ogac 

(Gadus ogac) and are now considered by taxonomists to be the same species [1, 2, 33].

Reproduction

Mode—Oviparous, external fertilization [14]. 

Spawning season—Can occur between January and July, although peak spawning takes place in spring in 

Alaska [13, 34]. 

Fecundity—225,000 –6,400,000 semi-adhesive eggs [14, 35]. The number of eggs produced per body length 

decreases with higher latitude [21].

Food and Feeding

Food items—Very wide range of benthic and water column organisms. Cod less than about 20 cm FL feed 

primarily on a wide range of crustaceans, such as copepods, gammarid and caprellid amphipods, mysids, and 

euphausiids, and some small fishes [36–39]. Larger fish add large numbers of fishes to their diets as well as 

shrimps, crabs, hermit crabs, polychaetes, snails, clams, squids, and octopuses [38, 40–42]. As Pacific Cod grow, 

they feed more heavily on fishes and less on invertebrates [43–44]. 

Trophic level—4.1 [45].

Biological Interactions

Predators—A large number of fishes, sea birds, and marine mammals. Major fish predators include Arrowtooth 

Flounder, Flathead Sole, Pacific Cod, Pacific Halibut, Sablefish, Spotted Spiny Dogfish, Walleye Pollock, and 

Yellowfin Sole [40, 46–48]. Seabirds include Common Murres, Horned and Tufted Puffins [49, 50]. Marine 

mammals include beluga, fin, minke, and sperm whales and orcas; bearded, harbor, northern fur, and ribbon 

seals; and Stellar sea lions [51–55]. 

Competitors—Other gadids, along with flatfishes, sculpins, poachers, and eelpouts.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years [56].
Pacific Cod 221

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Pacific Cod are not commercially harvested.

Potential Effects of Climate Change

It might be expected that Pacific Cod abundance will increase in Arctic waters if fish from the Bering Sea move 

northward. The probability of this species’ colonization of Arctic marine environments may be lower than for 

other gadid species because of its apparent fidelity to spawning locations in the Bering Sea. However, new 

experimental results indicate that Pacific Cod and Walleye Pollock grow at 2–3 times the rate of other Arctic 

gadids when exposed to increasing temperature regimes in laboratory that are similar to field conditions in 

summer in the coastal Chukchi and Beaufort Seas [57]. This suggests a potential competitive advantage for Pacific Cod under 

warming conditions.

Areas for Future Research [A]

Little is known about the ecology of this species in the study area. Whether Pacific Cod spawn in the Chukchi or 

Beaufort Seas is of major interest. Researchers believe that the cod display high fidelity to spawning areas in the 

Bering Sea and thus improved information about their migratory behavior is needed. 

Remarks

This species has long been known to be present in the Beaufort Sea (see [Walters, 1955], between Point Barrow and Smith Bay)

[58], under the name, Gadus ogac [3], and is common just over the U.S-Canadian border in Canadian waters. 

References Cited

Dunn, J.R., and Matarese, A.C., 1987, A review of the early life history of northeast Pacific gadoid fishes: Fisheries Research, v. 

5, nos. 2–3, p. 163–184. [17]

Forrester, C.R., 1964, Laboratory observations on embryonic development and larvae of the Pacific cod (Gadus macrocephalus 

Tilesius): Journal of the Fisheries Research Board of Canada, v. 21, no. 1, p. 9–16. [16]

Gustafson, R.G., Lenarz, W.H., McCain, B.B., Schmitt, C.C., Grant, W.S., Builder, T.L., and Methot, R.D., 2000, Status review 

of Pacific hake, Pacific cod, and walleye pollock from Puget Sound, Washington: Seattle, Washington, U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFS-NWFSC-44. [14]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [1]
222 Alaska Arctic Marine Fish Ecology Catalog

Palsson, W.A., 1990, Pacific cod (Gadus macrocephalus) in Puget Sound and adjacent waters—Biology and stock assessment: 

Olympia, State of Washington, Department of Fisheries, Technical Report no. 112, 137 p. [10]

Stark, J.W., 2007, Geographic and seasonal variations in maturation and growth of female Pacific cod (Gadus macrocephalus) in 

the Gulf of Alaska and Bering Sea: Fishery Bulletin, v. 105, no. 3, p. 396–407. [13]

Bibliography

1. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

2. Teletchea, F., Laudet, V., and Hänni, C., 2006, Phylogeny of the Gadidae (sensu Svetovidov, 1948) based on their 

morphology and two mitochondrial genes: Molecular Phylogenetics and Evolution, v. 38, no. 1, p. 189–199.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p.

6. Vinnikov, A.V., 1996, Pacific cod (Gadus macrocephalus) of the western Bering Sea, in Mathisen, O.A., and Coyle, K.O., 

eds., Ecology of the Bering Sea: Fairbanks, University of Alaska, Alaska Sea Grant College Program, p. 183–202.

7. Hamazaki, T., Fair, L., Watson, L., and Brennan, E., 2005, Analyses of Bering Sea bottom-trawl surveys in Norton Sound—

Absence of regime shift effects on epifauna and demersal fish: ICES Journal of Marine Science, v. 62, no. 8, p. 1,597–1,602.

8. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

9. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

10. Palsson, W.A., 1990, Pacific cod (Gadus macrocephalus) in Puget Sound and adjacent waters—Biology and stock 

assessment: Olympia, State of Washington, Department of Fisheries, Technical Report no. 112, 137 p.

11. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

12. Zenger, H.H., Jr., 2004, Data report—2002 Aleutian Islands bottom trawl survey: U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS-AFSC-143, 247 p.

13. Stark, J.W., 2007, Geographic and seasonal variations in maturation and growth of female Pacific cod (Gadus 

macrocephalus) in the Gulf of Alaska and Bering Sea: Fishery Bulletin, v. 105, no. 3, p. 396–407.

14. Gustafson, R.G., Lenarz, W.H., McCain, B.B., Schmitt, C.C., Grant, W.S., Builder, T.L., and Methot, R.D., 2000, Status 

review of Pacific hake, Pacific cod, and walleye pollock from Puget Sound, Washington: Seattle, Washington, U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFSNWFSC-44. 

15. North Pacific Fishery Management Council, 2010, Fishery management plan for groundfish of the Bering Sea and Aleutian 

Islands management area—Appendices—Fishery management plan for groundfish of the BSAI Management Area: North 

Pacific Fisheries Management Council, 233 p.
Pacific Cod 223

16. Forrester, C.R., 1964, Laboratory observations on embryonic development and larvae of the Pacific cod (Gadus 

macrocephalus Tilesius): Journal of the Fisheries Research Board of Canada, v. 21, no. 1, p. 9–16.

17. Dunn, J.R., and Matarese, A.C., 1987, A review of the early life history of northeast Pacific gadoid fishes: Fisheries 

Research, v. 5, no. 2–3, p. 163–184.

18. Abookire, A.A., Duffy-Anderson, J.T., and Jump, C.M., 2007, Habitat associations and diet of young-of-the-year Pacific 

cod (Gadus macrocephalus) near Kodiak, Alaska: Marine Biology, v. 150, no. 4, p. 713–726.

19. Matarese, A.C., Kendall, J., Arthur W., Blood, D.M., and Vinter, B.M., 1989, Laboratory guide to early life history stages 

of northeast Pacific fishes: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National 

Marine Fisheries Service Technical Report NMFS 80, 652 p.

20. Blackburn, J.E., and Jackson, P.B., 1982, Seasonal composition and abundance of juvenile and adult marine finfish and crab 

species in the nearshore zone of Kodiak Island’s eastside during April 1978 through March 1979: Outer Continental Shelf 

Environmental Assessment Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 377–570.

21. Alderdice, D.F., and Forrester, C.R., 1971, Effects of salinity, temperature, and dissolved oxygen on early development of 

the Pacific cod (Gadus macrocephalus): Journal of the Fisheries Research Board of Canada, v. 28, no. 6, p. 883–902.

22. Laurel, B.J., Stoner, A.W., Ryer, C.H., Hurst, T.P., and Abookire, A.A., 2007, Comparative habitat associations in juvenile 

Pacific cod and other gadids using seines, baited cameras and laboratory techniques: Journal of Experimental Marine 

Biology and Ecology, v. 351, nos. 1–2, p. 42–55.

23. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, nos. 1–3, p. 15–28.

24. Welch, D.W., and Foucher, R.P., 1988, A maximum likelihood for estimating length-at-maturity with application to Pacific 

cod (Gadus macrocephalus) population dynamics: Canadian Journal of Fisheries and Aquatic Sciences, v. 45, no. 2, 

p. 333–343.

25. Kimura, D.K., National Marine Fisheries Service, written commun., 2010.

26. Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution in the Kronitsky Bay in relation to abiotic factors: Zoologicheskii 

Zhurnal, v. 63, p. 1,363–1,373.

27. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

28. Van Hyning, J.M., and Cooney, R.T., 1974, Association of walleye pollock, Theragra chalcogramma, with the jellyfish, 

Cyanea: Copeia, v. 1974, no. 3, p. 791.

29. Shimada, A.M., and Kimura, D.K., 1994, Seasonal movements of Pacific cod, Gadus macrocephalus, in the eastern Bering 

Sea and adjacent waters based on tag-recapture data: Fishery Bulletin, v. 92, no. 4, p. 800–816.

30. Witherell, D., 2000, Groundfish of the Bering Sea and Aleutian Islands area—Species profiles 2001: Anchorage, Alaska, 

North Pacific Fishery Management Council, 15 p.

31. Tokranov, A.M., 1992, Features of feeding of benthic predatory fishes of the west Kamchatka shelf: Journal of Ichthyology, 

v. 32, no. 7, p. 45–55.

32. Gao, Y., Bargmann, G.G., Brand, U., and Noakes, D.L.G., 2005, Stable isotopic and trace elemental compositions of 

otoliths and the stock structure of Pacific cod, Gadus macrocephalus: Environmental Biology of Fishes, v. 74, nos. 3–4, 

p. 335–348.

33. Carr, S.M., Kivlichan, D.S., Pepin, P., and Crutcher, D.C., 1999, Molecular systematics of gadid fishes—Implications for 

the biogeographical origins of Pacific species: Canadian Journal of Zoology, v. 77, no. 1, p. 19–26.

34. Hirschberger, W.A., and Smith, G.B., 1983, Spawning of twelve groundfish species in the Alaska and Pacific Coast 

regions, 1975–81: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical 

Memorandum NMFS F/NWC-44, 50 p.
224 Alaska Arctic Marine Fish Ecology Catalog

35. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

36. Walters, G.E., 1984, Ecological aspects of larval and juvenile Pacific cod (Gadus macrocephalus), walleye pollock 

(Theragra chalcogramma), and Pacific tomcod (Microgadus proximus) in Port Townsend, Washington: Seattle, University 

of Washington, Master’s thesis.

37. Tokranov, A.M., and Vinnikov, A.B., 1991, Diet of the Pacific cod, Gadus morhua macrocephalus, and its position in the 

food chain in Kamchatkan coastal waters: Journal of Ichthyology, v. 31, no. 5, p. 84–98.

38. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.

39. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-158, 249 p.

40. Lang, G.M., Livingston, P.A., Pacunski, R.E., Parkhurst, J., and Yang, M.-S., 1991, Groundfish food habits and predation on 

commercially important prey species in the eastern Bering Sea from 1984 to 1986: Seattle, Washington, U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFS F/NWC-207, 240 p.

41. Ortiz, I., and Aydin, K., 2003, Feeding habits of demersal fish: Alaska Fisheries Science Center Quarterly Report, October–

December 2003, p. 49.

42. Yang, M.-S., 2004, Diet changes of Pacific cod (Gadus macrocephalus) in Pavlof Bay associated with climate changes in 

the Gulf of Alaska between 1980 and 1995: Fishery Bulletin, v. 102, no. 2, p. 400–405.

43. Lang, G.M., and Livingston, P.A., 1996, Food habits of key groundfish species in the eastern Bering Sea Slope region: 

Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical 

Memorandum NMFS-AFSC-67, 120 p.

44. Chuchukalo, V.I., Radchenko, V.I., Nadtochii, V.A., Koblikov, V.N. and Slabinskii, A.M., 1999, Feeding and some features 

of ecology of Gadidae of the western Kamchatka shelf in summer 1996: Journal of Ichthyology, v. 39, no. 4, p. 309–321.

45. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320. 

46. Bonham, K., 1954, Food of the dogfish Squalus acanthias: Washington Department of Fisheries, v. 1, no. 2, p. 25–36.

47. Yang, M.-S., 1996, Diets of the important groundfishes in the Aleutian Islands in summer 1991: Seattle, Washington, U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFSAFSC-60, 105 p.

48. Yang, M.-S., Dodd, K., Hibpshman, R., and Whitehouse, A., 2006, Food habits of groundfishes in the Gulf of Alaska 

in 1999 and 2001: U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical 

Memorandum, NMFS-AFSC-164, 189 p.

49. Krasnow, L.D., and Sanger, G.A., 1982, Feeding ecology of marine birds in the nearshore waters of Kodiak Island: 

Anchorage, Alaska, Final Report to the Outer Continental Shelf Environmental Assessment Program, U.S. Fish and Wildlife 

Service, National Fisheries Research Center, Migratory Bird Section, Contract 01-5-022-2538.

50. Hatch, S.A., and Sanger, G.A., 1992, Puffins as samplers of juvenile pollock and other forage fish in the Gulf of Alaska: 

Marine Ecology Progress Series, v. 80, p. 1–14.

51. Harry, G.Y., and Hartley, J.R., 1981, Northern fur seals in the Bering Sea, in Hood, D.W., and Calder, J.A., eds., The Eastern 

Bering Sea shelf—Oceanography and resources: Seattle, University of Washington Press, p. 847–867.

52. Antonelis, G.A., Melin, S.R., and Bukhtiyarov, Y.A., 1994, Early spring feeding habits of bearded seals (Erignathus 

barbatus) in the central Bering Sea, 1981: Arctic, v. 47, no. 1, p. 74–79.
Pacific Cod 225

53. Frost, K.J., and Lowry, L.F., 1981, Foods and trophic relationships of cetaceans in the Bering Sea, in Hood, D.W., and 

Calder, J.A., eds., The eastern Bering Sea shelf—Oceanography and resources: Seattle, University of Washington Press, 

p. 825–836.

54. Frost, K.J., and Lowry, L.F., 1981, Trophic importance of some marine gadids in northern Alaska and their body-otolith size 

relationships: Fishery Bulletin, v. 79, no. 1, p. 187–192.

55. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990.

56. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

57. Laurel, B.J., Spencer, M., Iseri, P., and Copeman, L.A., 2015, Temperature-dependent growth and behavior of juvenile 

Arctic cod (Boreogadus saida) and co-occurring North Pacific gadids: Polar Biology, no. 39, p. 1,127–1,135.

58. Walters, V., 1955, Fishes of western Arctic America and eastern Arctic Siberia: Bulletin of the American Museum of Natural 

History, v. 106, p. 259–368.
226 Alaska Arctic Marine Fish Ecology Catalog

Threespine Stickleback to Antlered Sculpin

Threespine Stickleback (Gasterosteus aculeatus) 

Linnaeus, 1758

Family Gasterosteidae 

Note: Exhibits highly variable life history patterns with marine, 

anadromous, riverine, and lacustrine populations [1, 2]. Data in 

this account are from marine and anadromous populations.

Colloquial Name: Iñuit—Kakalisauraq [3].

Ecological Role: In locations where common, for instance in 

Kotzebue Sound, this species may be of ecological importance in 

local food webs.

Physical Description/Attributes: Moderately elongate body. Anadromous type is blue-black to silvery or greenish with 

yellow, silvery, or white bellies. Breeding males become bright blue or green with red or orange throats and bellies. For specific 

diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 333) [4]. Swim bladder: Present, without 

pneumatic duct [5]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [6]. Elsewhere in Alaska, in all coastal waters. Worldwide, from Bering Sea south to 

Monterey Bay, central California, and to Seas of Okhotsk and Japan; in Atlantic Ocean from Hudson Bay to southern Greenland, 

Iceland, and southern Barents Sea to Novaya Zemlya, Russia [6].

Threespine Stickleback (Gasterosteus aculeatus), 42 mm 

TL, northeastern Chukchi Sea, 2007. Photograph by C.W. 

Mecklenburg, Point Stephens Research.
Threespine Stickleback 227

Relative Abundance: Common in brackish water at least as far north as Kotzebue Sound, U.S. Chukchi Sea, and occasional or 

rare east in U.S. Beaufort Sea [1, 9–11]. Rare in Northwest Territories, Canada [1]. Common in southwestern Barents Sea and 

Sea of Japan [6, 12].

tac14-5222_fig3-6-1m_Gasterosteus_aculeatus

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Threespine Stickleback

Gasterosteus aculeatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Threespine Stickleback (Gasterosteus aculeatus), within Arctic Outer Continental 

Shelf Planning Areas [7] based on review of published literature and specimens from historical and recent 

collections [6, 8]. 
228 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Intertidal to 27 m, as far as 805 km offshore for juveniles and adults [4, 13]. Larvae are abundant in surface 

waters [14]. Marine type spawns in shallow waters, such as tidepools [15]. 

Benthic and reproductive distribution of Threespine Stickleback (Gasterosteus aculeatus).

3-6-1ThreespineStickleback

Open ocean

Depth, in meters

500

400

300

200

100

0

Adults and juveniles

Gasterosteus aculeatus

Threespine Stickleback

Benthic distribution

Some fish 

to 27 m

Depth, in meters

Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Spawning and eggs

Reproductive distribution

Larvae

Data from outside U.S. Beaufort-Chukchi Seas 500

400

100

200

300

0

Habitats and Life History

Anadromous and marine [1, 2]. 

Eggs—Size: 0.11–0.16 cm [16]. Time to hatching: 5–20 days [12]. Habitat: Benthic, in tidepools for marine 

type [2]. 

Larvae—Size at hatching: 2.0–5.5 mm [17, 18]. Size at juvenile transformation: About 10.0 mm [19]. Days to 

juvenile transformation: About 30 days [19, 20]. Habitat: Benthic to pelagic [14]. 

Juveniles—Age and size: 1–12 months [19], and 11–30 mm TL [19]. Habitat: Benthic to pelagic, staying close 

to nests for 4–6 days [4], around eelgrass, filamentous algae, and other plants, as well as over sand and rocks [21, 

22]. 

Adults—Age and size at first maturity: Some mature after 1 year (fish spawn once and die), others take 2 years 

[16]. Growth rates vary with area [16, 23]. Maximum age: 5 years [18], typically 1–3.5 years. Varies with area 

[16, 23]. Maximum size: 11 cm TL [18]. Habitat: Benthic to pelagic [4], around eelgrass, filamentous algae, and 

other plants, as well as over sand and rocks, and in offshore waters [21, 22, 24, 25]. 

Substrate—Over rocks, silt, and sand for spawning [15]. 

Physical/chemical—Temperature: Marine type has been shown to tolerate temperatures as low as 4 °C and as 

high as 28 °C in laboratory experiments [26]. Salinity: Fresh to marine waters. More common in brackish than 

marine waters in Kotzebue and Norton sounds [9].

Behavior

Diel—In Puget Sound, both juveniles and adults inhabit surface waters at night [27, 28]. 

Seasonal—Some fish migrate into coastal water in autumn to over winter [15]. Other fish winter in deep 

water [16]. 

Reproductive—Off Alaska, anadromous fish enter freshwaters to spawn in late spring [1]. Anadromous fish may 

spawn in brackish or fresh waters [15]. Marine fish spawn in quiet areas such as tide pools [2]. Spawning occurs 

over rocks, eelgrass, silt, and sand [15]. Males construct nests composed of bits of plants and other debris held 

together by secretions formed in the kidneys. Through a series of courting behaviors, a male leads a female into 

the nest where she lays her eggs. Many nests contain eggs from more than one female and males guard the eggs 

until they hatch. Neighboring males not guarding eggs often “sneak” into a nearby nest and fertilize some of the 

eggs [16, 29]. Some individuals may spawn once and die [16, 30]. 

Schooling—Forms schools except in spawning season [12]. 

Feeding—Feeds throughout water column [12].
Threespine Stickleback 229

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous, separate sexes [20]. 

Spawning season—May–August in Alaska [1, 16]. 

Fecundity—Batch spawners, laying 50–200 eggs at a time with overall fecundity ranging from 65 to 1,300 

[16, 31]. 

Food and Feeding

Food items—Benthic and midwater prey. Small crustaceans (for example, copepods, euphausiids, mysids, and 

gammarid and caprellid amphipods) often are quite important, and crustacean larvae, insects, worms, mollusks, 

fish eggs, and small fishes are also frequently consumed [18, 32–35]. 

Trophic level—3.51 (standard error 0.49) [36].

Biological Interactions

Predators—Off Alaskan and British Columbia coasts, are a large number of fishes, sea birds, and marine 

mammals [12]. 

Competitors—Likely Polar and Saffron cods, whitefishes, and flatfishes.

1980 1990 2000 2010

Resilience

High, minimum population doubling time less than 15 months (K=0.6–1.8; t

m=1; tmax=4; Fecundity=80) [36].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Threespine Stickleback are not commercially harvested.

Potential Effects of Climate Change

Uncertain; however, this is a species with a very plastic life history pattern, with an ability to adapt to a wide 

range of environmental conditions. Increasing abundance is possible.
230 Alaska Arctic Marine Fish Ecology Catalog

Areas for Future Research [B]

Little studied species in the U.S. Chukchi and Beaufort Seas. Research needs include: (1) depth and location 

of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment; (3) preferred depth 

ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic movements; (6) population 

studies; (7) prey; and (8) predators.

References Cited

Evans, C.L., Reist, J.D., and Minns, C.K., 2002, Life history characteristics of freshwater fishes occurring in the Northwest 

Territories and Nunavut, with major emphasis on riverine habitat requirements: Canadian Manuscript Report of Fisheries and 

Aquatic Sciences, no. 2614, 169 p. [1]

Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p. [12]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [4]

Morrow, J.E., 1980, The freshwater fishes of Alaska: Anchorage, Alaska Northwest Publishing Company, 248 p. [16]

Richardson, E.S., Reist, J.D., and Minns, C.K., 2001, Life history characteristics of freshwater fishes occurring in the Northwest 

Territories and Nunavut, with major emphasis on lake habitat requirements: Canadian Manuscript Report of Fisheries and 

Aquatic Sciences no. 2569, 156 p. [15] 

Bibliography

1. Evans, C.L., Reist, J.D., and Minns, C.K., 2002, Life history characteristics of freshwater fishes occurring in the Northwest 

Territories and Nunavut, with major emphasis on riverine habitat requirements: Canadian Manuscript Report of Fisheries 

and Aquatic Sciences, no. 2614, 169 p.

2. Kume, M., Kuwahara, T., Arai, T., Okamoto, M., and Goto, A., 2006, A part of the Japan Sea form of the threespine 

stickleback, Gasterosteus aculeatus, spawns in the seawater tidal pools of western Hokkaido Island, Japan: Environmental 

Biology of Fishes, v. 77, no. 2, p. 169–175.

3. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

4. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

5. Moser, F., 1903, Comparative embryology of the swim-bladder: Journal of the Royal Microscopical Society, v. 25, p. 708.

6. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

7. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

8. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

9. Barton, L.H., 1978, Finfish resource surveys in Norton Sound and Kotzebue Sound: Alaska Department of Fish and Game, 

Commercial Fisheries Division, p. 75–313.

10. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.
Threespine Stickleback 231

11. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

12. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

13. Simenstad, C.A., Isakson, J.S., and Nakatani, R.E., 1977, Marine fish communities, in Merritt, M.L., and Fuller, R.G., eds., 

The environment of Amchitka Island, Alaska: National Technical Information Center, Energy Research and Development 

Administration TID-26712, p. 451–492.

14. Jump, C.M., Duffy-Anderson, J.T., and Mier, K.L., 2008, Comparison of the Sameoto, Manta and MARMAP neustonic 

ichthyoplankton samplers in the Gulf of Alaska: Fisheries Research, v. 89, no. 3, p. 222–229.

15. Richardson, E.S., Reist, J.D., and Minns, C.K., 2001, Life history characteristics of freshwater fishes occurring in the 

Northwest Territories and Nunavut, with major emphasis on lake habitat requirements: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences no. 2569, 156 p.

16. Morrow, J.E., 1980, The freshwater fishes of Alaska: Anchorage, Alaska Northwest Publishing Company, 248 p.

17. Wang, J.C.S., 1981, Taxonomy of the early life stages of fishes—Fishes of the Sacramento-San Joaquin Estuary and Moss 

Landing Harbor-Elkhorn Slough, California: Concord, California, Ecological Analysts, Inc., 168 p.

18. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

19. Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, M.E., 1991, Distribution and abundance of fishes and inverteprates 

in west coast estuaries, Volume II—Species life history summaries: National Oceanic and Atmospheric Administration/

National Ocean Service Strategic Environmental Assessments Division, ELMR Report no. 8, 327 p.

20. Garrison, K.J., and Miller, B.S., 1982, Review of the early life history of Puget Sound fishes: Seattle, University of 

Washington, Fisheries Research Institute, 729 p.

21. Murphy, M.L., Johnson, S.W., and Csepp, D.J., 2000, A comparison of fish assemblages in eelgrass and adjacent subtidal 

habitats near Craig, Alaska: Alaska Fishery Research Bulletin, v. 7, p. 11–21.

22. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

23. Jones, J.W., and Hynes, H.B.N., 1950, The age and growth of Gasterosteus aculeatus, Pygosteus pungitius and Spinachia 

vulgaris, as shown by their otoliths: Journal of Animal Ecology, v. 19, no. 1, p. 59–73.

24. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

25. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

26. Barrett, R.D.H., Paccard, A., Healy, T.M., Bergek, S., Schulte, P.M., Schluter, D., and Rogers, S.M., 2010, Rapid evolution 

of cold tolerance in stickleback: Proceedings of the Royal Society B, doi:10.1098/rspb.2010.0923.

27. Fresh, K.L., 1979, Distribution and abundance of fishes occurring in the nearshore surface waters of northern Puget Sound, 

Washington: Seattle, University of Washington, Master’s thesis.

28. Miller, B.S., Simenstad, C.A., Moulton, L.L., Fresh, K.L., Funk, F.C., Karp, W.A., and others, 1977, Puget Sound baseline 

program—Nearshore fish survey: University of Washington, Fishery Research Institute, FRI-UW-7710, 219 p.
232 Alaska Arctic Marine Fish Ecology Catalog

29. Zbinden, M., Mazzi, D., Künzler, R., Largiadèr, C.R., and Bakker, T.C.M., 2003, Courting virtual rivals increase ejaculate 

size in sticklebacks (Gasterosteus aculeatus): Behavioral Ecology and Sociobiology, v. 54, no. 3, p. 205–209.

30. Moyle, P.B., 2002, Inland fishes of California: Berkeley, University of California Press, 517 p.

31. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

32. Barraclough, W.E., Robinson, D.G., and Fulton, J.D., 1968, Number, size composition, weight and food of larval and 

juvenile fish caught with a two-boat surface trawl in Saanich Inlet April 23–July 21 (data record): Fisheries Research Board 

of Canada, Manuscript Report Series no. 1004, 305 p.

33. Tack, S.L., 1970, The summer distribution and standing stock of the fishes of Izembek Lagoon, Alaska: Fairbanks, 

University of Alaska, Master’s thesis.

34. Cross, J.N., Fresh, K., Miller, B.S., Simenstad, C.A., Steinfort, S.N., and Fegley, J.C., 1978, Nearshore fish and 

invertebrates assemblages along the Strait of Juan de Fuca including food habits of the common inshore fishes: U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, Environmental Research Laboratories, 

NOAA Technical Memorandum ERL MESA-32.

35. Simenstad, C.A., Miller, B.S., Nyblade, C.F., Thornburgh, K., and Bledsoe, L.J., 1979, Food web relationships of northern 

Puget Sound and the Strait of Juan de Fuca—A synthesis of the available knowledge: National Oceanic and Atmospheric 

Administration/Marine Ecosystems Analysis Puget Sound Project, Prepared for Office of Environmental Engineering and 

Technology, United States Environmental Protection Agency, 334 p.

36. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Ninespine Stickleback 233

Ninespine Stickleback (Pungitius pungitius) 

(Linnaeus, 1758)

Family Gasterosteidae 

Note on taxonomy: Some authors recognize more species in this 

complex than others. North American Arctic populations have been 

considered a separate species P. occidentalis or to comprise two 

subspecies: P. p. pungitius, and P. p. occidentalis [1].

Colloquial Name: Iñupiat—Kakaliqauraq [2].

Ecological Role: Likely of considerable seasonal importance as a 

prey of fishes, sea birds, and marine mammals; occurs in brackish 

and marine waters near the coast.

Physical Description/Attributes: Slender, elongate body. Olive to pale brown on back, silvery or brassy yellow on sides and 

belly. Breeding colors vary with population; spawning males often have a great deal of black on sides, belly, and chins [3, 4]. 

Pure black males have been noted [3]. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 

2002, p. 334) [4]. Swim bladder: Present, without pneumatic duct [5]. Antifreeze glycoproteins in blood serum: Unknown.

Range: Along shores of U.S. Chukchi and Beaufort Seas [1]. Elsewhere in Alaska, along Bering Sea and western and northern 

Gulf of Alaska coasts inland to northeastern British Columbia. Worldwide, western Pacific Ocean in Seas of Japan and Okhotsk, 

and along Arctic shores except absent from Canadian high Arctic Archipelago, Greenland, and Iceland [1]. 

Relative Abundance: Common, although overall abundance is poorly described. Occasionally taken in large numbers along 

coasts in U.S. Chukchi and Beaufort Seas and in Canadian Beaufort Sea [8–12].

Ninespine Stickleback (Pungitius pungitius) 62 mm, 

northeastern Bering Sea (2007). Photograph by C.W. 

Mecklenburg, Point Stephens Research.

Geographic distribution of Ninespine Stickleback (Pungitius pungitius), within Arctic Outer Continental 

Shelf Planning Areas [6] based on review of published literature and specimens from historical and recent 

collections [1, 4, 7]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Ninespine Stickleback

Pungitius pungitius

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
234 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Nearshore, surface waters to depths of 110 m [4]. In ocean, spawning occurs nearshore, in estuary tide pools 

[13]. Larvae remain near spawning nests [14].

Habitats and Life History

Anadromous, riverine, and lacustrine forms [15]. 

Eggs—Size: 1.0–1.5 mm [16, 17]. Time to hatching: Unknown. Habitat: Benthic, in nests made of algae and 

plant material [3, 16]. 

Larvae—Size at hatching: 5.7 mm on average [17]. Size at juvenile transformation: About 15 mm [14]. Days to 

juvenile transformation: Perhaps 3 months [14]. Habitat: Benthic, remaining near nests [14]. 

Juveniles—Size range: About 15–38 mm [14, 18]. Habitat: In ocean, benthic, and midwaters, often under ice [9, 

19, 20]. 

Adults—Age and size at first maturity: 1–2 years [16, 21, 22] and at least 38 mm in Baltic Sea [18]. Maximum 

age: At least 2 years in North America [21]. Freshwater fish in Great Britain live to 3.5 years [23]. Maximum 

size: 9 cm TL [4]. Habitat: In ocean, benthic, and midwaters, often under ice [9, 19, 20]. 

Substrate—Rocks and sand for spawning [3]. 

Physical/chemical—Temperature:–1.9–20 °C [19, 21]. Salinity: Fresh to marine [15]. In Baltic Sea, prefers 

warmer, brackish waters for spawning [14].

Behavior

Little is known of the behaviors of these fish in Arctic waters.

Diel—Once at sea, makes offshore excursions, as much as 6 km off the coast in Beaufort Sea [22, 24]. 

Seasonal—Toward autumn some fish migrate to marine waters. Over wintering can occur in estuaries and river 

deltas [10, 11, 25, 26]. 

Reproductive—Spawning occurs at shallow depths in fresh and brackish waters [3, 16]. Nesting occurs in dense 

vegetation or in more exposed areas, such as in the crevices of boulder fields or under rocks. Males construct 

tunnel-shaped nests of plant material and lure females to them through a series of courtship behaviors [3, 16]. 

Males often mate with more than one female [15–17]. Females are batch spawners. Males protect fertilized eggs 

and larvae through and somewhat after hatching, often retrieving errant young and spitting them back into the 

nest [3]. 

Schooling—Juveniles school, adults may form small groups [27]. 

Feeding—Appears to occur during daylight hours [27].

Populations or Stocks

There have been no studies.

Benthic and reproductive distribution of Ninespine Stickleback (Pungitius pungitius).

3-6-2g_NinespineStickleback

Open ocean

Depth, in meters

500

400

300

200

100

0

Adults and juveniles 

Pelagic under ice

Pungitius pungitius

Ninespine Stickleback

Benthic distribution

Some fish to 110 m

Depth, in meters

Overall benthic depth range. 

Specific depth range of either 

juveniles and adults is known

Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Spawning and eggs

Reproductive distribution

Larvae

Data from outside U.S. Beaufort-Chukchi Seas 500

400

100

200

300

0
Ninespine Stickleback 235

Reproduction

Mode—Oviparous, separate sexes. 

Spawning season—Spring and summer [3, 16]. 

Fecundity—350–960 eggs, in batches of 20–80 eggs [28]

Food and Feeding

Food items—Zooplankton (for example, mysids and ostracods), adult and larval insects, mollusks, and fish eggs 

[8, 29, 30]. 

Trophic level—3.29 (standard error 0.40) [31].

Biological Interactions

Predators—In U.S. Beaufort and Chukchi Seas, other fishes including Arctic Cisco, Least Cisco, Dolly Varden, 

Fourhorn Sculpin, and Humpback Whitefish [8, 32], as well as belugas (off Point Barrow in May) [33], and 

ringed seals (over much of the year in northeastern Chukchi Sea [34]. Generally, Ninespine Stickleback are an 

important prey species for other fishes, birds, and mammals. 

Competitors—Likely such zooplanktivores as whitefishes, Pacific Herring, sculpins, and gadids.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (t

m=1–2; tmax=5; Fecundity=350) [31].

Traditional and Cultural Importance

Historically, Ninespine Stickleback were used as both human and dog food, although currently this species is not 

used [2, 16].

Commercial Fisheries

Currently, Ninespine Stickleback are not commercially harvested.

Potential Effects of Climate Change

Ninespine Stickleback are a predominantly boreal species with widespread presence along Arctic shores [1], 

which could be expected to increase in abundance and continue expanding to localities where suitable habitat can 

be found, as the climate warms.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment; 

(3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic 

movements; (6) population studies; (7) prey; and (8) predators. 
236 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Griswold, B.L., and Smith, L.L., Jr., 1972, Early survival and growth of the ninespine stickleback, Pungitius pungitius: 

Transactions of the American Fisheries Society, v. 101, no. 2, p. 350–352. [17]

Jarvela, L.E., and Thorsteinson, L.K., 1999, The epipelagic fish community of Beaufort Sea coastal waters, Alaska: Arctic, v. 52, 

no. 1, p. 80–94. [20]

Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British Columbia, 

Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p. [8]

McPhail, J.D., and Lindsey, C.C., 1970, Freshwater fishes of northwestern Canada and Alaska: Bulletin of the Fisheries 

Research Board of Canada Bulletin 173, 381 p. [16]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [4]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [1]

Bibliography

1. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

2. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

3. McKenzie, J.A., and Keenleyside, M.H.A., 1970, Reproductive behavior of ninespine sticklebacks (Pungitius pungitius

[L.]) in South Bay, Manitoulin Island, Ontario: Canadian Journal of Zoology, v. 48, no. 1, p. 55–61.

4. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

5. Moser, F., 1903, Comparative embryology of the swim-bladder: Journal of the Royal Microscopical Society, v. 25, p. 708.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

8. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.

9. Barton, L.H., 1978, Finfish resource surveys in Norton Sound and Kotzebue Sound: Alaska Department of Fish and Game, 

Commercial Fisheries Division, p. 75–313.

10. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.

11. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

12. Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., and Colonell, J.M., 1996, The 1995 fish and oceanography 

study in Mikkelsen Bay, Alaska: Anchorage, Alaska, Prepared by LGL Alaska Research Associates and Woodward-Clyde 

Consultant for BP Exploration (Alaska) Inc., 102 p. plus apps.
Ninespine Stickleback 237

13. Poulin, R., and Fitzgerald, G.J., 1989, Early life histories of three sympatric sticklebacks in a salt-marsh: Journal of Fish 

Biology, v. 34, no. 2, p. 207–221.

14. Sokolowska, E., and Krzysztof, E.S., 2002, Reproductive cycle and the related spatial and temporal distribution of the 

ninespine stickleback (Pungitius pungitius L.) in Puck Bay: Oceanologia, v. 44, no. 4, p. 475–490.

15. Richardson, E.S., Reist, J.D., and Minns, C.K., 2001, Life history characteristics of freshwater fishes occurring in the 

Northwest Territories and Nunavut, with major emphasis on lake habitat requirements: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences no. 2569, 156 p.

16. McPhail, J.D., and Lindsey, C.C., 1970, Freshwater fishes of northwestern Canada and Alaska: Bulletin of the Fisheries 

Research Board of Canada Bulletin 173, 381 p.

17. Griswold, B.L., and Smith, L.L., Jr., 1972, Early survival and growth of the ninespine stickleback, Pungitius pungitius: 

Transactions of the American Fisheries Society, v. 101, no. 2, p. 350–352.

18. Sokolowska, E., and Krzysztof, E.S., 2001, Fecundity of ninespine stickleback (Pungitius pungitius L., 1758) in the Puck 

Bay: Acta Ichtyologica et Piscatoria, v. 31, no. 1, p. 45–60.

19. Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86.

20. Jarvela, L.E., and Thorsteinson, L.K., 1999, The epipelagic fish community of Beaufort Sea coastal waters, Alaska: Arctic, 

v. 52, no. 1, p. 80–94.

21. Cameron, J.N., Kostoris, J., and Penhale, P.A., 1973, Preliminary energy budget of the ninespine stickleback (Pungitius 

pungitius) in an Arctic lake: Journal of the Fisheries Research Board of Canada, v. 30, no. 8, p. 1,179–1,189.

22. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 

1119, 90 p.

23. Jones, J.W., and Hynes, H.B.N., 1950, The age and growth of Gasterosteus aculeatus, Pygosteus pungitius and Spinachia 

vulgaris, as shown by their otoliths: Journal of Animal Ecology, v. 19, no. 1, p. 59–73.

24. Houghton, J.P., and Whitmus, C.J., 1988, Shallow neritic fish of the central Beaufort Sea: Seattle, Washington, Report 

prepared for Standard Alaska Production Company by Dames an Moore, 17298-002-20.

25. Schmidt, D.R., Griffiths, W.B., and Martin, L.R., 1987, Importance of anadromous fish overwintering habitat in the 

Sagavanirktok River Delta, Alaska: Anchorage, Alaska, Report by Ecological Research Associates for Standard Alaska 

Production Company and North Slope Borough, 71 p.

26. Schmidt, D.R., Griffiths, W.R., and Martin, L.R., 1989, Overwintering biology of anadromous fish in the Sagavanirktok 

River Delta, Alaska: Biological Papers of the University of Alaska, v. 24, p. 55–74.

27. Collette, B.B., and Klein-MacPhee, G., 2002, Bigelow and Schroeder’s fishes of the Gulf of Maine (3rd ed.): Washington, 

D.C., Smithsonian Institution Press, 882 p.

28. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

29. Hatfield, C.T., Stein, J.N., Falk, M.R., and Jessop, C.S., 1972, Fish resources of the Mackenzie River Valley: Winnepeg, 

Environment Canada Fisheries Service, Interim report 1, v. 1.

30. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

31. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
238 Alaska Arctic Marine Fish Ecology Catalog

32. Griffiths, W.B., Craig, P., Walder, G., and Mann, G., 1975, Fisheries investigations in a coastal region of the Beaufort Sea 

(Nunaluk Lagoon, Yukon), in Craig, P.C., ed., Fisheries investigations in a coastal region of the Beaufort Sea: Arctic Gas, 

Biological Report Series, v. 34, p. 1–129.

33. Lowry, L.F., Frost, K.J., and Seaman, G.A., 1986, Investigations of belukha whales in coastal waters of western and 

northern Alaska: Outer Continental Shelf Environmental Program Unit 612, Final Report, p. 359–392.

34. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.
Whitespotted Greenling 239

Whitespotted Greenling (Hexagrammos stelleri) 

Tilesius, 1810

Family Hexagrammidae 

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Likely of limited abundance and little ecological 

importance in the U.S. Chukchi and Beaufort Seas. 

Physical Description/Attributes: Brown to green tinged with 

orange and yellow. Body and head usually have small white spots 

and anal fin is yellow often with brown bars. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, 

p. 392) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in 

blood serum: Unknown.

Range: U.S. Chukchi Sea and Beaufort Seas, reported eastward to Simpson Cove (about 70°N, 145°W) [1–4]. Northern Sea of 

Japan, Hokkaido Island to Commander-Aleutian chain and northeast to Simpson Cove, Beaufort Sea, and Bering Sea to Puget 

Sound, Washington [4].

Relative Abundance: Uncommon in U.S. Chukchi Sea and western Beaufort Sea [1–4, 6, 7]. Abundant from Sea of Japan [8] to 

eastern Bering Sea [9] and the Gulf of Alaska [10, 11].

Whitespotted Greenling (Hexagrammos stelleri) 32 cm TL, 

southern Kuril Islands, 2003. Photograph by B.A. Sheiko, 

Russian Academy of Sciences.

Geographic distribution of Whitespotted Greenling (Hexagrammos stelleri), within Arctic Outer Continental 

Shelf Planning Areas [5] based on review of published literature and specimens from historical and recent 

collections [2, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Whitespotted Greenling

Hexagrammos stelleri

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
240 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: In northern waters, intertidal to 175 m, usually less than 100 m [1]; down to 275 m in Sea of Japan [2]. Juveniles, 

very shallow waters to 50 m [12–14]. Documented from 14–50 m in U.S. Chukchi Sea [3, 6]. Spawning occurs in shallow waters 

to at least 8 m [15]. In Gulf of Alaska, larvae are abundant in surface waters [16].

Habitats and Life History

Eggs—Size: 1.6–1.9 mm [17]; colored green, blue, violet, or grey [15]. Time to hatching: 30 days [14, 18]. 

Habitat: Demersal, nearshore [15]. 

Larvae—Size at hatching: 7.0–9.0 mm SL [11, 19]. Size at juvenile transformation: 30–40 mm FL [14, 19]. 

Days to juvenile transformation: About 1 year [14]. Habitat: Pelagic, near surface [16, 20]. 

Juveniles—Age and size: 3–15 cm FL [9, 19]. Habitat: Demersal, nearshore, among rocks, often in heavy algae 

and eelgrass cover [1, 14, 20–22]. 

Adults—Age and size at first maturity: A few mature at age–1 (15.0 cm FL) and most at age–2 (17.0–20.0 cm 

FL) [9, 12, 17]. Maximum age: At least 6 years, probably more [17]. Maximum size: About 48 cm TL [1], and 

1.6 kg [23]. Habitat: Demersal, on continental shelf among complex substrates and, occasionally, soft bottoms, 

often in heavy algae and eelgrass cover [1, 14, 20–22, 24, 25]. 

Substrate—Sand, silt, gravel, cobble, shell hash [3, 26, 27]. 

Physical/chemical—Temperature: –1.5–11.7 °C in southeastern Bering Sea, (mainly 4–7.2 °C) [28]. Salinity: 

Marine and estuarine [14, 24].

Behavior

Diel—Substrate-oriented; remains within 1.5 m of sea floor, occasionally rising to 5–6 m in midwaters [29]. 

Strictly diurnal (Sea of Okhotsk) [29] and active, moving into shallow waters to feed at night (eastern Bering 

Sea) [9]. More agonistic toward each other than toward other species [29]. 

Seasonal—From late spring to autumn, pelagic larvae transform to juveniles and recruit to sea floor [12–14]. 

Mature fish winter in deeper waters of continental shelf and return to nearshore in the summer, whereas juveniles 

remain nearshore year-round [12, 14]. 

Reproductive—Spawning occurs in shallow waters [15]. Females lay adhesive eggs on algae and highly 

territorial males guard them until they hatch. Males may guard as many as 7 egg masses (1,200–5,200 eggs each) 

from multiples females [14, 15]. 

Schooling—Usually solitary though small schools of as much as 1 dozen individuals have been observed [29]. 

Feeding—Both juveniles and adults move into shallow, often intertidal waters, to feed [9]. Often roots around in 

substrate for prey [29].

Benthic and reproductive distribution of Whitespotted Greenling (Hexagrammos stelleri).

Open ocean Shore

Depth, in meters

375

300

225

150

75

0

Some fish to 275 m

Hexagrammus stelleri

Whitespotted Greenling

Benthic distribution

Overall depth data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas 500

400

300

200

100

0

Open ocean Shore

Depth, in meters

Spawning and eggs Adults and juveniles

Adults

Reproductive distribution

Depths documented in U.S. Chukchi Sea Larvae
Whitespotted Greenling 241

Populations or Stocks

There have been no studies.

Reproduction

Mode—Partial (heterochronal) spawners. 

Spawning season—Spawning occurs from autumn through spring in Puget Sound [15, 30] and from June to 

October in the Gulf of Alaska, Bering Sea, and off Asia [17, 31–33]. 

Fecundity—1,070–12,397, in batches [15, 17, 19].

Food and Feeding

Food items—A very diverse array of benthic and midwater prey in Bering Sea and Gulf of Alaska. Pelagic 

larvae: Such zooplankters as copepods, amphipods, ostracods, crab larvae, and euphausiids. Benthic juveniles 

and adults: Crustaceans (for example, gammarid and caprillid amphipods, calanoid and harpacticoid copepods, 

shrimps, crabs, and barnacles), snails, bivalves and bivalve siphons, polychaetes, fish eggs, and fishes [20, 26, 

34–36]. 

Trophic level—3.33 (standard error 0.41) [37].

Biological Interactions

Predators—Arctic Terns, horned and tufted puffins, and river and sea otters in Gulf of Alaska and eastern Bering 

Sea [38–41]. Predation by seals is likely in the Chukchi Sea. 

Competitors—Likely other benthic feeders such as flounders, sculpins, and eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (Fecundity=6,679–38,408) [37].

Traditional and Cultural Importance 

None reported. Elsewhere, Whitespotted Greenling was an important food fish for Alaska Natives living in the 

Gulf of Alaska, Aleutian Islands, and eastern Bering Sea [42, 43]. 

Commercial Fisheries

Currently, Whitespotted Greenlings are not commercially harvested. There is a commercial fishery for this 

species in the northern Sea of Okhotsk [44]. 

Potential Effects of Climate Change

As with other predominantly boreal Pacific fish species, Whitespotted Greenling are expected to expand their 

range in Arctic waters as the climate warms.
242 Alaska Arctic Marine Fish Ecology Catalog

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs for this species in the study 

area include: (1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year 

benthic recruitment; (3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and 

ontogenetic movements; (6) population studies; (7) prey; (and 8) predators. The vulnerability of Whitespotted 

Greenling to climate change should be assessed. It is a suitable indicator of changes in the nearshore marine and, 

if incorporated into a regional monitoring design, key population parameters should be studied.

References Cited

Antonenko, D.V., and Vdovin, A.V., 2001, Seasonal distribution of the common greenling Hexagrammos stelleri

(Hexagrammidae) in Peter the Great Bay (the Sea of Japan): Journal of Ichthyology, v. 41, p. 524–528. [14]

DeMartini, E.E., 1970, Reproductive colorations, paternal behavior and egg masses of kelp greenling, Hexagrammos 

decagrammus, and whitespotted greenling, H. stelleri: Northwest Science, v. 60, no. 1, p. 32–35. [15]

Gomelyuk, V.E., 2000, Comparative analysis of everyday behavior and mode of the life of three species of Hexagrammos

(Hexagrammidae, Scorpaeniformes) in summer: Journal of Ichthyology, v. 40, no. 1, p. 74–85. [29]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Shestakov, A.V., and Nazarkin, M.V., 2006, On the biology of the white-spotted greenling Hexagrammos stelleri and the masked 

greenling H. octogrammus (Hexagrammidae) from Taui Bay of the Sea of Okhotsk: Journal of Ichthyology, v. 46, no. 8, p. 

677–680. [17]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

7. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

8. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

9. Tack, S.L., 1970, The summer distribution and standing stock of the fishes of Izembek Lagoon, Alaska: Fairbanks, 

University of Alaska, Master’s thesis.
Whitespotted Greenling 243

10. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

11. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

12. Blackburn, J.E., and Jackson, P.B., 1982, Seasonal composition and abundance of juvenile and adult marine finfish and crab 

species in the nearshore zone of Kodiak Island’s eastside during April 1978 through March 1979: Outer Continental Shelf 

Environmental Assessment Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 377–570.

13. Isakson, J.S., Houghton, J.P., Rogers, D.E., and Parker, S.S., 1986, Fish use of inshore habitats north of the Alaska 

Peninsula, June–September 1984 and June–July 1985—Final report: U.S. Department of the Interior, Minerals Management 

Service, MMS Alaska OCS Region, OCS EIS/EA, 380 p.

14. Antonenko, D.V., and Vdovin, A.V., 2001, Seasonal distribution of the common greenling Hexagrammos stelleri

(Hexagrammidae) in Peter the Great Bay (the Sea of Japan): Journal of Ichthyology, v. 41, p. 524–528.

15. DeMartini, E.E., 1970, Reproductive colorations, paternal behavior and egg masses of kelp greenling, Hexagrammos 

decagrammus, and whitespotted greenling, H. stelleri: Northwest Science, v. 60, no. 1, p. 32–35.

16. Jump, C.M., Duffy-Anderson, J.T., and Mier, K.L., 2008, Comparison of the Sameoto, Manta and MARMAP neustonic 

ichthyoplankton samplers in the Gulf of Alaska: Fisheries Research, v. 89, no. 3, p. 222–229.

17. Shestakov, A.V., and Nazarkin, M.V., 2006, On the biology of the white-spotted greenling Hexagrammos stelleri and the 

masked greenling H. octogrammus (Hexagrammidae) from Taui Bay of the Sea of Okhotsk: Journal of Ichthyology, v. 46, 

no. 8, p. 677–680.

18. Gorbunova, N.N., 1970, Spawning and development of greenlings (family Hexagrammidae), in Rass, T.S., ed., 

Greenlings—Taxonomy, biology and interoceanic transplantation: Academy of Sciences of the U.S.S.R., Transactions of the 

Institute of Oceanology, Israel Program for Scientific Translations, Jerusalem, p. 121–185.

19. Washington, B.B., Moser, H.G., Laroche, W.A., and Richards, W.J., 1984, Scorpaeniformes—Development, in Moser, 

H.G., and others, eds., Ontogeny and systematics of fishes: Gainsville, Florida, American Society of Ichthyologists and 

Herpetolgists, Special Publication 1, 28 p.

20. Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, Fisheries Research Institute, FRI-UW-7719.

21. McConnaughey, T., and McRoy, C.P., 1979, 13C label identifies eelgrass (Zostera marina) carbon in an Alaskan estuarine 

food web: Marine Biology, v. 53, no. 3, p. 263–269.

22. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

23. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p. 

24. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

25. Gomelyuk, V.E., and Shchetkov, S.Y., 1992, Distribution of fish in coastal biotopes of Peter the Great Bay, Sea of Japan, in 

the summer period: Biologiya Morya, nos. 3–4, p. 26–32.

26. Simenstad, C.A., Miller, B.S., Nyblade, C.F., Thornburgh, K., and Bledsoe, L.J., 1979, Food web relationships of northern 

Puget Sound and the Strait of Juan de Fuca—A synthesis of the available knowledge: National Oceanic and Atmospheric 

Administration/Marine Ecosystems Analysis Puget Sound Project, Prepared for Office of Environmental Engineering and 

Technology, United States Environmental Protection Agency, 334 p.

27. Laur, D.R., and Haldorson, L.J., 1996, Coastal habitat studies—The effect of the Exxon Valdez oil spill on shallow subtidal 

fishes in Prince William Sound: American Fisheries Society Symposium 18, p. 659–670.
244 Alaska Arctic Marine Fish Ecology Catalog

28. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

29. Gomelyuk, V.E., 2000, Comparative analysis of everyday behavior and mode of the life of three species of Hexagrammos

(Hexagrammidae, Scorpaeniformes) in summer: Journal of Ichthyology, v. 40, no. 1, p. 74–85.

30. Busby, M.S., Matarese, A.C., and Mier, K.L., 2000, Annual, seasonal, and diel composition of larval and juvenile fishes 

collected by dip-net in Clam Bay, Puget Sound, Washington, from 1985 to 1995: Seattle, Washington, U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-111, 48 p.

31. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224.

32. Rogers, B.J., Wangerin, M.E., and Rogers, D.E., 1983, Seasonal composition and food web relationships of marine 

organisms in the nearshore zone of Kodiak Island, including ichthyoplankton, zooplankton, and fish—A report of the fish 

component of the study: National Oceanic and Atmospheric Administration, Environmental assessment of the Alaskan 

Continental Shelf, v. 17, p. 541–685.

33. Kolpakov, N.V., 2005, Diversity and seasonal dynamics of ichthyocenosis of the Circumlittoral of Russkaya Bight 

(Northern Primor’s): Journal of Ichthyology, v. 45, no. 9, p. 744–753.

34. Fraser, C.M., 1946, Food of fishes: Transactions of the Royal Society of Canada, v. 45, sec. 5, p. 33–39.

35. Cross, J.N., Fresh, K., Miller, B.S., Simenstad, C.A., Steinfort, S.N., and Fegley, J.C., 1978, Nearshore fish and 

invertebrates assemblages along the Strait of Juan de Fuca including food habits of the common inshore fishes: U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, Environmental Research Laboratories, 

NOAA Technical Memorandum ERL MESA-32.

36. Rosenthal, R.J., 1980, Shallow water fish assemblages in the northeastern Gulf of Alaska—Habitat evaluation, species 

composition, abundance, spatial distribution and trophic interaction, in Bureau of Land Management, Environmental 

assessment of the Alaskan Continental Shelf, final reports of principal investigators: U.S. Department of Commerce, 

National Oceanic and Atmospheric Administration, p. 451–540.

37. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

38. Baird, P.A., and Moe, R.A., 1978, The breeding biology and feeding ecology of marine birds in the Sitkalidak Strait area, 

Kodiak Island, 1977: U.S. Fish and Wildlife Service, Office of Biological Services–Coastal Ecosystems, National Oceanic 

and Atmospheric Administration-Outer Continental Shelf Environmental Assessment Program, Contract 01-5-022-2538. 

39. Baird, P.A., and Gould, P., 1983, The breeding biology and feeding ecology of marine birds in the Gulf of Alaska: U.S. Fish 

and Wildlife Service, Denver Wildlife Research Center, Final Report, Outer Continental Shelf Environmental Assessment 

Program, Research Unit 341.

40. Larsen, D.N., 1984, Feeding habits of river otters in coastal southeastern Alaska: Journal of Wildlife Management, v. 48, no. 

4, p. 1,446–1,452.

41. Hatch, S.A., and Sanger, G.A., 1992, Puffins as samplers of juvenile pollock and other forage fish in the Gulf of Alaska: 

Marine Ecology Progress Series, v. 80, p. 1–14.

42. Turner, L.M., 1886, Contributions to the natural history of Alaska—Arctic series of publications, no. 2, Washington: U.S. 

Government Printing Office, 226 p.

43. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

44. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.
Okhotsk Hookear Sculpin 245

Okhotsk Hookear Sculpin (Artediellus ochotensis) 

Gilbert & Burke, 1912

Family Cottidae

Note: Except for geographic range data, all information is from areas 

outside of the study area. 

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Largely unknown. However, Okhotsk Hookear 

Sculpin are unlikely to represent a significant prey resource to higherlevel organisms.

Physical Description/Attributes: Head and upper body light reddish 

brown, light reddish spots on body and small reddish brown blotches 

on first dorsal fin. For specific diagnostic characteristics, see Fishes of 

Alaska (Mecklenburg and others, 2002, p. 494) [1]. Swim bladder: 

Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea north of Lisburne Peninsula (one record only) [2]. Worldwide, Japan Sea at Peter the Great Bay to 

Okhotsk Sea, Kuril Islands, and Commander Islands, to Gulf of Anadyr, Russia, western Bering Sea [2].

Relative Abundance: Rare in U.S. Chukchi Sea [2]. Elsewhere, occasional in Sea of Okhotsk. [5] and rare in Sea of Japan [6].

Geographic distribution of Okhotsk Hookear Sculpin (Artediellus ochotensis), within Arctic Outer Continental 

Shelf Planning Areas [3] based on review of published literature and specimens from historical and recent 

collections [2, 4]. 

Okhotsk Hookear Sculpin (Artediellus ochotensis). From 

Mecklenburg and others (2002, p. 494) citing others; 

drawing of fish from off western Kamchatka Peninsula, 

Russia.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Okhotsk Hookear Sculpin

Artediellus ochotensis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
246 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Benthic, at least 4–100 m [1] and perhaps to 913 m [5]. Typically less than 50 m [1].

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Likely benthic [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 10.2 cm TL [1]. 

Habitat: Unknown. 

Substrate—Unknown. 

Physical/chemical—Temperature: Unknown. Salinity: Marine [1].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes; oviparous [7]. 

Spawning season—Unknown. 

Fecundity—Unknown.

Food and Feeding

Food items—Unknown. 

Trophic level—3.33 (standard error 0.40) [8].

3-6-4g_OkhotshHookearSculpin

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Juveniles and adults

Some fish

may go to

913 m

Artediellus ochotensis

Okhotsh Hookear Sculpin

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Overall benthic depth range.

Specific depth range of either 

juveniles or adults is unknown 

Depth range of spawning is uncertain. 

Potential location of spawning and eggs 

is based on depth range of benthic individuals

Characteristic location of sculpin larvae

Data from outside U.S. Beaufort-Chukchi Seas

Benthic distribution and reproductive distribution of Okhotsh Hookear Sculpin (Artediellus ochotensis). 
Okhotsk Hookear Sculpin 247

Biological Interactions

Predators—In the mid-eastern U.S. Chukchi Sea, Artediellus sp. are occasionally eaten by ringed seals [9]. 

Competitors—Presumably a wide range of other zoobenthos feeders such as Arctic Cod, Walleye Pollock, other 

sculpins, poachers, and eelpouts.

1980 1990 2000 2010

Resilience

High, minimum population doubling time less than 15 months (Preliminary K or Fecundity) [8].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Okhotsk Hookear Sculpin are not commercially harvested.

Potential Effects of Climate Change

As with other boreal Pacific species, climate warming would be expected to increase its abundance in the 

U.S. Chukchi Sea and possibly expand its range into the U.S. Beaufort Sea.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs in the study area include: 

(1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment; 

(3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic 

movements; (6) population studies; (7) prey; and (8) predators.

References Cited

Dudnik, Y.I., and Dolganov, V.N., 1992, Distribution and abundance of fish on the continental slopes of the Sea of Okhotsk and 

of the Kuril Islands during the summer of 1989: Journal of Ichthyology, v. 32, no. 9, p. 58–76. [5] 

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]
248 Alaska Arctic Marine Fish Ecology Catalog

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resoure 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Dudnik, Y.I., and Dolganov, V.N., 1992, Distribution and abundance of fish on the continental slopes of the Sea of Okhotsk 

and of the Kuril Islands during the summer of 1989: Journal of Ichthyology, v. 32, no. 9, p. 58–76.

6. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

7. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

8. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

9. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.
Hamecon 249

Hamecon (Artediellus scaber) 

Knipowitsch, 1907

Family Cottidae

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Largely unknown. However, the Hamecon is 

unlikely to represent a significant prey resource to higher-level 

organisms, but it is an important subsistence resource in some 

Alaskan communities.

Physical Description/Attributes: Grayish brown with large 

blotches and bars; fins have orange bars. Males have a dark 

blotch on posterior part of first dorsal fin. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, 

p. 491) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in 

blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas. Elsewhere in Alaska, in eastern Bering Sea to south of St. Lawrence Island. 

Worldwide, in western Bering Sea south to Cape Navarin, Russia; through Arctic Ocean eastward to Somerset Island, Canada, 

and westward to Barents and Kara Seas [1, 2].

Relative Abundance: Common, although patchily distributed in U.S. Chukchi Sea at least as far north as 71°N. [5–8]. 

Common in westernmost Beaufort Sea, although abundance in rest of Alaskan Beaufort Sea is unknown as few have been taken 

in nearshore areas [5–7, 9, 10]. Elsewhere, common in Canadian Beaufort Sea at Herschel Island, Yukon Territory [11] and in 

Russian Chukchi Sea [7].

Hamecon (Artediellus scaber), 83 mm TL, western Chukchi 

Sea, 2004. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Hamecon (Artediellus scaber), within Arctic Outer Continental Shelf Planning 

Areas [3] based on review of published literature and specimens from historical and recent collections [2, 4].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Hamecon

Artediellus scaber

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
250 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: At depths of 4–159 m, mostly shallower than 55 m [2]. Elsewhere, overall depth range given for benthic 

individuals is 7 m [9] to 290 m [12], although depths greater than 159 m are likely in error [2].

Some fish to 290 m

Open ocean Shore

Depth, in meters

375

300

225

150

75

0

375

300

225

150

75

0

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Artediellus scaber

Hamecon

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning and eggs

Depths in U.S. Beaufort Sea

Reproductive distribution

Characteristic location of sculpin larvae

Maximum depth data from outside U.S. Beaufort-Chukchi Seas

No data for this species

Juveniles and adults

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Benthic and reproductive distribution of Hamecon (Artediellus scaber). 

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [13]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Young-of-the-year may recruit to 

nearshore waters at lengths of 2.0 cm TL [9]. Days to juvenile transformation: Unknown. Habitat: Pelagic [1]. 

Juveniles—Age and size: Unknown and 2.0–5.6 cm TL [9, 14]. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Females larger than 5.6 cm TL are mature (about 3 or 4 years) [14]. 

Maximum age: At least seven years [13, 14]. Maximum size: 11.4 cm TL [15]. Habitat: Benthic, in coastal 

waters [13]. 

Substrate—Over sand, mud, and around rocks [7, 13]. Larger individuals may be found in deeper parts of 

species’ depth range [9]. 

Physical/chemical—Temperature: -1.8–9.8 °C or more [4]. Salinity: Brackish and marine, primarily brackish 

[16]. In U.S. Chukchi Sea, documented as much as 32.41 parts per thousand [7]. Off Russia and in western 

Chukchi Sea, documented between 10 and 32.87 parts per thousand [7, 13], 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes; oviparous [17]. 

Spawning season—Reproduction appears to take place at least in the autumn [13]. 

Fecundity—Females produce between 50 and 100 eggs [14].
Hamecon 251

Food and Feeding

Food items—Benthic individuals eat a variety of benthic and epibenthic prey, most importantly polychaetes and 

gammarid amphipods, as well as mysids, cumaceans, euphausiids, hyperiid amphipods, and isopods [14]. Food 

habits of larvae unknown. 

Trophic level—3.5 (standard error 0.38) [18].

Biological Interactions

Predators—In the mid-eastern U.S. Chukchi Sea Artediellus sp. are occasionally eaten by ringed seals [19].

Competitors—Presumably a wide range of other zoobenthos feeders such as Arctic Cod, Walleye Pollock, other 

sculpins, poachers, and eelpouts.

1980 1990 2000 2010

Resilience

High, minimum population doubling time less than 15 months (Preliminary K or Fecundity) [18].

Traditional and Cultural Importance 

None reported

Commercial Fisheries

Currently, Hamecon are not commercially harvested.

Potential Effects of Climate Change

An essentially Arctic species, climate warming would be expected to contract this species’ range northward from 

the Bering Sea.

Areas for Future Research [B]

Little is known about the ecology and life history of this species in the study area. Research needs include: 

(1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment; 

(3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic 

movements; (6) population studies; (7) prey; and (8) predators.

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [13]
252 Alaska Arctic Marine Fish Ecology Catalog

Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p. [9] 

Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western Beaufort 

Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA Technical Report 

NMFS-SSRF-764, 22 p. [14]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [7]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. 

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Alverson, D.L., and Wilimovsky, N.J., 1966, Fishery investigations of the southeastern Chukchi Sea, in Wilimovsky, N.J., 

and Wolfe, J.N., eds., Environment of the Cape Thompson region, Alaska: Oak Ridge, Tennessee, U.S. Atomic Energy 

Commission, Division of Technical Information, p. 843–860.

6. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

7. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

8. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

9. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

10. Logerwell, E.A., National Marine Fisheries Service, written commun., 2010.

11. McAllister, D.E., 1963, Fishes of the 1960 “Salvelinus” program from western Arctic Canada: National Museum of Canada 

Bulletin, no. 185, p. 17–39.

12. Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic marine fishes of Canada: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, no. 2674, 112 p.

13. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]
Hamecon 253

14. Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western 

Beaufort Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA 

Technical Report NMFS-SSRF-764, 22 p.

15. Norcross, B.L., Holladay, B.A., and Gleason, C., 2011, Appendix 7—Length-frequency plots of non-key species, in A 

synthesis of diversity, distribution, abundance, age, size and diet of fishes in the lease sale 193 area of the northeastern 

Chukchi Sea—Final report: Prepared for ConocoPhillips Alaska Inc., Shell Exploration and Production Company, and 

Statoil USA E&P, Inc.

16. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

17. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

18. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

19. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.
254 Alaska Arctic Marine Fish Ecology Catalog

Antlered Sculpin (Enophrys diceraus) 

(Pallas, 1787)

Family Cottidae 

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Antlered Sculpin are common in the U.S. Chukchi 

Sea and uncommon in the U.S. Beaufort Sea. Their respective roles 

in marine ecosystem dynamics, although unknown, probably are 

more significant than many other species, and correspond to this 

abundance pattern.

Physical Description/Attributes: Greenish and reddish brown 

mottling on back and sides on cream or pale yellow background. 

Often with three or four vague dark bands and some marbling and spotting and fins are barred [1]. Spawning males have dark 

dorsal, pectoral, and caudal fins [2]. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, 

p. 472) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: In U.S. Chukchi Sea [1, 3] and western U.S. Beaufort Sea [4]. Elsewhere in Alaska, in Bering Sea and Commander 

(Russia)–Aleutian islands chain, southeastwards to Fort Tongass, Alaska. Worldwide, in Sea of Japan to Sea of Okhotsk [1]. 

Relative Abundance: Fairly common in the northeastern U.S. Chukchi Sea [7, 8]. Elsewhere, common from the Sea of Japan 

[9] and Sea of Okhotsk [10] to Bering Sea [11].

Antlered Sculpin (Enophrys diceraus), 101 mm TL, Chukchi 

Sea, 2007. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Antlered Sculpin (Enophrys diceraus), within Arctic Outer Continental Shelf 

Planning Areas [5] based on review of published literature and specimens from historical and recent 

collections [3, 6].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Antlered Sculpin

Enophrys diceraus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Antlered Sculpin 255

Depth Range: Documented at 26–50 m in U.S. Chukchi Sea [7, 12]. In other areas, primarily between 2–120 m deep [3, 13], but 

has been reported to 600 m in Peter the Great Bay, Sea of Japan [13]. Depth range of larvae and juveniles is unknown. Spawning 

occurs in nearshore waters as shallow as 2–15 m [2, 13].

Habitats and Life History

Eggs—Size: 1.7–2.2 mm in diameter [2, 13]. Time to hatching: Unknown. Habitat: Nearshore, on rocks [2, 13]. 

Larvae—Size at hatching: 6.5–6.9 mm. Larvae hatch in spring [2, 13]. Size at juvenile transformation: 1.3–2.2 

cm TL in Sea of Japan [2]. Days to juvenile transformation: Unknown. Habitat: Pelagic [2]. 

Juveniles—Age and size: Age unknown and 1.3–2.2 cm TL to 18–21 cm TL [2, 13]. Habitat: Benthic [2]. 

Adults—Age and size at first maturity: Most mature at 18–21 cm TL and males grow larger than females [2, 13]. 

Maximum age: Unknown. Maximum size: 38 cm (15.2 in) TL [13]. Habitat: Benthic. In Prince William Sound, 

mainly found along protected beachlines and in shallow embayments dominated by seaweed and seagrasses [14]. 

Large aggregations have also been found over soft sea floors in Sea of Japan [13]. 

Substrate—Shell hash, rocks, mixed gravel, sand, and mud [12, 13]. 

Physical/chemical—Temperature: Between -1.5 and 10 °C, but may prefer temperatures greater than 0 °C [2, 12, 

13, 15]. Salinity: Marine [2].

Behavior

Diel—Unknown. 

Seasonal—In Sea of Okhotsk it moves into deeper waters in winter [13]. 

Reproductive—In Sea of Japan, spawning occurs nearshore on rocks. In autumn, large mature males migrate 

into spawning areas first, followed by smaller mature males and, lastly, females. Juvenile fish do not inhabit the 

spawning grounds. Females lay eggs on rocks and these are guarded by adult fish, most likely males. Multiple 

females may lay their eggs in one nest and egg masses can be as large as 30 × 20 cm [2, 13]. Following spawning 

migrations occur offshore into deeper waters [13, 16]. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Benthic and reproductive distribution of Antlered Sculpin (Enophrys diceraus). 

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

500

400

300

200

100

0

Enophrys diceraus

Antlered Sculpin

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Characteristic location of sculpin larvae

Depth range of eggs is uncertain.

Potential location is based on 

depth range of spawning individuals

Overall depth data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Depths documented in U.S. Chukchi Sea

Some fish 

to 600 m

Juveniles and adults

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 
256 Alaska Arctic Marine Fish Ecology Catalog

Reproduction

Mode—Separate sexes; oviparous [17]. 

Spawning season—Unknown in U.S. Chukchi and Beaufort Seas. Elsewhere, spawning is from November to 

February [2, 13] and in April and May in the more northerly waters of Sea of Japan [18]. Fecundity: Females 

produce between 9,523 and 17,160 crimson, orange, or purple eggs [2]. 

Food and Feeding

Food items—Food habits of larvae unknown. Benthic individuals eat a wide range of benthic prey. Important 

food items are crustaceans (for example, gammarid amphipods, brachyuran, and hermit crabs), limpets, sea 

urchins, and brittle stars [14, 16, 18]. 

Trophic level—3.26 (standard error 0.43) [19].

Biological Interactions

Predation—Off Kamchatka Peninsula, Russia, both great and plain sculpins eat this species [20]. At Tee Harbor, 

southeastern Alaska, commonly eaten by river otters [21]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other 

sculpins, poachers, and eelpouts.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (Preliminary K or Fecundity) [19].

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Antlered Sculpin are not commercially harvested.

Potential Effects of Climate Change

A boreal Pacific species that appears to be common in the Gulf of Alaska, and Bering Sea, and common in the 

northeastern Chukchi Sea, Antlered Sculpin would be expected to increase in abundance in abundance of shelf 

areas of both seas.

Areas for Future Research [B]

Little is known about the ecology and life history of this species in Arctic Alaska. Research needs for this 

species include: (1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year 

benthic recruitment; (3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and 

ontogenetic movements; (6) population studies; (7) prey; and (8) predators.
Antlered Sculpin 257

References Cited

Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p. [7]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Panchenko, V.V., 2005, Seasonal distribution of antlered sculpin, Enophrys diceraus (Cottidae) in Peter the Great Bay, Sea of 

Japan: Russian Journal of Marine Biology, v. 31, p. 273–278. [13]

Sokolovskii, A.S., and Sokolovskaya, T.G., 1999, The early ontogenesis of Enophrys diceraus (Cottidae) from the waters of 

Peter the Great Bay (the Sea of Japan): Journal of Ichthyology, v. 39, no. 3, p. 268–272. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Sokolovskii, A.S., and Sokolovskaya, T.G., 1999, The early ontogenesis of Enophrys diceraus (Cottidae) from the waters of 

Peter the Great Bay (the Sea of Japan): Journal of Ichthyology, v. 39, no. 3, p. 268–272.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Rand, K.M., and Logerwell, E.A., 2011, The first demersal trawl survey of benthic fish and invertebrates in the Beaufort Sea 

since the late 1970s: Polar Biology, v. 34, no. 4, p. 475–488.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

8. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

9. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

10. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

11. Dragoo, D.E., 2006, Seabird, fish, marine mammals and oceanography coordinated investigations (SMMOCI) near Bluff, 

Norton Sound, Alaska, July 2002: U.S. Fish and Wildlife Service Report AMNWR 06/03, 35 p.
258 Alaska Arctic Marine Fish Ecology Catalog

12. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

13. Panchenko, V.V., 2005, Seasonal distribution of antlered sculpin, Enophrys diceraus (Cottidae) in Peter the Great Bay, Sea 

of Japan: Russian Journal of Marine Biology, v. 31, p. 273–278.

14. Rosenthal, R.J., 1980, Shallow water fish assemblages in the northeastern Gulf of Alaska—Habitat evaluation, species 

composition, abundance, spatial distribution and trophic interaction, in Bureau of Land Management, Environmental 

assessment of the Alaskan Continental Shelf, final reports of principal investigators: U.S. Department of Commerce, 

National Oceanic and Atmospheric Administration, p. 451–540.

15. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.

16. Kim, S.T., 2001, Winter migrations of shelf fish to the continental slope: Journal of Ichthyology, v. 41, no. 8, p. 564–574.

17. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

18. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

19. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

20. Tokranov, A.M., 1987, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok

(Cuvier) (Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114.

21. Mecklenburg, T.A., and Mecklenburg, C.W., Point Stephens Research, written commun., 2012.
Arctic Staghorn Sculpin 259

Arctic Staghorn Sculpin to Ribbed Sculpin

Arctic Staghorn Sculpin (Gymnocanthus tricuspis) 

(Reinhardt, 1830)

Family Cottidae 

Colloquial Name: No colloquial name within U.S. Chukchi and 

Beaufort Seas.

Ecological Role: Largely unknown. Current information about 

the precise occurrence in the U.S. Chukchi and Beaufort Seas is 

limited, although it appears to be one of the more common benthic 

fish species in the Arctic Ocean [1, 2]. Information regarding 

most of the biology and ecology of this species within the U.S. 

Chukchi and Beaufort Seas is not available. However, as one of 

the most common species in the Chukchi Sea, the Arctic Staghorn 

Sculpin is likely to represent a significant prey resource to higher 

level organisms. 

Physical Description/Attributes: Dark brown backs with dark blotches extending below lateral line and yellowish lower sides. 

Dark bars on dorsal and pectoral fins. First dorsal fin of males is blackish with scattered white spots. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 464) [3]. Swim bladder: Absent [3]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: Throughout U.S. Chukchi and Beaufort Seas. Elsewhere in Alaska, in eastern Bering Sea southwards to Saint Matthew 

Island (61°03’N, 173°40’W) [2]. Worldwide, circumpolar in distribution [3], in the Atlantic Ocean southward to Gulf of Maine 

(Eastport, Maine) and northern Norway [1]. 

Arctic Staghorn Sculpin (Gymnocanthus tricuspis) female, 

146 mm TL, Chukchi Sea, 2007. Photograph by B. Sheiko, 

Russian Academy of Sciences and C.W. Mecklenburg, Point 

Stephens Research.
260 Alaska Arctic Marine Fish Ecology Catalog

Relative Abundance: Common throughout U.S. Chukchi Sea at least as far north as 72°19’N, 175°57’W [2, 6, 7], and in the 

U.S. Beaufort Sea to at least Herschel Island, Yukon Territory [8, 9]. In the northern Bering Sea, common to just south of Bering 

Strait [10]. 

tac14-5222_fig3-7-1m_Gymnocanthus_tricuspis

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Staghorn Sculpin

Gymnocanthus tricuspis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Arctic Staghorn Sculpin (Gymnocanthus tricuspis) within Arctic Outer Continental 

Shelf Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [1, 5].
Arctic Staghorn Sculpin 261

Some fish to 451 m

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

500

400

300

200

100

0

Juveniles and adults

Gymnocanthus tricuspis

Arctic Staghorn Sculpin

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Depths common in U.S. Chukchi Sea

Reproductive distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Characteristic location of sculpin larvae

Overall depth data from outside U.S. Beaufort-Chukchi Seas

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Benthic and reproductive distribution of Arctic Staghorn Sculpin (Gymnocanthus tricuspis).

Depth Range: In northern U.S. Chukchi Sea common from 40–100 m. One juvenile taken in midwaters at 37 m to surface and 

one at 81 m to surface. A few juveniles and larvae documented from midwaters with maximum tow depths varying from 51 to 

29 m [11]. Elsewhere, 2–451 m, but uncommon less than 10–20 m [12–15].

Habitats and Life History

Eggs—Size: 1–2 mm [16, 17]. Time to hatching: Unknown. Habitat: Benthic [3, 17]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 18–30 mm [16, 17]. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [17]. 

Juveniles—Age: Unknown. Size: 1.8–9.0 cm SL [16]. Habitat: Benthic. 

Adults—Age and size at first maturity: Males mature at 6.0–7.0 cm SL (2–3 years) and females at about 9.0 cm 

SL (3–4 years). Females grow faster than males, reach a larger size, and are heavier at length [16]. Maximum 

age: In northeastern Chukchi Sea, females live to at least 9 years and males to 8 years [18]. Maximum size: 

29.9 cm TL [3]. Habitat: Benthic [17]; soft or low relief sea floors, sometimes in association with algal beds [1, 

16, 19]. 

Substrate—Documented on mud to gravel and rock in U.S. Chukchi Sea [2]. Elsewhere, sand, sand–mud, and 

gravel [1, 16, 20]. 

Physical/chemical—Temperature: -1.9–12.5 °C. Tolerant of a fairly broad range of temperatures but mainly 

about 0 °C or less [16, 21]. Salinity: Marine and brackish water as low as 8.1 ppt [16, 21].

Behavior

Diel—Partly buries itself in bottom substrates [17]. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.
262 Alaska Arctic Marine Fish Ecology Catalog

Reproduction

Mode—Separate sexes; oviparous [22]. Internal fertilization likely [16]. 

Spawning season—Autumn and early winter in Atlantic Ocean and White and Kara Seas [13, 16, 23]. 

Fecundity—In U.S. Chukchi Sea, 3,030–5,414 eggs [18]; Elsewhere, 2,060–3,512 eggs in an unidentified Arctic 

location [16]. 

Food and Feeding

Food items—In U.S. Chukchi Sea, prey varies with site, but crustaceans (for example, amphipods, cumaceans, 

and hermit crabs), polychaetes, clam siphons, echiurioids, gastropods, bivalves, and larvaceans are all 

important [24]. In general, benthic and water column invertebrates [24, 25]. 

Trophic level—3.46 standard error 0.49 [26]. 

Biological Interactions

Predators—In U.S. Chukchi Sea off Point Barrow, Alaska, predators include Polar Cod, Bering Flounder, and 

Estuarine Eelpout [18]. In Canadian Arctic, Black Guillemots, Thick-billed Murres, and bearded seals [27, 28]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (t

m=2–4; tmax=9; Fecundity=3,030) [26].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Arctic Staghorn Sculpin are not commercially harvested.

Potential Effects of Climate Change

As a predominantly Arctic species with a low temperature preference [16] a northward shift this species 

distribution is possible. 

Areas for Future Research [A]

Life history information is limited; however, distribution and abundance data suggest its potential as an 

indicator of changing conditions. Research needs include: (1) depth and location of pelagic larvae, (2) depth, 

location, and timing of young-of-the-year benthic recruitment, (3) preferred depth ranges for juveniles and 

adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population studies, (7) prey, and 

(8) predators. Arctic Staghorn Sculpin should be considered in vulnerability assessments of Arctic marine fish to 

climate change.
Arctic Staghorn Sculpin 263

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [16]

Ennis, G.P., 1968, Occurrences of the staghorn sculpin (Gymnocanthus tricuspis) in Newfoundland waters: Journal of the 

Fisheries Research Board of Canada, v. 25, no. 12, p. 2,729–2,731. [13]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [1]

Smith, R.L., Barber, W.E., Vallarino, M., Gillispie, J., and Ritchie, A., 1997a, Population biology of the Arctic staghorn sculpin 

in the northeastern Chukchi Sea: American Fisheries Society Symposium 19, p. 133–139. [18]

Bibliography

1. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

2. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Alverson, D.L., and Wilimovsky, N.J., 1966, Fishery investigations of the southeastern Chukchi Sea, in Wilimovsky, N.J., 

and Wolfe, J.N., eds., Environment of the Cape Thompson region, Alaska: Oak Ridge, Tennessee, U.S. Atomic Energy 

Commission, Division of Technical Information, p. 843–860.

7. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

8. McAllister, D.E., 1963, Fishes of the 1960 “Salvelinus” program from western Arctic Canada: National Museum of Canada 

Bulletin, no. 185, p. 17–39.

9. Logerwell, E.A., National Marine Fisheries Service, written commun., 2010.

10. Cui, X., Grebmeier, J.M., Cooper, L.W., Lovvorn, J.R., North, C.A., Seaver, W.L., and Kolts, J.M., 2009, Spatial 

distributions of groundfish in the northern Bering Sea in relation to environmental variation: Marine Ecology Progress 

Series, v. 393, p. 147–160.

11. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

12. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.
264 Alaska Arctic Marine Fish Ecology Catalog

13. Ennis, G.P., 1968, Occurrences of the staghorn sculpin (Gymnocanthus tricuspis) in Newfoundland waters: Journal of the 

Fisheries Research Board of Canada, v. 25, no. 12, p. 2,729–2,731.

14. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

15. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

16. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

17. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

18. Smith, R.L., Barber, W.E., Vallarino, M., Gillispie, J., and Ritchie, A., 1997, Population biology of the Arctic staghorn 

sculpin in the northeastern Chukchi Sea: American Fisheries Society Symposium 19, p. 133–139.

19. Bond, W.A., and Erickson, R.N., 1993, Fisheries investigations in coastal waters of Liverpool Bay, Northwest Territories: 

Winnipeg, Manitoba, Canada Department of Fisheries and Oceans, Central and Arctic Region, Canadian Manuscript Report 

of Fisheries and Aquatic Sciences No. 2204, 59 p.

20. McAllister, D.E., 1975, Ecology of the marine fishes of Arctic Canada, in Proceedings of the Circumpolar Conference on 

Northern Ecology, September 15–18, 1975: Ottawa, National Research Council of Canada, p. II-49–II-65.

21. Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86.

22. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

23. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

24. Coyle, K.O., Gillispie, J.A., Smith, R.L., and Barber, W.E. 1997, Food habits of four demersal Chukchi Sea fishes, in

Reynolds, J.B., ed., Symposium 19, Proceedings of the Fish Ecology in Arctic North America Symposium: American 

Fisheries Society, Fairbanks, Alaska, May 19–21, 1992, p. 310–318.

25. Atkinson, E.G., and Percy, J.A., 1992, Diet comparison among demersal marine fish from the Canadian Arctic: Polar 

Biology, v. 11, no. 8, p. 567–573.

26. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

27. Finley, K.J., and Evans, C.R., 1983, Summer diet of the bearded seal (Erignathus barbatus) in the Canadian High Arctic: 

Arctic, v. 36, no. 1, p. 82–89.

28. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.
Butterfly Sculpin 265

Butterfly Sculpin (Hemilepidotus papilio)

(Bean, 1880)

Family Cottidae 

Note: Except for physical description, relative abundance, and 

geographic range data, all information is from areas outside of the 

study area.

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Largely unknown. Current information about 

the occurrence of this fish is limited to the Chukchi Sea. The 

Butterfly Sculpin is unlikely to represent a significant prey resource 

to higher level organisms but may be an important consumer of 

benthic invertebrates.

Physical Description/Attributes: Reddish brown, yellow, and white, with metallic gold sheen and four more or less distinct 

blackish bars on upper sides and back extending onto dorsal fin [1]. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 431) [2]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea as far north as 69°55’N, 168°00’W [3]. Elsewhere in Alaska, eastern Bering Sea and along Aleutian 

Islands west to Buldir Island. Worldwide, from western Bering Sea to Sea of Okhotsk and Sea of Japan off Hokkaido, Japan [3].

Relative Abundance: Fairly common but not abundant in U.S. Chukchi Sea [6, 7]. Common in the Bering Sea [8], most 

common south of St. Matthew Island [6] and from Kamchatka Peninsula [9] to Sea of Okhotsk off eastern Sakhalin Island, 

Russia [10].

Butterfly Sculpin (Hemilepidotus papilio) 167 mm, Chukchi 

Sea, 2004. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Butterfly Sculpin (Hemilepidotus papilio) within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [5, 6]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Butterfly Sculpin

Hemilepidotus papilio

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
266 Alaska Arctic Marine Fish Ecology Catalog

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Likely pelagic. Larvae of the related Hemilepidotus hemilepidotus are found 

over continental shelf, slope, and in oceanic waters [12]. 

Juveniles—Age and size: Unknown. Habitat: Benthic, primarily on low relief sea floors [13]. Frequently in tide 

pools [2]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: About 42 cm TL 

[21]. Habitat: Benthic, primarily on low relief sea floors [13]. Frequently in tide pools [2]. 

Substrate—Mainly on stony-gravel bottoms [13]. 

Physical/chemical—Temperature: -1.8–11.8 °C [3, 5, 13, 14]; prefers 2.0 °C or less [9]. Salinity: Marine [13].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes; oviparous [15]. 

Spawning season—July in Bering Sea [16]. 

Fecundity—Unknown. 

Depth Range: From intertidal zone to 320 m [2], and typically less than 150 m [11]. Larvae are found over continental shelf, 

slope, and in oceanic waters [12].

3-7-2g_ButterflySculpin

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

500

400

300

200

100

0

Juveniles and adults

Hemilepidotus papilio

Butterfly Sculpin

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown 

Characteristic location of sculpin larvae

Data from outside U.S. Beaufort-Chukchi Seas

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Some fish to 320 m

Benthic and reproductive distribution of Butterfly Sculpin (Hemilepidotus papilio).
Butterfly Sculpin 267

Food and Feeding

Food items—In eastern Bering Sea, primarily benthic and epibenthic prey such as Tanner crabs, gammarid 

amphipods, young Walleye Pollock, and ostracods [17]. 

Feeding—Unknown. 

Trophic level—4.0 [18].

Biological Interactions

Predators—Unknown. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other 

sculpins, poachers, and eelpouts.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (Preliminary K or Fecundity) [19].

Traditional and Cultural Importance

None reported. Occasionally used as food by Alaska Natives on the Pribilof Islands. [20].

Commercial Fisheries

Currently, Butterfly Sculpin are not commercially harvested.

Potential Effects of Climate Change

As a predominantly Boreal species, Butterfly Sculpin would be expected to increase in abundance in the U.S. 

Chukchi Sea and to expand its range into the Beaufort Sea.

Areas for Future Research [B]

Little is known about the ecology and life history of this species in the U.S. Arctic marine environment. Research 

needs include: (1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year 

benthic recruitment, (3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and 

ontogenetic movements, (6) population studies, (7) prey, and (8) predators.
268 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [13]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [6]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [3]

Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237. [9]

Bibliography

1. Mecklenburg, C.W., and Mecklenburg, T.A., 2011, Butterfly Sculpin—Hemilepidotus papilio (Bean, 1880): Arctic Ocean 

Diversity Web site, accessed April 2011, at http://www.arcodiv.org/Fish/Hemilepidotus_papilio.html.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

7. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

8. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

9. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.

10. Kim, S.T., and Shepeleva, O.N., 2001, The structure of shelf ichthyocenoses of northeastern Sakhalin and Terpeniya Bay: 

Journal of Ichthyology, v. 41, no. 9, p. 711–722.

11. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

12. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.
Butterfly Sculpin 269

13. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

14. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

15. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

16. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224.

17. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

18. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.

19. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

20. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

21. Tokranov, A.M., 2014, Some features of biology of butterfly sculpin Melletes papilio (Cottidae) in the pre-Kamchatka 

waters of the Sea of Okhotsk: Journal of Ichthyology, no. 54, p. 558–565.
270 Alaska Arctic Marine Fish Ecology Catalog

Twohorn Sculpin (Icelus bicornis)

(Reinhardt, 1840)

Family Cottidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Twohorn Sculpin are uncommon in the U.S. Chukchi 

and Beaufort Seas. Their ecological role in benthic ecosystems, though 

not fully known, is thought to be minor with respect to predation and 

energy flows.

Physical Description/Attributes: Yellowish brown with brown spots. 

Nasal tubes and cirri are pale. For specific diagnostic characteristics, 

see Fishes of Alaska (Mecklenburg and others, 2002, p. 456) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood 

serum: Unknown.

Ranges: U.S. Beaufort Sea; presence in U.S. Chukchi Sea assumed from confirmed presence in adjacent waters of the 

East Siberian Sea, the slope north of the U.S. Chukchi Sea, and reported presence in western Chukchi Sea [2]. Worldwide: 

Predominantly Arctic; circumpolar [2].

Relative Abundance: Uncommon on outer continental shelf of U.S. Beaufort Sea [2]. 

Twohorn Sculpin (Icelus bicornis) 88 mm, Beaufort Sea, 

2011. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Twohorn Sculpin (Icelus bicornis) within Arctic Outer Continental Shelf Planning 

Areas [3] based on review of published literature and specimens from historical and recent collections [1, 

2, 4].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Twohorn Sculpin

Icelus bicornis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Twohorn Sculpin 271

Depth Range: Documented in U.S. Beaufort Sea at depths of 41–360 m [4]. At depths of 14–560 m, but mostly at 50–180 m on 

outer shelf [2, 4, 5]. Larvae of Icelus sp. are abundant in surface waters [6]. Juvenile fish live in shallow waters [5]. 

Habitats and Life History

Eggs—Size: Almost ripe eggs are 3.1 mm [5]. Time to hatching: Unknown. Habitat: Likely benthic based on life 

history patterns of many other members of this family [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Likely pelagic based on life history patterns of many other members of this 

family [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic, frequently in algal beds [5]. 

Adults—Age and size at first maturity: In U.S. Beaufort Sea, females grew larger than males and matured at 

about 4 years and 6 cm TL [7]. Maximum age: At least 5 years, based on a small sample from the Beaufort Sea 

[7]. Maximum size: 17 cm TL [4]. 8.8 cm TL in the U.S. Beaufort Sea [4]. Habitat: Benthic [1]. 

Substrate—Mud, shell hash, or cobble sea floors [5]. 

Physical/chemical—Temperature: -1.8–8.8 °C, preferably around 0 °C [5]. Salinity: Marine and brackish-water 

at salinities as low as 25.4 ppt [5].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes; oviparous [9]. 

Spawning season—August –October [5]. 

Fecundity—79–1,300 eggs [7, 8]. 

3-7-3g_TwohornSculpin

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

500

400

300

200

100

0

Juveniles

Icelus bicornus

Twohorn Sculpin

Benthic distribution

Adults

Some fish to 560 m

Open Ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Overall benthic depth range.

Specific depth range of either 

juveniles or adults is unknown

Overall data from outside U.S. Beaufort-Chukchi Seas

Depths documented in

U.S. Beaufort Sea

Characteristic

location of sculpin larvae

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Benthic and reproductive distribution of Twohorn Sculpin (Icelus bicornis).
272 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Food habits of larvae unknown. Major prey includes gammarid amphipods: polychaetes, 

euphausiids, hyperiid amphipods, isopods, shrimps, cumaceans, and sipunculids also are consumed [5, 7]. 

Trophic level—3.13 standard error 0.35 [10].

Biological Interactions

Predators—Unknown. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [10].

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Twohorn Sculpin are not commercially harvested.

Potential Effects of Climate Change

The Twohorn Sculpin is a predominantly Arctic species. The potential effects of climate change could be 

expected to shift the species distribution farther northwards. 

Areas for Future Research [B]

Little is known about the ecology and life history of this species from this region. 

Research needs include (1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-theyear benthic recruitment, (3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal 

and ontogenetic movements, (6) population studies, (7) prey, and (8) predators.
Twohorn Sculpin 273

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [5]

Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western Beaufort 

Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA Technical Report 

NMFS-SSRF-764, 22 p. [7]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

6. Jump, C.M., Duffy-Anderson, J.T., and Mier, K.L., 2008, Comparison of the Sameoto, Manta and MARMAP neustonic 

ichthyoplankton samplers in the Gulf of Alaska: Fisheries Research, v. 89, no. 3, p. 222–229.

7. Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western 

Beaufort Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA 

Technical Report NMFS-SSRF-764, 22 p.

8. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

9. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

10. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
274 Alaska Arctic Marine Fish Ecology Catalog

Spatulate Sculpin (Icelus spatula) 

Gilbert & Burke, 1912

Family Cottidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Largely unknown. Specific information regarding 

biology and ecology is sparse. This species is not a dominant 

species in the U.S. Chukchi and Beaufort Seas and is unlikely to be 

ecologically significant in regional food webs.

Physical Description/Attributes: Light brown backs with four 

or five indistinct dark saddles and white underside. For specific 

diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, p. 455) [1]. Swim bladder: Absent [1]. Antifreeze 

glycoproteins in blood serum: Unknown.

Ranges: U.S. Chukchi and Beaufort Seas [1–4]. Elsewhere in Alaska, in Bering Sea and Aleutian Islands west to Atka Island, 

and eastern Gulf of Alaska at Glacier Bay [1]. Worldwide, in Sea of Okhotsk, around Kuril Islands, Russia, and in western North 

Pacific off Kamchatka Peninsula to Arctic seas off Russia, and Canada to western Greenland and Labrador [1]. In Arctic Ocean, 

documented to 77°26’N [3] north of Siberia and to about 81°N in the Canadian High Arctic archipelago [5].

Relative Abundance: Common in U.S. Chukchi and Beaufort Seas [1–4, 7]. Elsewhere, common in Sea of Okhotsk, along Kuril 

Islands and Kamchatka Peninsula, Russia, and in Bering Sea [1, 3, 8, 9].

Spatulate Sculpin (Icelus spatula) 79 mm, western Chukchi 

Sea, 2004. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Spatulate Sculpin (Icelus spatula) within Arctic Outer Continental Shelf Planning 

Areas [6] based on review of published literature and specimens from historical and recent collections [1, 7]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Spatulate Sculpin

Icelus spatula

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Spatulate Sculpin 275

Depth Range: In U.S. Chukchi Sea, documented in 100 m or less [3]. Elsewhere, 12–930 m [10], mainly shallower than 200 m 

[11]. Larvae of Icelus sp. are abundant in surface waters in Gulf of Alaska [12]. In U.S. Chukchi Sea, one Icelus sp. juvenile was 

found in midwaters between 37 m and the surface [4].

Habitats and Life History

Eggs—Size: 1.4 mm. Pale yellow in color [11]. Time to hatching: Unknown. Habitat: Benthic [11]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [12]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [11]. 

Adults—Age and size at first maturity: Unknown. Maximum age: 10 years [11]. Maximum size: About 21 cm 

TL. Females grow larger than males [11]. Habitat: Benthic [11]. 

Substrate—Complex substrates (for example, rocks and sponges) and soft sea floor [3, 11, 13]. 

Physical/chemical—Temperature: -1.8–10.5 °C [3, 7]. Salinity: Primarily marine, documented as low as 

24.7 ppt [13]. 

Behavior

Diel—Unknown. 

Seasonal—In autumn, fish along the northern Kuril Islands migrate from the mid-continental shelf to the shelfslope break [11]. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Have been observed feeding in water column to 1 m above the bottom [14].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes; oviparous [15]. 

Spawning season—August –December [11, 16]. 

Fecundity—110–9,100 eggs, in a single batch [11, 16].

Benthic and reproductive distribution of Spatulate Sculpin (Icelus spatula).

Open ocean Shore

Depth, in meters 1000

800

600

400

200

0

1000

800

600

400

200

0

Juveniles and adults

Some fish to 930 m

Icelus spatula

Spatulate Sculpin

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Depth range documented 

in U.S. Chukchi Sea Characteristic location of 

sculpin larvae

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Overall depth data from outside U.S. Beaufort-Chukchi Seas

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 
276 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—In U.S. Chukchi and Beaufort Seas, epibenthic and benthic prey, such as mysids, gammarid 

amphipods, shrimps, and polychaetes [16]. In Russia, similar prey, as well as fishes and mollusks [11, 13]. 

Trophic level—3.92 standard error 0.67 [17].

Biological Interactions

Predators—Bearded seals and Thick-billed Murres in Canadian Arctic [18, 19], river otters in southeastern 

Alaska [20], and Great Sculpin off Kamchatka Peninsula [21]. 

Competitors—Presumably a wide range of other zoobenthos feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, flatfish, and other sculpins.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [17].

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Spatulate Sculpin are not commercially harvested.

Potential Effects of Climate Change

The Spatulate Sculpin is an Arctic Boreal species [10]. Although climate warming may not greatly affect the 

species current distribution, abundance patterns are likely to change and probably increase.

Areas for Future Research [B]

Little is known about the ecology and life history of this species from this region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators. 
Spatulate Sculpin 277

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [13]

Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western Beaufort 

Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA Technical Report 

NMFS-SSRF-764, 22 p. [16]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [3]

Tokranov, A.M., and Orlov, A.M., 2005, Some features of the biology of Icelus spatula (Cottidae) in Pacific waters off the 

northern Kuril Islands: Journal of Ichthyology, v. 45, no. 3, p. 229–236. [11]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Arctic Laboratories Limited, 1987, Beaufort ocean dumpsite characterization: Prepared for Environment Protection, 

Conservation and Protection, Environment Canada, Yellowknife, Northwest Territories, by Arctic Laboratories Limited and 

LGL Limited.

3. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

4. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

5. Hunter, J.G., Leach, S.T., McAllister, D.E., and Steigerwalk, M.B., 1984, A distributional atlas of records of the marine 

fishes of Arctic Canada in the National Museums of Canada and Arctic Biological Station: National Museums of Canada 

Syllogues, no. 52, 35 p.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

8. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

9. Orlov, A.M., 2003, Impact of eddies on spatial distributions of groundfishes along waters off the northern Kuril Islands, and 

southeastern Kamchatka (north Pacific Ocean): Indian Journal of Marine Sciences, v. 32, no. 2, p. 95–111.

10. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

11. Tokranov, A.M., and Orlov, A.M., 2005, Some features of the biology of Icelus spatula (Cottidae) in Pacific waters off the 

northern Kuril Islands: Journal of Ichthyology, v. 45, no. 3, p. 229–236.
278 Alaska Arctic Marine Fish Ecology Catalog

12. Jump, C.M., Duffy-Anderson, J.T., and Mier, K.L., 2008, Comparison of the Sameoto, Manta and MARMAP neustonic 

ichthyoplankton samplers in the Gulf of Alaska: Fisheries Research, v. 89, no. 3, p. 222–229.

13. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

14. Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86.

15. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

16. Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western 

Beaufort Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA 

Technical Report NMFS-SSRF-764, 22 p.

17. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

18. Bradstreet, M.S.W., 1980, Thick-billed murres and black guillemots in the Barrow Strait area, N.W.T., during spring—Diets 

and food availability along ice edges: Journal of Zoology, v. 58, no. 11, p. 2,120–2,140.

19. Finley, K.J., and Evans, C.R., 1983, Summer diet of the bearded seal (Erignathus barbatus) in the Canadian High Arctic: 

Arctic, v. 36, no. 1, p. 82–89.

20. Larsen, D.N., 1984, Feeding habits of river otters in coastal southeastern Alaska: Journal of Wildlife Management, v. 48, no. 

4, p. 1,446–1,452.

21. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.
Belligerent Sculpin 279

Belligerent Sculpin (Megalocottus platycephalus)

(Pallas, 1814)

Family Cottidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown but likely minimal, especially in the 

U.S. Beaufort Sea.

Physical Description/Attributes: Olive-brown or gray-brown 

with white or yellow markings. Lower sides have a series of light 

spots, the belly is white, the underside of head dark, and all fins 

are spotted and barred. For specific diagnostic characteristics, 

see Fishes of Alaska (Mecklenburg and others, 2002, p. 475) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood 

serum: Unknown.

Ranges: U.S. Chukchi Sea to western U.S. Beaufort Sea off Point Barrow, Alaska, at about 71°22’N, 156°19’W [2]. Elsewhere 

in Alaska, in eastern Bering Sea south to Herendeen Bay [1]. Worldwide, from Sea of Japan at Peter the Great Bay to Sea of 

Okhotsk and western Bering Sea [1, 3–5]. 

Relative Abundance: Common in eastern U.S. Chukchi Sea [7] and rare in U.S. Beaufort Sea [1, 2]. Common in Sea of Japan, 

Sea of Okhotsk, and parts of Bering Sea [3–5].

Belligerent Sculpin (Megalocottus platycephalus). 

Photograph by Doyne W. Kessler, National Marine Fisheries 

Service.

Geographic distribution of Belligerent Sculpin (Megalocottus platycephalus) within Arctic Outer Continental 

Shelf Planning Areas [6] based on review of published literature and specimens from historical and recent 

collections [1, 2, 7]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Belligerent Sculpin

Megalocottus platycephalus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
280 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Documented in U.S. Beaufort Sea at 0.3 m [2]. Juveniles and adults are found from shallow subtidal to 40 m 

[1, 7, 9].

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [9]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [9]. 

Juveniles—Age and size: Unknown. Habitat: Benthic, shallow nearshore waters [9]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 42 cm TL [5]. 

Habitat: Benthic, shallow coastal waters, estuaries, and occasionally lower parts of rivers [3, 9]. 

Substrate—Unknown. 

Physical/chemical—Temperature: 2.6–12.6 °C [7]. Salinity: Brackish to marine waters [9].

Behavior

Diel—Unknown. Seasonal—Along western Kamchatka Peninsula (Russia) coast, lives in coastal waters 

in winter and spring and ascends well up estuaries and short distances up rivers in summer and fall [5]. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes; oviparous [10]. 

Spawning season—Unknown. 

Fecundity—Unknown. 

Food and Feeding

Food items—Off western Kamchatka Peninsula, smaller fish feed on algae, and crustaceans such as copepods, 

cumaceans, amphipods, and mysids. Larger fish feed on similar prey as well as small fishes [11]. 

Trophic level—4.08 standard error 0.70 [12].

Benthic and reproductive distribution of Belligerent Sculpin (Megalocottus platycephalus).

3-7-5g_BelligerentSculpin

Open ocean Shore

Depth, in meters

Juveniles and adults

Megalocottus platycephalus

Belligerent Sculpin

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Spawning and eggs

Reproductive distribution

Both juveniles and adults 

occupy same depths 

Characteristic 

location of 

sculpin larvae

Documented in U.S. 

Beaufort Sea at 0.3 m

Data from outside U.S. Beaufort-Chukchi Seas

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Some fish to 30 m
Belligerent Sculpin 281

Biological Interactions

Predators—Unknown. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other 

sculpins, and eelpouts. 

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (assuming fecundity = 10–100) [12]. 

Traditional and Cultural Importance

Historically, an important subsistence species in northeastern Bering Sea and U.S. Chukchi Sea [13, 14].

Commercial Fisheries

Currently, Belligerent Sculpin are not commercially harvested.

Potential Effects of Climate Change

A predominantly Boreal Pacific species [2] that appears to be common in parts of the Bering Sea and uncommon 

in the U.S. Chukchi Sea. Belligerent Sculpin would be expected to increase in abundance in the U.S. Chukchi 

Sea and potentially the Beaufort Sea as waters warm.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [9]

Maksimenkov, V.V., and Tokranov, A.M., 1993, The diet of the pond smelt, Hypomesus olidus, in the Bol’shaya River Estuary 

(Western Kamchatka): Journal of Ichthyology, v. 33, no. 9, p. 11–21. [11]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]
282 Alaska Arctic Marine Fish Ecology Catalog

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

4. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of 

Japan): Journal of Ichthyology, v. 38, no. 1, p. 1–11.

5. Tokranov, A.M., 1994, Distribution and population of the Northern Far East belligerent sculpin, Megalocottus platycephalus 

platycephalus (Cottidae), in the Bol’shaya River Estuary (Western Kamchatka): Journal of Ichthyology, v. 34, no. 5, 

p. 149–153.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

8. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

9. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

10. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

11. Maksimenkov, V.V., and Tokranov, A.M., 1992, The feeding of the northern flathead sculpin in the Bolshaya River estuary 

(western Kamchatka): Biologiya Morya, v. 1–2, p. 34–42. [In Russian.]

12. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

13. Turner, L.M., 1886, Contributions to the natural history of Alaska—Arctic series of publications, no. 2, Washington: U.S. 

Government Printing Office, 226 p.

14. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.
Brightbelly Sculpin 283

Brightbelly Sculpin (Microcottus sellaris)

(Gilbert, 1896)

Family Cottidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown in benthic ecosystems over the 

continental shelf.

Physical Description/Attributes: Purplish body spotted and 

mottled with either black and white or with red or black. Body has 

two or more white bars and bright yellow belly. Pelvic fins have 

black or red spots along rays. For specific diagnostic characteristics, 

see Fishes of Alaska (Mecklenburg and others, 2002, p. 483) [1]. 

Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Ranges: Southern U.S. Chukchi Sea [2–4]. Elsewhere in Alaska, to southern Bering Sea and Commander–Aleutian Islands 

chain. Worldwide: From Gulf of Anadyr southward to the northern Sea of Japan, southern Okhotsk Sea, and Kuril Islands, 

Russia [1].

Relative Abundance: Rare in U.S. Chukchi Sea [2–4]; absent from the U.S Beaufort Sea. Common as far north as Norton 

Sound, eastern Bering Sea and southward to Sea of Japan and Kamchatka Peninsula, Russia [6–8].

Brightbelly Sculpin (Microcottus sellaris). Photograph by 

Doyne W. Kessler, National Marine Fisheries Service.

Geographic distribution of Brightbelly Sculpin (Microcottus sellaris) within Arctic Outer Continental Shelf 

Planning Areas [5] based on review of published literature and specimens from historical and recent collections 

[1, 3, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Brightbelly Sculpin

Microcottus sellaris

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
284 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Coastal, shallow waters down to 50 m, most often from shallow subtidal to about 40 m [1, 6, 9–11]. Some 

juveniles are found in nearshore waters [6].

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Likely benthic, based on other members of this 

family [1, 9]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Likely pelagic, based on other members of this family [1]. 

Juveniles—Age and size: Unknown. Habitat: Likely benthic, based on other members of this family [1, 9]. Some 

juveniles are found in nearshore eelgrass beds [6]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 17 cm TL [15]. 

Habitat: Likely benthic, based on other members of this family [1, 9]. 

Substrate—Unknown. 

Physical/chemical—Temperature: 2–10 °C in summer off western Kamchatka Peninsula, primarily 6–8 °C [7]. 

Salinity: Marine and estuarine fish [9].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—At night may move off bottom to feed [6].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes; oviparous [12]. 

Spawning season—Unknown. 

Fecundity—Unknown. 

Benthic and reproductive distribution of Brightbelly Sculpin (Microcottus sellaris).

3-7-6gBrightbellySculpin

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Juveniles and adults

Microcottus sellaris

Brightbelly Sculpin

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Overall benthic depth range.

Specific depth range of either 

juveniles or adults is unknown 

Characteristic location of sculpin larvae

Data from outside U.S. Beaufort-Chukchi Seas

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 
Brightbelly Sculpin 285

Food and Feeding

Food items—Juveniles living in eelgrass beds in eastern Bering Sea feed on polychaetes, amphipods and other 

crustaceans, snails, eelgrass, and fishes [6, 13]. 

Trophic level—3.4 standard error 0.53 [14].

Biological Interactions

Predators—Unknown. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [14].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Brightbelly Scuplin are not commercially harvested.

Potential Effects of Climate Change

As a predominantly Boreal Pacific species that appears to be common in the eastern Bering Sea and relatively 

rare in the Chukchi Sea, Brightbelly Sculpin could be expected to increase in abundance in the U.S. Chukchi Sea 

and perhaps to expand their distribution into the Beaufort Sea as waters warm.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [9]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]
286 Alaska Arctic Marine Fish Ecology Catalog

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Tack, S.L., 1970, The summer distribution and standing stock of the fishes of Izembek Lagoon, Alaska: Fairbanks, University of 

Alaska, Master’s thesis. [6]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Tack, S.L., 1970, The summer distribution and standing stock of the fishes of Izembek Lagoon, Alaska: Fairbanks, 

University of Alaska, Master’s thesis.

7. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.

8. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

9. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

10. Fedorov, V.V., 2000, Species composition, distribution and habitation depths of the northern Kuril Islands fish and fish like 

species, in Koteneva, B.N., ed., Commercial and biological studies of fishes in the Pacific waters of the Kuril Islands and 

adjacent areas of the Okhotsk and Bering Seas in 1992–1998: Moscow, VNIRO Publishing, collected papers, p. 7–41. [In 

Russian.]

11. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

12. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

13. McConnaughey, T., and McRoy, C.P., 1979, 13C label identifies eelgrass (Zostera marina) carbon in an Alaskan estuarine 

food web: Marine Biology, v. 53, no. 3, p. 263–269.

14. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

15. Tokranov, A.M., 2013, Specific features of distribution and size indices of four poorly studied species of sculpins (Cottidae) 

in the Okhotsk Sea waters off Kamchatka: Journal of Ichthyology, no. 53, p. 404–415.
Plain Sculpin 287

Plain Sculpin (Myoxocephalus jaok) 

(Cuvier, 1829)

Family Cottidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Plain Sculpin have been only rarely observed. 

There ecological significance is suspected to be of minor 

importance in nearshore and shelf habitats.

Physical Description/Attributes: Gray-brown body covered in 

small black spots. Lower sides have white spots. Fins are barred and 

spotted. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 481) [1]. Swim bladder: Absent 

[1]. Antifreeze glycoproteins in blood serum: Unknown.

Ranges: U.S. Chukchi Sea and western Beaufort Sea eastward to at least 71°N, 153°W [1, 2]. Elsewhere in Alaska, throughout 

Bering Sea to eastern Gulf of Alaska at Limestone Inlet [1] and Glacier Bay [3]. Worldwide, from Sea of Japan off North Korea 

to Sea of Okhotsk [1].

Relative Abundance: Uncommon in U.S. Chukchi Sea and western Beaufort Sea [2, 5]. Common throughout eastern Bering 

Sea as far north as Norton Sound and southwards to about Kodiak Island, Alaska, and westwards to Sakhalin Island (Russia), 

Sea of Okhotsk [6–10].

Plain Sculpin (Myoxocephalus jaok) 133 mm, northeastern 

Chukchi Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Plain Sculpin (Myoxocephalus jaok), within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [1, 2, 5].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Plain Sculpin

Myoxocephalus jaok

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
288 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Intertidal to 680 m, mainly less than depth of 50 m off North America [6] and deeper than 5 m in western Pacific 

Ocean [11]. Larvae live over coastal and continental shelf waters [12]. Juveniles are in shallow nearshore waters [11, 13]. 

Spawning has been noted at nearshore at 6–8 m and on mid-continental shelf at 80–130 m [14].

Habitats and Life History

Eggs—Size: 1.9–2.4 mm [14, 15]. Time to hatching: Unknown. Habitat: Benthic and adhesive [14]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 1.3–2.0 cm SL [13, 16]. Days to juvenile 

transformation: About two months in Peter the Great Bay [15]. Habitat: Pelagic [12]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1, 6]. 

Adults—Age and size at first maturity: Few males mature at 3 years and most by 6 years. Females mature later, 

some as early as 5 years and most by 8 years [14, 15]. Maximum age: At least 15 years in Sea of Japan and 

western Pacific Ocean. Females may live longer than males [14, 15]. Maximum size: 70 cm TL [17]. Females 

grow faster than males [14, 15]. Habitat: Benthic [1, 6]. 

Substrate—Sand and mud bottoms [1]. 

Physical/chemical—Temperature: -1.9–13.1 °C [2]. Off Kamchatka Peninsula, most common at 2–4 °C [17]. 

Salinity: Marine waters [18–20].

Behavior

Diel—Unknown. 

Seasonal—Young-of-the-year settle into shallow nearshore waters as early as May [13]. Some adults occupy 

shallower shelf waters in summer and retreat to continental slope during winter [17, 21]. 

Reproductive—Females lay eggs in shallow waters on plants and mussel clusters. Males guard nests until the 

ova hatch [14]. Schooling: Unknown. 

Feeding—Off Kamchatka Peninsula, fish feed most heavily during summer months [22].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes; oviparous [23]. 

Spawning season—December–March [14, 15, 24]. 

Fecundity—From 25,411–147,029 eggs, however it is unclear if all eggs are deposited at one time [14].

Benthic and reproductive distribution of Plain Sculpin (Myoxocephalus jaok).

Open ocean

Depth, in meters

500

400

300

200

100

0

500

400

300

200

100

0 Juveniles

Myoxocephalus jaok

Plain Sculpin

Benthic distribution

Adults

Some fish to 

680 m

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Specific depth range of juveniles uncertain.

Reported as shallow and near shore

Larvae

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Depth range of eggs is uncertain.

Potential location is based on depth

range of spawning individuals 
Plain Sculpin 289

Food and Feeding

Food items—A wide range of benthic and water column prey. Fishes and crabs often are most important prey, 

but in eastern Bering Sea, shrimps, hermit crabs, brittle stars, polychaetes, clams, squids, copepods, mysids, 

gammarid amphipods, and euphausiids also are consumed [22, 25, 26]. 

Trophic level—4.2 standard error 0.73 [27].

Biological Interactions

Predators—Mostly unknown. Great Sculpins [22] and probably river otters [28]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (assuming fecundity = 10–100) [27].

Traditional and Cultural Importance

Historically, this species was caught as a food fish in the northeastern Bering Sea and southeastern U.S. Chukchi 

Sea [29]. Like other nearshore species, they were taken mainly by elderly men and women who were not able to 

go great distances to procure other food [30].

Commercial Fisheries

Currently, Plain Sculpin are not commercially harvested.

Potential Effects of Climate Change

A predominantly Boreal Pacific species that appears to be common in the eastern Bering Sea but uncommon in 

the Chukchi Sea. Plain Sculpin could be expected to increase in abundance in the Chukchi Sea and perhaps to 

expand their range into the Beaufort Sea.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators. 
290 Alaska Arctic Marine Fish Ecology Catalog

References Cited 

Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p. [6]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Panchenko, V.V., 2001, Reproduction peculiarities of plain sculpin Myoxocephalus jaok in Peter the Great Bay, Sea of Japan: 

Russian Journal of Marine Biology, v. 27, p. 111–112. [14]

Panchenko, V.V., 2002, Age and growth of sculpins of the genus Myoxocephalus (Cottidae) in Peter the Great Bay (the Sea of 

Japan): Journal of Ichthyology, v. 42, no. 7, p. 516–522. [15]

Tokranov, A.M., 1987a, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok (Cuvier) 

(Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114. [22]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

3. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

6. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

7. Kim, S.T., and Shepeleva, O.N., 2001, The structure of shelf ichthyocenoses of northeastern Sakhalin and Terpeniya Bay: 

Journal of Ichthyology, v. 41, no. 9, p. 711–722.

8. Napazakov, V.V., and Chuchukalo, V.I., 2003, Diet and trophic status of mass species of sculpins (Cottidae) in the western 

part of the Bering Sea in autumn: Journal of Ichthyology, v. 43, no. 3, p. 236–244.

9. Hamazaki, T., Fair, L., Watson, L., and Brennan, E., 2005, Analyses of Bering Sea bottom-trawl surveys in Norton Sound—

Absence of regime shift effects on epifauna and demersal fish: ICES Journal of Marine Science, v. 62, no. 8, p. 1,597–1,602.

10. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

11. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

12. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.
Plain Sculpin 291

13. Blackburn, J.E., and Jackson, P.B., 1982, Seasonal composition and abundance of juvenile and adult marine finfish and crab 

species in the nearshore zone of Kodiak Island’s eastside during April 1978 through March 1979: Outer Continental Shelf 

Environmental Assessment Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 377–570.

14. Panchenko, V.V., 2001, Reproduction peculiarities of plain sculpin Myoxocephalus jaok in Peter the Great Bay, Sea of 

Japan: Russian Journal of Marine Biology, v. 27, p. 111–112.

15. Panchenko, V.V., 2002, Age and growth of sculpins of the genus Myoxocephalus (Cottidae) in Peter the Great Bay (the Sea 

of Japan): Journal of Ichthyology, v. 42, no. 7, p. 516–522.

 16. Burke Museum of Natural History and Culture, 2013, Ichthyology collections database—Record UW 49094: Burke 

Museum of Natural History database, accessed July 29, 2013, at http://biology.burke.washington.edu/ichthyology/database/

search.php.

17. Tokranov, A.M., 1992, Features of feeding of benthic predatory fishes of the west Kamchatka shelf: Journal of Ichthyology, 

v. 32, no. 7, p. 45–55.

18. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.

19. Gomelyuk, V.E., and Shchetkov, S.Y., 1992, Distribution of fish in coastal biotopes of Peter the Great Bay, Sea of Japan, in 

the summer period: Biologiya Morya, nos. 3–4, p. 26–32.

20. Cui, X., 2009, Climate-driven impacts of groundfish on food webs in the northern Bering Sea: Knoxville, University of 

Tennessee, Ph.D. Dissertation 162 p.

21. Kim, S.T., 2001, Winter migrations of shelf fish to the continental slope: Journal of Ichthyology, v. 41, no. 8, p. 564–574.

22. Tokranov, A.M., 1987a, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok

(Cuvier) (Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114.

23. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

24. Kolpakov, N.V., 2005, Diversity and seasonal dynamics of ichthyocenosis of the Circumlittoral of Russkaya Bight 

(Northern Primor’s): Journal of Ichthyology, v. 45, no. 9, p. 744–753.

25. Brodeur, R.D., and Livingston, P.A., 1988, Food habits and diet overlap of various Eastern Bering Sea fishes: National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS F/NWC-127, 76 p.

26. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

27. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

28. Heaton, T.H., and Grady, F., 2003, The Late Wisconsin vertebrate history of Prince of Wales Island, southeast Alaska, in

Shubert, B.W., Mead, J.I., and Graham, R.W., eds., Ice Age cave faunas of North America: Bloomington, Indiana University 

Press, p. 17–53.

29. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

30. Turner, L.M., 1886, Contributions to the natural history of Alaska—Arctic series of publications, no. 2: Washington, D.C., 

U.S. Government Printing Office, 226 p.
292 Alaska Arctic Marine Fish Ecology Catalog

Great Sculpin (Myoxocephalus polyacanthocephalus) 

(Pallas, 1814)

Family Cottidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area.

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Minor role. Great Sculpin are rare within the U.S. 

Chukchi Sea and absent from the Beaufort Sea. 

Physical Description/Attributes: Brown or gray with three darker 

saddles. Belly and lower sides have small white spots, and fins 

are barred and spotted. For specific diagnostic characteristics, see 

Fishes of Alaska (Mecklenburg and others, 2002, p. 482) [1]. Swim bladder absent [1]. Antifreeze glycoproteins in blood serum: 

Unknown. Unlike that of Arctic and shorthorn sculpins, blood serum is clear, rather than blue or green [2].

Ranges: One record from northeastern Chukchi Sea near Icy Cape at 70°20’N, 163°06’W [3]. Elsewhere in Alaska, throughout 

Bering Sea, Aleutian Islands and entire southeastern coast. Worldwide, in western Pacific Ocean, Gulf of Andyr to sea of 

Okhotsk Sea and eastern Sea of Japan and in eastern Pacific Ocean to southern Puget Sound, Washington [4–7]. 

Relative Abundance: Rare in eastern U.S. Chukchi Sea [3, 9, 10]. Elsewhere in Alaska, uncommon in northern most Bering 

Sea [10, 11] and common along Aleutian Islands, in eastern Bering Sea, and to at least southeastern Alaska [6, 7]. Worldwide, 

common in Sea of Japan eastwards along eastern Kuril Island and Kamchatka Peninsula, Russia [4, 5].

Great Sculpin (Myoxocephalus polyacanthocephalus) 

146 mm, Semidi Islands, western Gulf of Alaska, 2007. 

Photograph by C.W. Mecklenburg, Point Stephens Research.

Geographic distribution of Great Sculpin (Myoxocephalus polyacanthocephalus) within Arctic Outer 

Continental Shelf Planning Areas [8] based on review of published literature and specimens from historical 

and recent collections [1, 3, 9].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Great Sculpin

Myoxocephalus polyacanthocephalus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

Occurrence records

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Great Sculpin 293

Depth Range: Intertidal to 825 m, primarily 20–250 m [12–17]. Adults occasionally taken in subtidal nearshore waters [18]. 

Larvae are found over coastal and continental shelf waters [19]. Juveniles usually 20 m or less [7, 20]. Spawning occurs on 

lower continental shelf and upper continental slope [4, 21]. Found spawning on slope at depths of 415 m off Sakhalin Island, 

Russia [21].

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic and adhesive [4]. 

Larvae—Size at hatching: As small as 1.1 cm SL [22]. Size at juvenile transformation: Unknown. Days to 

juvenile transformation: Unknown. Habitat: Pelagic [19, 23]. 

Juveniles—Age and size: Unknown. Habitat: Benthic, neritic [20, 24], complex habitats such as rocks, cobbles, 

eelgrass, and other aquatic plants, and over soft sea floors [5–7, 20, 25, 26]. 

Adults—Age and size at first maturity: In Sea of Okhotsk, males first mature at 5 years (33–34 cm TL), fifty 

percent at 6 years (40 cm TL), and all at 8 years (50 cm TL). Females mature when older and larger, first at 7 

years (49–50 cm), fifty percent at 8 years (55 cm) and all at 9 years (65 cm TL). Maximum age: 16 years, off 

North America [27]. Maximum size: 76 cm TL [1] and more than 9 kg [12, 28]. Habitat: Benthic, neritic to 

oceanic [12–14, 16, 17]. Complex habitats, such as rocks, cobbles, eelgrass, and other aquatic plants as well as 

soft sea floors [5–7, 20, 25, 26]. 

Substrate—Rocks, cobbles, eelgrass, other aquatic plants and soft sea floors [5–7, 20, 25, 26]. 

Physical/chemical—Temperature: -0.5–12 °C, most abundant in 8 °C or less [4, 7, 12, 29, 30]. In Sea of 

Okhotsk spawning occurs between 0.8 and 1.9 °C [4, 21]. Salinity: Marine, or slightly estuarine waters [4, 7, 12, 

29, 30].

Behavior

Diel—Unknown. 

Seasonal—Beginning in May, young-of-the-year recruit from plankton to intertidal and shallow subtidal waters 

[23, 31]. In western Pacific Ocean and Sea of Japan, adults migrate into deeper waters in winter [15, 21]. 

Reproductive—Sea of Okhotsk fish spawn over cobble sea floors [4, 21]. 

Schooling—Little is known although adults have been noted forming dense aggregations in Sea of Okhotsk [4]. 

Feeding behavior—In western Pacific Ocean, feeding is heavier during summer [28, 29].

Populations or Stocks

There have been no studies.

Benthic and reproductive distribution of Great Sculpin (Myoxocephalus polyacanthocephalus).

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

Juveniles

Some fish to 825 m

Myoxocephalus polyacanthocephalus

Great Sculpin

Benthic distribution

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas 500

400

300

200

100

0

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Typical location of sculpin larvae

Adults
294 Alaska Arctic Marine Fish Ecology Catalog

Reproduction

Mode—Unknown. 

Spawning season—January–February in Sea of Okhotsk [4]. 

Fecundity—48,000–423,000 adhesive and yellowish-orange to red eggs [4, 21]. 

Food and Feeding

Food items—In eastern Bering Sea, a wide range of benthic and epibenthic prey. Fishes (for example, Capelin, 

Pacific Cod, Walleye Pollock, snailfishes, and flatfishes) and crabs are most important, followed by polychaetes, 

clams, octopi, mysids, gammarid and caprellid amphipods, isopods, euphausiids, shrimps, hermit crabs, brittle 

stars, and echiuroids [31–33]. Under certain circumstances, adults prey on eggs of conspecifics [28]. Smaller fish 

feed primarily on invertebrates and larger individuals on fishes [29, 34, 35]. 

Trophic level—4.1 standard error 0.67 [36].

Biological Interactions

Predators—In northeastern Pacific Ocean and Bering Sea, a wide range including other Great Sculpin, Pacific 

Cod, Pacific Halibut, Pink Salmon, Red Irish Lord, Rock Greenling, Walleye Pollock, Pigeon Guillemot, mink, 

harbor seals, and Steller sea lions [26, 35, 37–41]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins. 

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (t

m=6–8; tmax=9–13; K=0.08–0.21; Fecundity = 2,00–

10,000) [36].

Traditional and Cultural Importance

Historically, Great Sculpin were an important subsistence fish in Alaskan waters [42]. Currently, they are an 

unmarketed bycatch in Russian trawl fisheries [43].

Commercial Fisheries

Currently, Plain Sculpin are not commercially harvested.

Potential Effects of Climate Change

A Boreal Pacific species [9], Great Sculpin would be expected to increase in abundance in the U.S. Chukchi Sea 

and possibly expand their range into the Beaufort Sea.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, 4) spawning season, (5) seasonal and ontogenetic movements, 

(6) population studies, (7) prey, and (8) predators.
Great Sculpin 295

References Cited 

Abookire, A.A., Piatt, J.F., and Norcross, B.L., 2001, Juvenile groundfish habitat in Kachemak Bay, Alaska, during late summer: 

Alaska Fishery Research Bulletin, v. 8, no. 1, p. 45–56. [7]

Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p. 

[20]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Tokranov, A.M., 1987a, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok (Cuvier) 

(Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114. [28]

Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution in the Kronitsky Bay in relation to abiotic factors: Zoologicheskii 

Zhurnal, v. 63, p. 1,363–1,373. [4]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Low, P.S., and Bada, J.L., 1974, Bile pigments in the blood serum of fish from the family Cottidae: Comparative 

Biochemistry and Physiology, Part A—Physiology, v. 47, no. 2, p. 411–418.

3. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

4. Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution in the Kronitsky Bay in relation to abiotic factors: Zoologicheskii 

Zhurnal, v. 63, p. 1,363–1,373.

5. Gomelyuk, V.E., and Shchetkov, S.Y., 1992, Distribution of fish in coastal biotopes of Peter the Great Bay, Sea of Japan, in 

the summer period: Biologiya Morya, nos. 3–4, p. 26–32.

6. Murphy, M.L., Johnson, S.W., and Csepp, D.J., 2000, A comparison of fish assemblages in eelgrass and adjacent subtidal 

habitats near Craig, Alaska: Alaska Fishery Research Bulletin, v. 7, p. 11–21.

7. Abookire, A.A., Piatt, J.F., and Norcross, B.L., 2001, Juvenile groundfish habitat in Kachemak Bay, Alaska, during late 

summer: Alaska Fishery Research Bulletin, v. 8, no. 1, p. 45–56.

8. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

9. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

10. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

11. Dragoo, D.E., 2006, Seabird, fish, marine mammals and oceanography coordinated investigations (SMMOCI) near Bluff, 

Norton Sound, Alaska, July 2002: U.S. Fish and Wildlife Service Report AMNWR 06/03, 35 p.

12. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.

13. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.
296 Alaska Arctic Marine Fish Ecology Catalog

14. Panchenko, V.V., 1999, Distribution of sculpins of the genus Myoxocephalus in Peter the Great Bay, the Sea of Japan, in the 

summer period: Russian Journal of Marine Biology, v. 25, no. 172–173.

15. Panchenko, V.V., 2002, Age and growth of sculpins of the genus Myoxocephalus (Cottidae) in Peter the Great Bay (the Sea 

of Japan): Journal of Ichthyology, v. 42, no. 7, p. 516–522.

16. Zenger, H.H., Jr., 2004, Data report—2002 Aleutian Islands bottom trawl survey: U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS-AFSC-143, 247 p.

17. Orlov, A.M., 2005, Bottom trawl-caught fishes and some features of their vertical distribution in the Pacific waters off the 

north Kuril Islands and south-east Kamchatka, 1993–1999: Aqua, Journal of Ichthyology and Aquatic Biology of Fishes, 

v. 9, no. 4, p. 139–160.

18. Yakimishyn, J., University of British Columbia, written commun., 2010.

19. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.

20. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

21. Kim, S.T., 2001, Winter migrations of shelf fish to the continental slope: Journal of Ichthyology, v. 41, no. 8, p. 564–574.

22. Burke Museum of Natural History and Culture, 2013, Ichthyology collections database—Record UW 49094: Burke 

Museum of Natural History database, accessed July 29, 2013, at http://biology.burke.washington.edu/ichthyology/database/

search.php.

23. Blackburn, J.E., and Jackson, P.B., 1982, Seasonal composition and abundance of juvenile and adult marine finfish and crab 

species in the nearshore zone of Kodiak Island’s eastside during April 1978 through March 1979: Outer Continental Shelf 

Environmental Assessment Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 377–570.

24. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

25. Miller, B.S., Simenstad, C.A., Moulton, L.L., Fresh, K.L., Funk, F.C., Karp, W.A., and others, 1977, Puget Sound baseline 

program—Nearshore fish survey: University of Washington, Fishery Research Institute, FRI-UW-7710, 219 p.

26. Simenstad, C.A., Isakson, J.S., and Nakatani, R.E., 1977, Marine fish communities, in Merritt, M.L., and Fuller, R.G., eds., 

The environment of Amchitka Island, Alaska: National Technical Information Center, Energy Research and Development 

Administration TID-26712, p. 451–492.

27. Kimura, D.K., National Marine Fisheries Service, written commun., 2010.

28. Tokranov, A.M., 1987a, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok

(Cuvier) (Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114.

29. Tokranov, A.M., 1992, Features of feeding of benthic predatory fishes of the west Kamchatka shelf: Journal of Ichthyology, 

v. 32, no. 7, p. 45–55.

30. Orlov, A.M., and Maukhametov, I.N., 2003, Feeding characteristics of Greenland halibut Reinhardtius hippoglossoides

matsuurae and Kamchatka flounder Atheresthes evermanni in the northwestern part of the Pacific Ocean: Journal of 

Ichthyology, v. 43, no. 9, p. 789–801.

31. Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, Fisheries Research Institute, FRI-UW-7719.

32. Brodeur, R.D., and Livingston, P.A., 1988, Food habits and diet overlap of various Eastern Bering Sea fishes: National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS F/NWC-127, 76 p.

33. Yang, M.-S., 2003, Food habits of the important groundfishes in the Aleutian Islands in 1994 and 1997: National Marine 

Fisheries Service, AFSC Processed Report 2003-07, 233 p.
Great Sculpin 297

34. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

35. Napazakov, V.V., and Chuchukalo, V.I., 2003, Diet and trophic status of mass species of sculpins (Cottidae) in the western 

part of the Bering Sea in autumn: Journal of Ichthyology, v. 43, no. 3, p. 236–244.

36. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

37. Drent, R.H., 1965, Breeding biology of the pigeon guillemot, Cepphus columba: Ardea, v. 53, nos. 3–4, p. 99–159.

38. Hatler, D.F., 1976, The coastal mink of Vancouver Island, British Columbia: Vancouver, University of British Columbia, 

Ph.D. dissertation.

39. Hunter, M.A., 1979, Food resource partitioning among demersal fishes in the vicinity of Kodiak Island, Alaska: Seattle, 

University of Washington, Master’s thesis, 120 p.

40. Olesiuk, P.F., Bigg, M.A., Ellis, G.M., Crockford, S.J., and Wigen, R.J., 1990, An assessment of the feeding habits of 

harbour seals (Phoca vitulina) in the Strait of Georgia, British Columbia, based on scat analysis: Department of Fisheries 

and Oceans, Canadian Technical Report of Fisheries and Aquatic Sciences No. 1730.

41. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990.

42. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

43. Orlov, A.M., 2010, Russian Federal Research Institute of Fisheries and Oceanography, written commun., 2010.

44. Kontula, T., and Väinölä, R., 2003, Relationships of Palearctic and Nearctic ‘glacial relict’ Myoxocephalus sculpins from 

mitochondrial DNA data: Molecular Ecology, v. 12, no. 11, p. 3,179–3,184.
298 Alaska Arctic Marine Fish Ecology Catalog

Fourhorn Sculpin (Myoxocephalus quadricornis) 

(Linnaeus, 1758)

Family Cottidae 

Colloquial Names: Iñupiat: Kanaiok [1]; Kanayuq [2].

Ecological Role: This species is ubiquitous in the shallow 

nearshore waters of the U.S. Beaufort and Chukchi Seas. Found in 

the nearshore throughout the year, it is consumed by a wide variety 

of fishes, birds, and mammals.

Physical Description/Attributes: Dark gray on backs and sides 

and lighter on belly. Saddles on the back and bars on the fins. 

Elongate, tapering body with dorsally compressed head. For specific 

diagnostic characteristics, see Fishes of Alaska (Mecklenburg

 and others, 2002, p. 477) [3]. Swim bladder: Absent [3]. Antifreeze glycoproteins in blood serum: Unknown.

Ranges: In shallow waters of the U.S. Chukchi and Beaufort Seas. Elsewhere, Arctic; Circumpolar; southward to Gulf of 

Anadyr, St. Lawrence Island, and northern Bristol Bay, Bering Sea, and to northern Greenland and Baltic Sea in the Atlantic [4].

Relative Abundance: Common throughout shallow waters of U.S. Beaufort Sea [7–9] and Chukchi Sea to Point Lay [10, 

11]. Abundance south of Point Lay is unknown. Extremely abundant in shallow waters in Canadian Beaufort Sea eastwards to 

Tuktoyaktuk Peninsula, Canada [9, 12, 13].

Fourhorn Sculpin (Myoxocephalus quadricornis) 

Photograph by Doyne W. Kessler, National Marine 

Fisheries Service.

Geographic distribution of Fourhorn Sculpin (Myoxocephalus quadricornis) within Arctic Outer Continental Shelf 

Planning Areas [5] based on review of published literature and specimens from historical and recent collections 

[3, 4, 6].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Fourhorn Sculpin

Myoxocephalus quadricornis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Fourhorn Sculpin 299

Depth Range: Intertidal to 47 m, mainly 15–20 m or less [10, 14]. Elsewhere, in Baltic Sea common to at least 40 m [15]. 

Spawning occurs nearshore as shallow as 4 m [16–18]. In northeast Pacific Ocean, larvae are pelagic in coastal and continental 

shelf waters [19]. Juveniles are found in very shallow waters, often 1 m or less [16].

Habitats and Life History

Eggs—Size: 1.5–2.9 mm [15, 20, 21]. Color can be green, yellow, or brown according to type of prey eaten by 

females [22]. Time to hatching: Baltic Sea eggs hatched in 97 days at 1.5 °C and 55 days at 4.7 °C, and no eggs 

hatched at 10.5 °C [23]. Habitat: Benthic and adhesive [15, 20, 21]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 10–12 mm FL [16, 17, 24–28]. Days to 

juvenile transformation: Unknown. Habitat: Pelagic in coastal and continental shelf waters [19]. 

Juveniles—Age and size: 10–12 mm to 100–250 mm FL [12, 16, 17, 24–30]. Habitat: Same as adults. 

Adults—Age and size at first maturity: Varies with location. In Simpson Lagoon (near Colville River), males 

matured between 2–4 years and females between 4–6 years [16]. On Yukon Coast a few of both sexes were 

mature at 5 years, all males at 7 years, and all females at 10 years [29]. Generally, few fish mature at 100 mm FL

and all by 250 mm FL [12, 16, 17, 25, 29, 30]. Growth rates are variable between locations and females tend to 

grow larger at age than males [12, 18, 29]. A comparison of growth rates at two Baltic Sea locations indicated 

that growth was faster at higher ambient temperatures [31]. Maximum age: 16 years [17] and females may live 

longer [12, 18, 29]. Maximum size: 36.5 cm TL [3]. Habitat: Benthic habitats in protected, nearshore waters of 

river mouths, bay and inlets, and along open coasts [7, 29]. Occasionally, travels upstream in rivers and occupies 

lakes; reported in Mackenzie River as far upstream as 193 km, in Meade River as far upstream as 129 to 145 km 

[14] and in deep, cold Scandinavian lakes [32]. Eelgrass [33]. 

Substrate—Mud sea floors [23, 34, 35]. 

Physical/chemical—In U.S. Beaufort Sea, most abundant in colder, more saline marine waters [36, 37]. 

Temperature: In Alaskan and Canadian waters fish have been found at water temperatures between -2.0 and 

15 °C [14, 38]. Spawning occurs often at temperatures of less than 0 °C [16–18]. In Baltic Sea, upper lethal 

temperatures are between 17.5 to 25.5 °C [39]. Salinity: Fresh waters to 32 ppt [16, 27]. In a saline lake on 

Cornwallis Island, Canadian Arctic, fish survive in hypersaline conditions of as much as 36 ppt [40]. 

Benthic and reproductive distribution of Fourhorn Sculpin (Myoxocephalus quadricornis).

Open ocean Shore

Depth, in meters

125

100

75

50

25

0 Juveniles

Adults

Some fish to 47 m

Myoxocephalus quadricornus

Fourhorn Sculpin

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Larvae

125

100

75

50

25

0

Data from U.S. Beaufort-Chukchi Seas Data from U.S. Beaufort-Chukchi Seas

Depth range of eggs is uncertain.

Potential location is based on depth

range of spawning individuals 
300 Alaska Arctic Marine Fish Ecology Catalog

Behavior

Diel—Only limited coastal movements occur aside from seasonal, ice-related, inshore-offshore excursions [16, 

41, 42]. At night, small individuals may rise up into water column to feed [43]. 

Seasonal—In U.S. Beaufort Sea, fish over winter along shallow, bottom ice-free coastlines [12, 13, 16, 18, 25, 

44] and river deltas (for example, Colville and Sagavanirktok) [13, 45, 46]. In June and early July, after inshore 

bottom ice melts, migrations occur from just offshore into shallow waters [16, 28]. Young-of-the-year recruit to 

very shallow waters from June to August [16, 17, 24–28]. 

Reproductive—Females spawn annually [16] and may spawn more than one batch per season [30]. In Baltic 

Sea, males become territorial a few months before spawning and prepare nests by digging holes in mud sea 

floors. Females lay eggs in these depressions and males guard eggs until they hatch by lying on or next to eggs 

and fanning them with pectoral fins. Along with raising and opening fins, guarding males also make warning 

sounds and bite intruders. In aquaria, harassed males will move eggs to another location by pushing them, 

tucking them into the angle between head and pectoral fin, or taking eggs into their mouths [23, 34, 35]. 

Schooling—Probably does not school. 

Feeding—Smaller fish prey primarily on invertebrates, whereas larger ones add fishes to their diets [29]. 

Populations or Stocks: There have been no studies, although genetic research in the Canadian Arctic suggests 

little population structure in this region [47]. 

Reproduction

Mode—Though described as having external fertilization, a number of females containing fertilized eggs have 

been found [23]. 

Spawning season—January–March in U.S. Beaufort Sea, [12, 17, 18] and occasionally during summer 

months [30]. 

Fecundity—792–6,150 [15, 48] or as much as 18,000 eggs [49].

Food and Feeding

Food items—Benthic and epibenthic organisms. Dominant prey are crustaceans, primarily isopods and 

amphipods, and fishes (for example, Fourhorn Sculpin, Saffron Cod, and Arctic Cod). Other prey include 

polychaetes, insects, ascidians, fish eggs, clams, plant material, copepods, and shrimps [10, 11, 13, 17, 50]. 

Trophic level—3.7 standard error 0.59 [51].

Biological Interactions

Predators—A wide range of fishes, such as Arctic Cisco, Arctic Sculpin, Dolly Varden, Fourhorn Sculpin, 

Pacific Herring, Arctic Smelt, Saffron Cod, Shorthorn Sculpin, various eelpouts, as well as grebes, herons, loons, 

mergansers, Mew Gulls, Thick-billed Murres, ringed and probably bearded seals, and polar bears [10, 18, 27, 29, 

40, 52–55]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (t

m=3–5; tmax=14; Fecundity=792) [51]. 
Fourhorn Sculpin 301

Traditional and Cultural Importance

Generally, although they are widely taken as bycatch in a number of Canadian Arctic subsistence fisheries, 

Fourhorn Sculpin only are occasionally consumed [11, 56, 57]. In the past, particularly during times of poor 

fishing, they were more often targeted and eaten [1, 58]. Hudson Bay Inuits reportedly catch and consume in 

moderate numbers [59].

Commercial Fisheries

Currently, Fourhorn Sculpin are not commercially harvested.

Potential Effects of Climate Change

It is unclear what effects climate change will have on this species. This species appears to be somewhat 

eurythermal, implying that it would not be negatively affected by warming temperatures. However, its apparent 

rarity south of the Bering Strait suggests (1) physiological adaptation to colder water or (2) a competitive 

advantage for cold-temperate species, or both. If these assumptions are true, climate change effects might include 

a northerly shift in the distribution of the species and increased interactions with other marine species.

Areas for Future Research [A]

The Fourhorn Sculpin is one of the more dominant species in the U.S. Beaufort Sea nearshore , as it is relatively 

well-studied in coastal monitoring programs. It is a potential indicator species and, as such, information needs 

include: (1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic 

recruitment, (3) preferred depth ranges for juveniles and adults, (4) seasonal and ontogenetic movements, and (5) 

population dynamics.

References Cited

Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special reference 

to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 1119, 90 p. [17]

Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, Simpson 

Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649. [16]

Griffiths, W.B., Craig, P., Walder, G., and Mann, G., 1975, Fisheries investigations in a coastal region of the Beaufort Sea 

(Nunaluk Lagoon, Yukon), in Craig, P.C., ed., Fisheries investigations in a coastal region of the Beaufort Sea: Arctic Gas, 

Biological Report Series, v. 34, p. 1–129. [29]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Westin, L., 1968, The fertility of fourhorn sculpin, Myoxocephalus quadricornis (L.): Drottningholm, Sweden, Report of the 

Institute of Freshwater Research, v. 48, p. 67–70. [15]

Bibliography

1. Nelson, R.K., 1969, Hunters of the Northern Ice: Chicago, University of Chicago Press, 429 p.

2. Nelson, R.K., 1981, Harvest of the sea—Coastal subsistence in modern Wainwright: North Slope Borough, 112 p.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.
302 Alaska Arctic Marine Fish Ecology Catalog

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Bendock, T.N., 1977, Beaufort Sea estuarine fishery study: Alaska Department of Fish and Game Annual Report, Contract 

#03-5-022-69, p. 670–729.

8. Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., and Colonell, J.M., 1996, The 1995 fish and oceanography 

study in Mikkelsen Bay, Alaska: Anchorage, Alaska, Prepared by LGL Alaska Research Associates and Woodward-Clyde 

Consultant for BP Exploration (Alaska) Inc., 102 p. plus apps.

9. Wiswar, D.W., and Frugé, D.J., 2006, Fisheries investigations in western Camden Bay, Arctic National Wildlife Refuge, 

Alaska, 1987: Alaska Fisheries Data Series, U.S. Fish and Wildlife Service, 2006-10, 49 p.

10. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

11. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

12. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

13. Jones, M.L., and Den Beste, J., 1977, Tuft Point and adjacent coastal areas fisheries projects: Calgary, Alberta, Canada, 

Aquatic Environments, Ltd., 152 p.

14. McAllister, D.E., 1961, The origin and status of the deepwater sculpin, Myoxocephalus thompsoni, a Nearctic glacial relict: 

National Museum of Canada Bulletin, v. 172, p. 44–65.

15. Westin, L., 1968, The fertility of fourhorn sculpin, Myoxocephalus quadricornis (L.): Drottningholm, Sweden, Report of the 

Institute of Freshwater Research, v. 48, p. 67–70.

16. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

17. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, 

no. 1119, 90 p.

18. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.

19. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.

20. Washington, B.B., Moser, H.G., Laroche, W.A., and Richards, W.J., 1984, Scorpaeniformes—Development, in Moser, 

H.G., and others, eds., Ontogeny and systematics of fishes: Gainsville, Florida, American Society of Ichthyologists and 

Herpetolgists, Special Publication 1, 28 p.

21. Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p.

22. Westin, L., 1968, Environmentally determined variation in the roe colour in the fourhorn sculpin Myoxocephalus 

quadricornis (L.): Oikos, v. 19, no. 2, p. 403–407.

23. Westin, L., 1969, The mode of fertilization, parental behaviour and time of egg development in fourhorn sculpin, 

Myoxocephalus quadricornis (L): Drottningholm, Sweden, Report of the Institute of Freshwater Research, v. 49, 

p. 175–182.
Fourhorn Sculpin 303

24. Kendel, R.E., Johnston, R.A.C., Kozak, M.D., and Lobsiger, U., 1974, Movements, distribution, populations and food 

habits of fish in the western coastal Beaufort Sea: Department of the Environment, Canada, Interim Report of Beaufort Sea 

Project, Study B1, 64 p.

25. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.

26. Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbour, N.W.T.: Canadian Technical 

Report of Fisheries and Aquatic Sciences, no. 1218, 21 p.

27. Bond, W.A., and Erickson, R.N., 1987, Fishery data from Phillips Bay, Yukon, 1985: Winnipeg, Manitoba, Canadian Data 

Report of Fisheries and Aquatic Sciences, Central and Artic Region, Department of Fisheries and Oceans, no. 635, 47 p.

28. Bond, W.A., and Erickson, R.N., 1989, Summer studies of the nearshore fish community at Phillips Bay, Beaufort Sea coast, 

Yukon: Winnepeg, Manitoba, Canadian Technical Report of Fisheries and Aquatic Sciences, Central and Arctic Region, 

Department of Fisheries and Oceans, no. 1676, 102 p.

29. Griffiths, W.B., Craig, P., Walder, G., and Mann, G., 1975, Fisheries investigations in a coastal region of the Beaufort Sea 

(Nunaluk Lagoon, Yukon), in Craig, P.C., ed., Fisheries investigations in a coastal region of the Beaufort Sea: Arctic Gas, 

Biological Report Series, v. 34, p. 1–129.

30. Goldberg, S.R., Yasutake, W.T., and West, R.L., 1987, Summer spawning in the fourhorn sculpin, Myoxocephalus 

quadricornis, from Alaska: The Canadian Field-Naturalist, v. 101, p. 457.

31. Timola, O., and Luotonen, H., 1986, Age distribution and growth of the fourhorn sculpin, Myoxocephalus quadricornis, in 

the NE Bothnian Bay and the Archipelago Sea: Bothian Bay Report, v. 4, p. 25–32.

32. Savolainen, E., 1975, Distribution and food of Myoxocephalus quadricornis (L.) (Teleostei, Cottidae) in fresh waters of 

eastern Finland: Annales Zoologici Fennici, v. 12, p. 271–274.

33. Laur, D.R., and Haldorson, L.J., 1996, Coastal habitat studies—The effect of the Exxon Valdez oil spill on shallow subtidal 

fishes in Prince William Sound: American Fisheries Society Symposium 18, p. 659–670.

34. Westin, L., 1970, Observations on the nest digging of fourhorn sculpin Myoxocephalus quadricornis (L.): Drottningholm, 

Sweden, Report of the Institute of Freshwater Research, v. 50, p. 211–214.

35. Westin, L., 1971, Locomotory activity patterns of fourhorn sculpin, Myoxocephalus quadricornis (L.) (Pisces): 

Drottningholm, Sweden, Report of the Institute of Freshwater Research, v. 51, p.184–196.

36. Griffiths, W.B., Fechhelm, R.G., Gallaway, B.J., and Fissel, D.B., 1988, Yukon fish and effects of causeway: For Indian and 

Northern Affairs Canada, LGL Ltd. and Arctic Science Ltd., 124 p. 

37. Wiswar, D.W., West, R.L., and Winkleman, W.N., 1995, Fisheries investigation in Oruktalik Lagoon, Arctic National 

Wildlife Refuge, Alaska, 1986: U.S. Fish and Wildlife Service, Alaska Fisheries Technical Report, no. 30, 38 p.

38. Schmidt, D.R., Griffiths, W.R., and Martin, L.R., 1989, Overwintering biology of anadromous fish in the Sagavanirktok 

River Delta, Alaska: Biological Papers of the University of Alaska, v. 24, p. 55–74.

39. Westin, L., 1968, Lethal limits of temperature for fourhorn sculpin Myoxocephalus quadricornis (L.): Drottningholm, 

Sweden, Report of the Institute of Freshwater Research, v. 48, p. 71–76.

40. Dickman, M., 1995, An isolated population of fourhorn sculpin (Myoxocephalus quadricornis, family Cottidae) in a 

hypersaline high Arctic Canadian lake: Hydrobiologia, v. 312, no. 1, p. 27–35.

41. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

42. Underwood, T.J., Gordon, J.A., Millard, M.J., Thorpe, L.A., and Osborne, B.M., 1995, Characteristics of selected fish 

populations of Arctic National Wildlife Refuge coastal waters, final report, 1988–91: Fairbanks, Alaska, U.S. Fish and 

Wildlife Service, Fairbanks Fishery Resource Office, Alaska Fisheries Technical Report, no. 28.
304 Alaska Arctic Marine Fish Ecology Catalog

43. Hammar, J., Bergstrand, E., and Enderlein, O., 1996, Why do juvenile fourhorn sculpin, Triglopsis quadricornis, appear in 

the pelagic habitat at night?: Environmental Biology of Fishes, v. 46, no. 2, p. 185–195.

44. Houston, J., 1990, Status of the fourhorn sculpin, Myoxocephalus quadricornis, in Canada: The Canadian Field-Naturalist, 

v. 104, p. 7–13.

45. Craig, P.C., 1989a, An introduction to anadromous fishes in the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.

46. Schmidt, D.R., Griffiths, W.B., and Martin, L.R., 1987, Importance of anadromous fish overwintering habitat in the 

Sagavanirktok River Delta, Alaska: Anchorage, Alaska, Report by Ecological Research Associates for Standard Alaska 

Production Company and North Slope Borough, 71 p.

47. Kontula, T., and Väinölä, R., 2003, Relationships of Palearctic and Nearctic ‘glacial relict’ Myoxocephalus sculpins from 

mitochondrial DNA data: Molecular Ecology, v. 12, no. 11, p. 3,179–3,184.

48. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

49. Morrow, J.E., 1980, The freshwater fishes of Alaska: Anchorage, Alaska Northwest Publishing Company, 248 p.

50. Griffiths, W.B., 1983, Fish, in Truett, J.C., ed, Final reports of principal investigators—Volume 24—Environmental 

characterization and biological use of lagoons in the eastern Beaufort Sea: Anchorage, Alaska, National Oceanic and 

Atmospheric Administration, Office of Oceanography and Marine Assessment, Ocean Assessments Division, p. 325–366.

51. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed 

July 8, 2012, at http://www.fishbase.org.

52. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.

53. Bradstreet, M.S.W., 1980, Thick-billed murres and black guillemots in the Barrow Strait area, N.W.T., during spring—Diets 

and food availability along ice edges: Journal of Zoology, v. 58, no. 11, p. 2,120–2,140.

54. Finley, K.J., and Evans, C.R., 1983, Summer diet of the bearded seal (Erignathus barbatus) in the Canadian High Arctic: 

Arctic, v. 36, no. 1, p. 82–89.

55. Dyck, M.G., and Romberg, S., 2007, Observations of a wild polar bear (Urus maritimus) successfully fishing Arctic charr 

(Salvelinus alpinus) and fourhorn sculpin (Myoxocephalus quadricornis): Polar Biology, v. 30, p. 1,625–1,628.

56. George, J.C., and Nageak, B.P., 1986, Observations on the Colville River subsistence fishery at Nuiqsut, Alaska for the 

period 4 July–1 November 1984: Alaska Department of Wildlife Manangement, 70 p.

57. Moulton, L.L., Field, L.J., and Kovalsky, R., 1991, Predictability in the catch of Arctic cisco in the Colville River, Alaska: 

American Fisheries Society Symposium no. 11, p. 145–156.

58. MacGinitie, G.E., 1955, Distribution and ecology of the marine invertebrates of Point Barrow, Alaska: Smithsonian 

Miscellaneous Collections, v. 128, no. 9, p. 1–201.

59. Wein, E.E., Freeman, M.M.R., and Makus, J.C., 1996, Use of and preference for traditional foods among the Belcher Island 

Inuit: Arctic, v. 49, no. 3, p. 256–264.
Arctic Sculpin 305

Geographic distribution of Arctic Sculpin (Myoxocephalus scorpioides) within Arctic Outer Continental 

Shelf Planning Areas [8] based on review of published literature and specimens from historical and recent 

collections [4, 9]. 

Arctic Sculpin (Myoxocephalus scorpioides)

(Fabricius, 1780) 

Family Cottidae 

Colloquial Name: Iñupiat: Tivaqiq (Coronation Gulf) [1].

Ecological Role: Largely unknown. However, Arctic Sculpin are 

unlikely to represent a significant prey resource to other fish, birds, 

or mammals.

Physical Description/Attributes: Purplish-blue or blackish body 

with dark bands, white mottling and spots, and with barred and 

spotted fins. For specific diagnostic characteristics, see Fishes of 

Alaska (Mecklenburg and others, 2002, p. 479) [2]. Swim bladder: 

Absent [2]. Antifreeze glycoproteins in blood serum: Unknown. 

Blood serum is green to blue-green in color, likely caused by 

biliverdin build-up [3].

Range: Along coasts of U.S. Chukchi and Beaufort Seas. Elsewhere 

in Alaska, in Bering Sea south to Norton Sound and St. Lawrence Island. Worldwide, in western Bering Sea south to Gulf of 

Anadyr; from East Siberian Sea and along Arctic Canadian coast eastward to Greenland and Gulf of St. Lawrence in the Atlantic 

[4–7].

Relative Abundance: Uncommon in U.S. Chukchi and Beaufort Seas [4].Reported as common in Canadian Arctic Ocean but 

likely misidentified, confused with the more numerous M. scorpius [4].

Arctic Sculpin (Myoxocephalus scorpioides). Photograph 

by Andrey Vinnikov, Chukotka Branch, Pacific Research 

Institute of Fisheries and Oceanography.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Sculpin

Myoxocephalus scorpioides

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
306 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Shallow nearshore to depth of 40 m off Alaska [2, 21] and to 40 m elsewhere [10]. Larval Myoxocephalus spp. live 

in coastal and continental shelf waters [11].

Benthic and reproductive distribution of Arctic Sculpin (Myoxocephalus scorpioides).

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Likely benthic, based on other members of this 

family [2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [11]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [12]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 23.8 cm TL [2]. 

Habitat: Benthic [11, 12]. 

Substrate—Rock and algae [13]. 

Physical/chemical—Temperature: Below 0 to 8 °C or higher [14]. Salinity: Marine and estuarine waters as low 

as 6 ppt [7, 14–17].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Unknown. 

Spawning season—Autumn, in Russian Arctic Ocean [14]. 

Fecundity—Unknown. 

3-7-10g_ArcticSculpin

Open ocean Shore

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

Juveniles and adults

Myoxocephalus scorpioides

Arctic Sculpin

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Larvae

Data from outside U.S. Beaufort-Chukchi Seas Larvae data from outside U.S. Beaufort-Chukchi Seas

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 
Arctic Sculpin 307

Food and Feeding

Food items—Amphipods [12, 14]. 

Trophic level—3.39 standard error 0.52 [18].

Biological Interactions

Predators—Black Guillemots and possibly bearded seals in Canadian Arctic Ocean [19, 20]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins.

1980 1990 2000 2010

Resilience 

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [18].

Traditional and Cultural Importance

None reported. Not commonly caught and retained in subsistence fisheries [12].

Commercial Fisheries

Currently, Arctic Scuplin are not commercially harvested.

Potential Effects of Climate Change

The Arctic Sculpin is an endemic species that lives in shallow waters close to shore. A distributional shift to the 

north, in this case contraction from the northern Bering Sea, and increased species abundance in coastal waters 

are likely outcomes of climate warming.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.
308 Alaska Arctic Marine Fish Ecology Catalog

References Cited 

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [14]

Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton assemblages 

in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281. [11]

Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126. [12]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]

Bibliography

1. McAllister, D.E., Legendre, V., and Hunter, J.G., 1987, List of Inuktitut (Eskimo), French, English and scientific names of 

marine fishes of Arctic Canada: Rapport Manuscrit Canadien des Sciences Halieutiques et Aquatiques, no. 1932, 106 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Low, P.S., and Bada, J.L., 1974, Bile pigments in the blood serum of fish from the family Cottidae: Comparative 

Biochemistry and Physiology, Part A—Physiology, v. 47, no. 2, p. 411–418.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

6. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

7. Dunton, K.H., and Schonberg, S.V., 2000, The benthic faunal assemblage of the boulder patch kelp community, in Truett, 

J.C., and Johnson, S.R., eds., The natural history of an Arctic oil field—Development and the biota: San Diego, Academic 

Press, p. 371–397.

8. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

9. Mecklenburg, C.W., and Mecklenburg, T.A., Point Stephens Research, unpub. data, 2010.

10. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

11. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.

12. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.
Arctic Sculpin 309

13. McAllister, D.E., 1975, Ecology of the marine fishes of Arctic Canada, in Proceedings of the Circumpolar Conference on 

Northern Ecology, September 15–18, 1975: Ottawa, National Research Council of Canada, p. II-49–II-65.

14. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

15. Bond, W.A., and Erickson, R.N., 1993, Fisheries investigations in coastal waters of Liverpool Bay, Northwest Territories: 

Winnipeg, Manitoba, Canada Department of Fisheries and Oceans, Central and Arctic Region, Canadian Manuscript Report 

of Fisheries and Aquatic Sciences No. 2204, 59 p.

16. Morin, R., and Dodson, J.J., 1986, The ecology of fishes in James Bay, Hudson Bay and Hudson Strait, in Martini, I.P., ed., 

Canadian Inland Seas: Amsterdam, Elsevier Science Publishers B.V., p. 293–323.

17. Schneider-Vieira, F., Baker, R., and Lawrence, M., 1993, The estuaries of Hudson Bay—A case study of the physical and 

biological characteristics of selected sites: Report prepared for the Hudson Bay Program by North-South Consultants Inc., 

26 p.

18. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

19. Finley, K.J., and Evans, C.R., 1983, Summer diet of the bearded seal (Erignathus barbatus) in the Canadian High Arctic: 

Arctic, v. 36, no. 1, p. 82–89.

20. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.

21. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arctic-marine-fishes.
310 Alaska Arctic Marine Fish Ecology Catalog

Shorthorn Sculpin (Myoxocephalus scorpius) 

(Linnaeus, 1758)

Family Cottidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Shorthorn Sculpin is one of the most common 

benthic species in the shallower waters of the U.S. Chukchi Sea 

and is common in the Beaufort Sea. Although research is lacking, 

available information implies that this species is likely of moderate 

ecological importance as prey for some marine mammals and 

sea birds. 

Physical Description/Attributes: Body variably colored from 

black to greenish brown to pale olive with yellow and orange tinges and dark mottling. Belly is white, red (particularly in 

males), yellow, orange, or brown. Fins are brown, green, or yellow, with paler spots and bars [1–3]. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 478) [3]. Swim bladder: Absent [3]. Antifreeze 

glycoproteins in blood serum: Present [4, 5]. Blood serum is colored blue-green, likely due to biliverdin build-up [6].

Ranges: U.S. Chukchi and Beaufort Seas. Elsewhere in Alaska, southward through Bering Sea and Commander–Aleutian 

Islands chain and all southeastern Gulf of Alaska. Worldwide, circumpolar; southward to Gulf of Maine, Bay of Biscay, 

southwestern Kamchatka Peninsula, Russia, and northern British Columbia, Canada [7].

Relative Abundance: Common throughout much of the Chukchi Sea [10] (to at least 71°55’N, 175°18’W [11], and in U.S. 

Beaufort Sea [12–14]. Common in eastern Bering Sea and Norton Sound [15, 16] along with James Bay, Hudson Bay, and 

Hudson Strait, Canada [17].

Shorthorn Sculpin (Myoxocephalus scorpius) 237 mm, 

Bering Strait, 2006. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Shorthorn Sculpin

Myoxocephalus scorpius

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Shorthorn Sculpin (Myoxocephalus scorpius) within Arctic Outer Continental 

Shelf Planning Areas [8] based on review of published literature and specimens from historical and recent 

collections [3, 7, 9].
Shorthorn Sculpin 311

Depth Range: In Russian Chukchi Sea to at least 84 m [10, 11, 13]. Overall, intertidal (juveniles) [18] to 550 m, mainly less 

than 70 m [3]. Off Newfoundland, adults have been observed at less than 3 m [19]. Spawning occurs between 3 and 11 m [2, 

20]. Larval Myoxocephalus spp. live in coastal and continental shelf waters [21].

Habitats and Life History

Eggs—Size: 1.8–2.5 mm [22]. Time to hatching: 7–8 weeks at 3.3 °C [23] and less than 3 months at -1.5–0 °C

[2, 20]. Habitat: Benthic and adhesive, in crevices of boulders and other rocky formations [2, 20]. 

Larvae—Size at hatching: 6.0–8.8 mm [23]. Size at juvenile transformation: 1.0–2.0 cm TL, [24, 25]. Days to 

juvenile transformation: Unknown. Habitat: Pelagic, in coastal and continental shelf waters [21]. 

Juveniles—Age and size: 1–2 to 21–39 cm TL [24, 25]. Age unknown. Habitat: Young juveniles are pelagic and 

neritic [21], and older juveniles are benthic and neritic [3]. 

Adults—Age and size at first maturity: Off Newfoundland, a few of both sexes mature at 3 years (21 cm TL), 

all males at 6 years (30 cm TL); 60 percent of females at 6 years (34–35 cm TL), and all by 8 years (39 cm 

TL). Females grow larger than males and males mature at a slightly younger age and smaller size [2, 19, 20]. 

In European waters, males mature at 1 year (8–14 cm TL) and all females at 3 years (20 cm) [26]. In Russian 

Arctic Ocean, both sexes mature at 2–4 years and 15 cm TL (males) and 20 cm (females) long TL [25]. Egg 

development within female may take as long as 2.5 years in Newfoundland waters [20] and 1 year in European 

waters [27]. Generally, Newfoundland fish live much longer, grow more slowly, and mature later in life than 

do those across the Atlantic Ocean [19, 20, 28]. Based on Baltic Sea research, fish on both sides of the Atlantic 

Ocean appear to weigh about the same at length with females heavier at length than males [27]. Maximum age: 

15 years [2] (rarely more than 6 years in Europe) [26]. Maximum size: 60 cm TL [3] and 1.1 kg [29]. Habitat: 

Benthic, mainly shallow shelf waters [3, 25, 30]. 

Substrate—A wide range, including sand, mud, boulder, cobble, and algal beds [2, 11, 12, 31]. 

Physical/chemical—Temperature: -1.8–18.9 °C [11, 32], most commonly less than 0 °C [25]. Salinity: From 

5 ppt to full sea water [25, 32, 33].

Benthic and reproductive distribution of Shorthorn Sculpin (Myoxocephalus scorpius).

Open ocean

Depth, in meters

500

400

300

200

100

0

125

100

75

50

25

0

Juveniles

Myoxocephalus scorpius

Shorthorn Sculpin

Benthic distribution

Adults

Some fish 

to 550 m

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Specific depth range of juveniles

is uncertain. Reported as shallow 

and nearshore

Characteristic location of sculpin larvae

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Depths documented Russian Chukchi Sea
312 Alaska Arctic Marine Fish Ecology Catalog

Behavior

Diel—Off Newfoundland, juveniles move into shallower waters (less than 1–2 m) at night. [34]. 

Seasonal—Young-of-the-year recruit to shallow, often estuarine, waters [1] and some stay nearshore year round

[13]. Off Newfoundland, seasonal inshore-offshore movements of juveniles (shallow in summer and deeper 

depths in winter) have been reported [20]. 

Reproductive—Spawning occurs in shallow waters. Adhesive eggs are laid in crevices of boulders and other 

rocky formations. Nests are 1–4 m apart and may contain clutches of more than one female. Males aggressively 

guard eggs until hatching. After spawning, females move into deep waters (likely deeper than 100 m) and return 

in April and May after eggs hatch [2, 20]. 

Schooling—Unknown. 

Feeding—Semi-sedentary, lies in wait for prey [7]. When drifting ice darkens the waters, diets shifts from 

drabber benthic prey to brightly colored midwater amphipods [35]. Females feed less just before and after 

spawning [2].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Unknown. 

Spawning season—November and December in Newfoundland [2, 20] and as late as March in Europe [23]. 

Fecundity—4,205–60,976 (as few as 1,200 in Norway) [23], pink, orange, or red colored eggs [20, 23]. 

Food and Feeding

Food items—A diverse array of vertebrate and invertebrate prey. In Newfoundland, the Canadian Arctic Ocean, 

eastern and western Bering Sea, and Norway, fishes and crabs are often most important in the diet, although 

many other crustaceans (for example, isopods, amphipods, and shrimps), polychaetes, snails, sea urchins, brittle 

stars, and sea cucumbers also are consumed [2, 23, 35–37]. 

Trophic level—3.9 standard error 0.40 [38].

Biological Interactions

Predators—Ringed seals in U.S. Chukchi Sea [39]. Black Guillemots, Thick-billed Murres, and bearded seals in 

Canadian Arctic Ocean [40–42]. River otters in southeastern Alaska [43]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (t

m=2; Fecundity=2,742) [38]. 

Traditional and Cultural Importance 

Seem to have relatively little importance in the U.S. Chukchi and Beaufort Seas. Historically, this was a fairly 

important species to the Inuits of the Canadian Arctic where it is frequently used as food if other sources are in 

short supply and used as dog-feed [1].
Shorthorn Sculpin 313

Commercial Fisheries

Currently, Shorthorn Sculpin are not commercially harvested.

Potential Effects of Climate Change

The Shorthorn Sculpin reproduces and is common in Arctic and Boreal waters [7] and the possible effects 

of warming on its distribution are difficult to predict. This species has already reestablished the circumpolar 

distribution it had in pre-Wisconsinan glacial and Bering Land Bridge times [7].

Areas for Future Research [B]

Although there has been substantial life history work conducted off Newfoundland [2, 19, 20] and in European 

waters [23, 26, 27] little is known about the ecology and life history of this species in the study area. Research 

needs include: (1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year 

benthic recruitment, (3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and 

ontogenetic movements, (6) population studies, (7) prey, and (8) predators. The distribution of the species in 

regional habitats suggests its potential suitability as an indicator species for monitoring changes associated with 

global warming.

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [25]

Ennis, G.P., 1969, The biology of the shorthorn sculpin Myoxocephalus scorpius (L.) in Newfoundland waters: Saint John’s, 

Newfoundland, Memorial University, Master’s thesis. [2]

Ennis, G.P., 1970, Reproduction and associated behaviour in the shorthorn sculpin, Myoxocephalus scorpius in Newfoundland 

waters: Journal of the Fisheries Research Board of Canada, v. 27, no. 11, p. 2,037–2,045. [20]

Luksenburg, J.A., Pedersen, T., and Falk-Petersen, I.-B., 2004, Reproduction of the shorthorn sculpin Myoxocephalus scorpius

in northern Norway: Journal of Sea Research, v. 51, no. 2, p. 157–166. [23]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Bibliography

1. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.

2. Ennis, G.P., 1969, The biology of the shorthorn sculpin Myoxocephalus scorpius (L.) in Newfoundland waters: Saint John’s, 

Newfoundland, Memorial University, Master’s thesis.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Fletcher, G.L., Addison, R.F., Slaughter, D., and Hew, C.L., 1982, Antifreeze proteins in the Arctic shorthorn sculpin 

(Myoxocephalus scorpius): Arctic, v. 35, no. 2, p. 302–306.

5. Power, G., 1997, A review of fish ecology in Arctic North America: American Fisheries Society Symposium, no. 19, 

p. 13–39.
314 Alaska Arctic Marine Fish Ecology Catalog

6. Low, P.S., and Bada, J.L., 1974, Bile pigments in the blood serum of fish from the family Cottidae: Comparative 

Biochemistry and Physiology, Part A—Physiology, v. 47, no. 2, p. 411–418.

7. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

8. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

9. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

10. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

11. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

12. Alverson, D.L., and Wilimovsky, N.J., 1966, Fishery investigations of the southeastern Chukchi Sea, in Wilimovsky, N.J., 

and Wolfe, J.N., eds., Environment of the Cape Thompson region, Alaska: Oak Ridge, Tennessee, U.S. Atomic Energy 

Commission, Division of Technical Information, p. 843–860.

13. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

14. Logerwell, E.A., National Marine Fisheries Service, written commun., 2010.

15. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

16. Hamazaki, T., Fair, L., Watson, L., and Brennan, E., 2005, Analyses of Bering Sea bottom-trawl surveys in Norton Sound—

Absence of regime shift effects on epifauna and demersal fish: ICES Journal of Marine Science, v. 62, no. 8, p. 1,597–1,602.

17. Morin, R., and Dodson, J.J., 1986, The ecology of fishes in James Bay, Hudson Bay and Hudson Strait, in Martini, I.P., ed., 

Canadian Inland Seas: Amsterdam, Elsevier Science Publishers B.V., p. 293–323.

18. McAllister, D.E., 1975, Ecology of the marine fishes of Arctic Canada, in Proceedings of the Circumpolar Conference on 

Northern Ecology, September 15–18, 1975: Ottawa, National Research Council of Canada, p. II-49–II-65.

19. Ennis, G.P., 1970a, Age, growth, and sexual maturity of the shorthorn sculpin, Myoxocephalus scorpius, in Newfoundland 

waters: Journal of the Fisheries Research Board of Canada v. 27, no. 12, p. 2,155–2,158.

20. Ennis, G.P., 1970b, Reproduction and associated behaviour in the shorthorn sculpin, Myoxocephalus scorpius in 

Newfoundland waters: Journal of the Fisheries Research Board of Canada, v. 27, no. 11, p. 2,037–2,045.

21. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.

22. Washington, B.B., Moser, H.G., Laroche, W.A., and Richards, W.J., 1984, Scorpaeniformes—Development, in Moser, 

H.G., and others, eds., Ontogeny and systematics of fishes: Gainsville, Florida, American Society of Ichthyologists and 

Herpetolgists, Special Publication 1, 28 p.

23. Luksenburg, J.A., Pedersen, T., and Falk-Petersen, I.-B., 2004, Reproduction of the shorthorn sculpin Myoxocephalus 

scorpius in northern Norway: Journal of Sea Research, v. 51, no. 2, p. 157–166.

24. Burke Museum of Natural History and Culture, 2013, Ichthyology collections database—Record no. UW 043069: Burke 

Museum of Natural History and Culture, Icthyology Collections Database, accessed July 1, 2013, at http://biology.burke.

washington.edu/ichthyology/database.
Shorthorn Sculpin 315

25. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] 

26. Luksenburg, J.A., and Pedersen, T., 2002, Sexual and geographical variation in life history parameters of the shorthorn 

sculpin: Journal of Fish Biology, v. 61, no. 6, p. 1453–1464.

27. Lamp, V.F., 1966, Beitrage zur biologie der seeskorpione Myoxocephalus scorpius (L.) und Taurulus bubalis (Euphr.) in der 

Kieler Forde: Kieler Meeresforch, v. 22, p. 98–120.

28. Ennis, G.P., 1968, Occurrences of the staghorn sculpin (Gymnocanthus tricuspis) in Newfoundland waters: Journal of the 

Fisheries Research Board of Canada, v. 25, no. 12, p. 2,729–2,731.

29. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

30. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

31. Norderhaug, K.M., Christie, H., Fossa, J.H., and Fredriksen, S., 2005, Fish—Macrofauna interactions in a kelp (Laminaria 

hyperborea) forest: Journal of the Marine Biological Association of the UK, v. 85, p. 1,279–1,286.

32. Moring, J.R., 2001, Appearance and possible homing of two species of sculpins in Maine tidepools: Northeastern Naturalist, 

v. 8, no. 2, p. 207–218.

33. Schneider-Vieira, F., Baker, R., and Lawrence, M., 1993, The estuaries of Hudson Bay—A case study of the physical and 

biological characteristics of selected sites: Report prepared for the Hudson Bay Program by North-South Consultants Inc., 

26 p.

34. Methven, D.A., Haedrich, R.L., and Rose, G.A., 2001, The fish assemblage of a Newfoundland estuary—Diel, monthly and 

annual variation: Estuarine, Coastal and Shelf Science, v. 52, no. 6, p. 669–687.

35. Moore, I.A., and Moore, J.W., 1974, Food of shorthorn sculpin, Myoxocephalus scorpius, in the Cumberland Sound area of 

Baffin Island: Journal of the Fisheries Research Board of Canada, v. 31, no. 3, p. 355–359.

36. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p, 537–580.

37. Napazakov, V.V., and Chuchukalo, V.I., 2003, Diet and trophic status of mass species of sculpins (Cottidae) in the western 

part of the Bering Sea in autumn: Journal of Ichthyology, v. 43, no. 3, p. 236–244.

38. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

39. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: United States Atomic Energy Commission, Division of 

Technical Information, p. 877–924.

40. Bradstreet, M.S.W., 1980, Thick-billed murres and black guillemots in the Barrow Strait area, N.W.T., during spring—Diets 

and food availability along ice edges: Journal of Zoology, v. 58, no. 11, p. 2,120–2,140.

41. Finley, K.J., and Evans, C.R., 1983, Summer diet of the bearded seal (Erignathus barbatus) in the Canadian High Arctic: 

Arctic, v. 36, no. 1, p. 82–89.

42. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.

43. Larsen, D.N., 1984, Feeding habits of river otters in coastal southeastern Alaska: Journal of Wildlife Management, v. 48, 

no. 4, p. 1,446–1,452.
316 Alaska Arctic Marine Fish Ecology Catalog

Hairhead Sculpin (Trichocottus brashnikovi) 

Soldatov & Pavlenko, 1915 

Family Cottidae 

Note: Except for physical description, relative abundance, and 

geographic range data, all information is from areas outside of the 

study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Largely unknown. However, Hairhead Sculpin 

are unlikely to represent a significant prey resource to higher level 

organisms except for the possibility of greater importance in the 

southeastern Chukchi Sea. 

Physical Description/Attributes: Head and body mottled reddish 

and blackish brown with vague dark saddles; numerous white spots make pattern indistinct; white underside, and white band 

around caudal peduncle [1]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea [3] and U.S. Beaufort Sea eastward at least to 72°N, 155°W, northeast of Dease Inlet [1]. Elsewhere 

in Alaska, in northeastern Bering Sea from Seward Peninsula south to St Lawrence Island. Worldwide, from western Bering Sea 

off Cape Navarin, Russia, south to Sea of Okhotsk and northern Sea of Japan [2].

Relative Abundance: Common, although perhaps not abundant, in U.S. Chukchi Sea and present but uncommon in U.S. 

Beaufort Sea [1, 3].

Hairhead Sculpin (Trichocottus brashnikovi) 164 mm 

TL, northeastern Bering Sea, 2006. Photograph by C.W. 

Mecklenburg, Point Stephens Research.

Geographic distribution of Hairhead Sculpin (Trichocottus brashnikovi), within Arctic Outer Continental 

Shelf Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [1-3]. 

200

200

50

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50

YUKON

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TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Hairhead Sculpin

Trichocottus brashnikovi

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Hairhead Sculpin 317

Depth Range: 7–87 m [2]. Documented in U.S. Chukchi and Beaufort Seas at 13–60 m [1, 3].

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Likely benthic, based other members of this 

family [2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Likely pelagic, based other members of this family [2]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [2]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 22.5 cm [2]. 

Habitat: Benthic [2]. 

Substrate—Sandy bottoms [2]. 

Physical/chemical—Temperature: -1.2–6.5 °C [1]. Salinity: Marine [2].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Unknown. 

Spawning season—Unknown. 

Fecundity—Unknown.

Benthic and reproductive distribution of Hairhead Sculpin (Trichocottus brashnikovi).

3-7-12g_HairheadSculpin

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Adults and juveniles 

Some fish

to 87 m

Trichocottus brashnikori

Hairhead Sculpin

Benthic distribution

Open ocean No data for this species

Depths documented in U.S.

Beaufort-Chukchi Seas

Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Overall benthic depth range.

Specific depth range of either 

juveniles or adults is unknown 

Depth range of spawning is uncertain. 

Potential location of spawning and eggs

is based on depth range of benthic individuals

Overall depth data from outside U.S. Beaufort-Chukchi Seas
318 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Unknown. 

Trophic level—3.45 standard error 0.48 [5].

Biological Interactions

Predators—Unknown. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other 

sculpins, poachers, and eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [5].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Hairhead Sculpin are not commercially harvested.

Potential Effects of Climate Change

As a Boreal Pacific species, a northerly shift in distribution and increased abundance are probable expectations 

of warming. The effects of increased abundance on biological interactions, especially competition with similar 

species, presently cannot be assessed.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include:

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators. 
Hairhead Sculpin 319

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
320 Alaska Arctic Marine Fish Ecology Catalog

Bigeye Sculpin (Triglops nybelini) 

Jensen, 1944

Family Cottidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Largely unknown. However, the Bigeye Sculpin 

is an uncommon species and is unlikely to represent a significant 

prey resource to higher level organisms

Physical Description/Attributes: Dark brown back without dark 

saddles, cream-colored belly and broken or complete black lines 

along sides. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 426) [1]. Swim 

bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Ranges: U.S. Chukchi Sea slope and U.S. Beaufort Sea slope eastward to 149°W [2]. Practically circumpolar in Arctic 

Ocean [3].

Relative Abundance: Unknown, likely uncommon, especially at depths shallower than 200 m.

Bigeye Sculpin (Triglops nybelini) 105 mm, Chukchi 

Borderland, 2009. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Bigeye Sculpin (Triglops nybelini) within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [1-3]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Bigeye Sculpin

Triglops nybelini

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Bigeye Sculpin 321

Depth Range: 9–1,354 m, usually deeper than 200 m [1, 3]. As shallow as 101 m in U.S. Beaufort Sea [2].

Habitats and Life History

Eggs—Size: 3 mm, almost ripe eggs in Kara Sea [5]. Time to hatching: Unknown. Habitat: Likely benthic, based 

on other members of this family [1, 5]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Likely pelagic, based on other members of this family [1, 5]. [1]. 

Juveniles—Age and size: Unknown. Habitat: Likely benthic, based on other members of this family [1, 5]. 

Adults—Age and size at first maturity: Starting at 3–4 years, at latest by 7 years, and 14 cm in the Barents Sea. 

[6]. Maximum age: Unknown. Maximum size: 17 cm TL [1]. Habitat: Likely benthic, based on other members of 

this family [1, 5]. 

Substrate—Soft sea floors [5, 7]. 

Physical/chemical—Temperature: -1.8 to -0.1 °C, in Russian Arctic Ocean [5]. Salinity: Marine [5, 7], prefers 

less than 34 ppt [6].

Behavior

Diel—Unknown. Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Unknown. 

Spawning season—Probably summer and early autumn, off Europe and Asia [5, 6]. 

Fecundity—600–100 eggs [6]. Mature females contain more than one size class of eggs. In Kara Sea two 

females contained 307 and 600 eggs of the largest class size [5]. 

Benthic and reproductive distribution of Bigeye Sculpin (Triglops nybelini).

3-7-13g_BigeyeSculpin

Open ocean

Depth, in meters

Juveniles and adults

Triglops nybelini

Bigeye Sculpin

Benthic distribution

Overall benthic depth range. 

Depth documented in U.S.

Beaufort Sea.

Specific depth range of either 

juveniles or adults is unknown

Some fish to 1,354 m

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Characteristic location of sculpin larvae

1,500

1,200

900

600

300

0

1,500

1,200

900

600

300

0

 Data from outside 

U.S. Beaufort-Chukchi Seas 

Depth range of spawning is 

uncertain. Potential location of

spawning and eggs is based on

depth range of benthic individuals
322 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—In Russian Arctic Ocean, zooplanktors such as euphausiids and calanoid copepods [5] and fish [6]. 

Trophic level—3.29 standard error 0.41 [8].

Biological Interactions

Predators—In Hudson Bay, Thick-billed Murres and Black Guillemots [9]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [8].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Bigeye Sculpin are not commercially harvested.

Potential Effects of Climate Change

The Bigeye Sculpin is an endemic Arctic species [3]. A northerly shift in its distribution to colder and deeper 

offshore waters may be an outcome of climate warming.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.
Bigeye Sculpin 323

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [5]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

6. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

7. Arctic Laboratories Limited, 1987, Beaufort ocean dumpsite characterization: Prepared for Environment Protection, 

Conservation and Protection, Environment Canada, Yellowknife, Northwest Territories, by Arctic Laboratories Limited and 

LGL Limited.

8. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

9. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.
324 Alaska Arctic Marine Fish Ecology Catalog

Ribbed Sculpin (Triglops pingelii) 

(Reinhardt, 1837)

Family Cottidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Largely unknown. However, the Ribbed Sculpin 

is unlikely to represent a significant prey resource to higher level 

organisms and does not play a significant role in regional 

food webs.

Physical Description/Attributes: Olive brown-back with vague blackish saddles, white belly, and dark spots or streaks on 

sides below lateral line. In males the dark streaks form a broken or continuous stripe. For specific diagnostic characteristics, 

see, Fishes of Alaska (Mecklenburg and others, 2002, p. 428) [1] Swim bladder: Absent [1]. Antifreeze glycoproteins in blood 

serum: Unknown.

Range: U.S. Beaufort and Chukchi Seas. Elsewhere in Sea of Okhotsk, Bering Sea and Commander–Aleutian Islands chain 

and entire southeast coast [4]. Worldwide, circumpolar in Arctic Ocean; south to Sea of Japan, Puget Sound, Strait of Belle Isle, 

southern Greenland, Jan Mayen Island, Norway, and White Sea [2].

Relative Abundance: Common in U.S. Chukchi Sea [1, 4–8] but uncommon in U.S. Beaufort Sea. Elsewhere, common in 

Canadian Beaufort Sea off Herschel Island [9] and from off western Kamchatka Peninsula, Russia, along Aleutian Islands, in 

central and eastern Bering Sea, and eastward to Kodiak Island region [10–12].

Ribbed Sculpin (Triglops pingelii) 145 mm, U.S. Chukchi Sea. 

Photograph by C.W. Mecklenburg, Point Stephens Research.

Geographic distribution of Ribbed Sculpin (Triglops pingelii) within Arctic Outer Continental Shelf Planning 

Areas [3] based on review of published literature and specimens from historical and recent collections [1, 2, 4].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Ribbed Sculpin

Triglops pingelii

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Ribbed Sculpin 325

Depth Range: Documented in U.S. Chukchi Sea at 30–48 m [8]. Elsewhere, 4–930 m [13], typically less than 200 m [2].

Habitats and Life History

Eggs—Size: 2–3 mm, red in color [14–16]. Time to hatching: Unknown. Habitat: Benthic [14, 15]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 35 mm TL [6]. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [6]. 

Juveniles—Age and size: Unknown. Habitat: Benthic to epibenthic [11, 17]. 

Adults—Age and size at first maturity: 3–5 years at 7–9 cm [16]. Maximum age: 9 years [16]. Maximum size: 

20.2 cm SL, about 23.3 cm TL; reported but not verifiable to 24.2 cm [1]. Females grow larger and are heavier at 

length [18]. Habitat: Benthic to epibenthic [11, 17]. 

Substrate—Soft sea floors and rocky bottoms [8, 17, 19]. 

Physical/chemical—Temperature: -1.5–10.5 °C [1, 8, 17, 19]. Salinity: Mainly marine [8, 15], also taken in 

brackish waters [20].

Behavior

Diel—Juveniles and adults have been captured in midwaters at night [17]. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Fish have been observed feeding as much as 1 m above the bottom [19].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Unknown. 

Spawning season—In autumn off Kodiak Island, Alaska [21, 22]. 

Fecundity—About 300–450 eggs [16]. 

Benthic and reproductive distribution of Ribbed Sculpin (Triglops pingelii).

3-7-14g_RibbedSculpin

Open ocean

Depth, in meters

1,000

800

600

400

200

0

1,000

800

600

400

200

0

Juveniles and adults

Depths documented in

U.S. Chukchi Sea

Triglops pingelii

Ribbed Sculpin

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Characteristic location of 

sculpin larvae 

 Overall depth data from 

outside U.S. Beaufort-Chukchi Seas 

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Some fish to 930 m
326 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—In southeastern Beaufort Sea and northwest Atlantic Ocean, primarily planktonic or epibenthic 

organisms such as amphipods, mysids, copepods, larval ascidians, euphausiids, and arrow worms [23]. 

Trophic level—3.41 standard error 0.51 [24].

Biological Interactions

Predators—Ringed seals and probably bearded seals in Canadian Arctic Ocean [25–27] as well as Thickbilled Murres during summer [28]. Great and Plain sculpins, Pacific Cod, Pacific Halibut, Pigeon Guillemots, 

Tufted Puffins, and Thick-billed Murres in Bering Sea, Gulf of Alaska, and off British Columbia, Canada and 

Kamchatka Peninsula, Russia [29–34]. 

Competitors—Presumably a wide range of other zoobenthos feeders such as Arctic Cod, Walleye Pollock, 

poachers, eelpouts, and other sculpins.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [24].

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Ribbed Scuplin are not commercially harvested.

Potential Effects of Climate Change

The Ribbed Sculpin reproduces in Arctic and Boreal waters [2], so it is difficult to predict how climate 

warming might affect its distribution or abundance. A trend of increasing abundance with latitudinal change is 

hypothesized.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.
Ribbed Sculpin 327

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [15]

Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., 190 p. [6]

Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, Fisheries Research Institute, FRI-UW-7719. [17]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [8]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Alverson, D.L., and Wilimovsky, N.J., 1966, Fishery investigations of the southeastern Chukchi Sea, in Wilimovsky, N.J., 

and Wolfe, J.N., eds., Environment of the Cape Thompson region, Alaska: Oak Ridge, Tennessee, U.S. Atomic Energy 

Commission, Division of Technical Information, p. 843–860.

6. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

7. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

8. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

9. McAllister, D.E., 1963, Fishes of the 1960 “Salvelinus” program from western Arctic Canada: National Museum of Canada 

Bulletin, no. 185, p. 17–39.

10. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.

11. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.
328 Alaska Arctic Marine Fish Ecology Catalog

12. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

13. Coad, B.W., Waszczuk, H., and Labignan, I., 1995, Encyclopedia of Canadian fishes: Canadian Museum of Nature and 

Canadian Sportfishing Productions, Inc., 928 p.

14. Washington, B.B., Moser, H.G., Laroche, W.A., and Richards, W.J., 1984, Scorpaeniformes—Development, in Moser, 

H.G., and others, eds., Ontogeny and systematics of fishes: Gainsville, Florida, American Society of Ichthyologists and 

Herpetolgists, Special Publication 1, 28 p.

15. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

16. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

17. Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, Fisheries Research Institute, FRI-UW-7719.

18. Tokranov, A.M., 1995, The size and sex structure of sculpins of the genus Triglops (Cottidae) in the coastal waters of 

Kamchatka: Journal of Ichthyology, v. 35, no. 4, p. 140–144.

19. Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, National Research Council of Canada, 

p. II-77–II-86.

20. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

21. Blackburn, J.E., and Jackson, P.B., 1982, Seasonal composition and abundance of juvenile and adult marine finfish and crab 

species in the nearshore zone of Kodiak Island’s eastside during April 1978 through March 1979: Outer Continental Shelf 

Environmental Assessment Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 377–570.

22. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

23. Atkinson, E.G., and Percy, J.A., 1992, Diet comparison among demersal marine fish from the Canadian Arctic: Polar 

Biology, v. 11, no. 8, p. 567–573.

24. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

25. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.

26. McLaren, I.A., 1958, The biology of the ringed seal (Phoca hispida Schreber) in the eastern Canadian Arctic: Fisheries 

Research Board of Canada, Bulletin No. 118, 97 p.

27. Finley, K.J., and Evans, C.R., 1983, Summer diet of the bearded seal (Erignathus barbatus) in the Canadian High Arctic: 

Arctic, v. 36, no. 1, p. 82–89.

28. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.

29. Tokranov, A.M., 1981, Rate of sexual maturation and fecundity of the staghorn sculpin, Gymnocanthus detrisus (Cottidae), 

off the east coast of Kamchatka: Journal of Ichthyology, v. 21, no. 1, p. 76–82.

30. Drent, R.H., 1965, Breeding biology of the pigeon guillemot, Cepphus columba: Ardea, v. 53, nos. 3–4, p. 99–159.
Ribbed Sculpin 329

31. Hunter, M.A., 1979, Food resource partitioning among demersal fishes in the vicinity of Kodiak Island, Alaska: Seattle, 

University of Washington, Master’s thesis, 120 p.

32. Hunt, G.L., Burgeson, B., and Sanger, G.A., 1981, Feeding ecology of seabirds of the Eastern Bering Sea, in Hood, D.W., 

and Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and Resources: National Oceanic and Atmospheric 

Administration, p. 629–641.

33. Hatch, S.A., and Sanger, G.A., 1992, Puffins as samplers of juvenile pollock and other forage fish in the Gulf of Alaska: 

Marine Ecology Progress Series, v. 80, p. 1–14.

34. Yang, M.-S., 1996, Diets of the important groundfishes in the Aleutian Islands in summer 1991: Seattle, Washington, U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFSAFSC-60, 105 p.
330 Alaska Arctic Marine Fish Ecology Catalog

Crested Sculpin to Leatherfin Lumpsucker

Crested Sculpin (Blepsias bilobus)

Cuvier, 1829

Family Hemitripteridae

Note: Except for the geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown, but given what is known about the 

species distribution and abundance, likely to be minor with respect 

to marine ecosystem function in the Chukchi Sea.

Physical Description/Attributes: Compressed body densely 

covered in prickles; orange-brown to olive green with darker 

blotches dorsally, or white with olive to black mottling. For specific 

diagnostic characteristics, see Fishes of Alaska (Meckenburg and 

others, 2002, p. 509) [1]. Swim bladder: Absent [2]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea [3]. Elsewhere in Alaska, Bering Sea, and Aleutian Islands from Amchitka Island along coasts to 

southern British Columbia near Port Hardy. Worldwide, Seas of Okhotsk and Japan [3]. 

Crested Sculpin (Blepsias bilobus), 128 mm TL, eastern 

Chukchi Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.
Crested Sculpin 331

Relative Abundance: Uncommon in U.S. Chukchi Sea. [6]. Elsewhere, common in Sea of Japan and eastern Bering Sea north to 

about St. Lawrence Island [3, 7, 8]. 

Geographic distribution of Crested Sculpin (Blepsias bilobus) within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [3, 5]. 

tac14-5222_fig3-8-1m_Blepsias_bilobus

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Crested Sculpin

Blepsias bilobus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
332 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 4–250 m [9, 15], typically less than 120 m [1]. Juveniles collected at the shallower depths and near the surface [9].

3-8-1g-CrestedSculpin

Open ocean

Depth, in meters

500

400

300

200

100

0

Adults and juveniles

Blepsias bilobus

Crested Sculpin

Benthic distribution

Some fish to 225 m

Depth, in meters

Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Spawning and eggs

Reproductive distribution

Characteristic location of sculpin larvae

No data for this species

500

400

100

200

300

0

Adults

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Likely benthic [10]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [10]. 

Juveniles—Age and size: Unknown. Habitat: Benthic, nearshore to upper slope [1]; over bare bottoms, bedrock, 

and among kelp and eelgrass [11]. 

 Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 27 cm TL [1]. 

Habitat: Benthic [1–3], nearshore to upper slope [1]; over bare bottoms, bedrock, and among kelp and 

 eelgrass [11] 

 Substrate—At least bedrock [11]. 

 Physical/chemical—Temperature: -1.4–11.4 °C [15]. Salinity: Marine.

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Probably oviparous [10]. 

Spawning season—Unknown. 

Fecundity—Unknown. 

Benthic and reproductive distribution of Crested Sculpin (Blepsias bilobus).
Crested Sculpin 333

Food and Feeding

Food items—Amphipods and Walleye Pollock (Gadus chalcogrammus) in Russia [13]. Small invertebrates in 

general for members of the Hemitripteridae family [2]. 

Trophic level—4.13 standard error ±0.74 [13].

Biological Interactions

Predators—Great Sculpin [14]. 

Competitors—Likely other sculpins, as well as flatfishes, gadids, and eelpouts.

1980 1990 2000 2010

Resilience

Unknown.

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Crested Sculpin are not commercially harvested.

Potential Effects of Climate Change

This Boreal Pacific species could be expected to increase in abundance and become more widespread in the 

Chukchi Sea, and perhaps to expand its distribution to the Beaufort Sea.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators.

References Cited

Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p. 

[11]

Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and estuarine 

fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California to the 

Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological Resources 

Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p. [9]
334 Alaska Arctic Marine Fish Ecology Catalog

Mecklenburg, C.W., 2003, Family Hemitripteridae Gill 1872—Sea ravens or sailfin sculpins: California Academy of Sciences 

Annotated Checklists of Fishes no. 5, 6 p. [2]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., 2003, Family Hemitripteridae Gill 1872—Sea ravens or sailfin sculpins: California Academy of 

Sciences Annotated Checklists of Fishes no. 5, 6 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

6. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

7. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

8. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

9. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

10. Matarese, A.C., Kendall, J., Arthur W., Blood, D.M., and Vinter, B.M., 1989, Laboratory guide to early life history stages 

of northeast Pacific fishes: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National 

Marine Fisheries Service Technical Report NMFS 80, 652 p.

11. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

12. Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution in the Kronitsky Bay in relation to abiotic factors: Zoologicheskii 

Zhurnal, v. 63, p. 1,363–1,373.

13. Froese, R., and Pauly, D., eds., 2012, FishBase—World Wide Web electronic publication—Global information system on 

fishes: FishBase, accessed July 8, 2012, at http://www.fishbase.org.

14. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.

15. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.
Eyeshade Sculpin 335

Eyeshade Sculpin (Nautichthys pribilovius) 

(Jordan & Gilbert, 1898)

Family Hemitripteridae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unlikely to be of significant importance in 

regional foods webs. 

Physical Description/Attributes: Brown to gray-brown body with 

vague dark blotches and covered with prickles that extend onto 

dorsal and pectoral fins; black bar from eye diagonally downward 

on cheek. For specific diagnostic characteristics, see Fishes of 

Alaska (Mecklenburg and others, 2002, p. 512) [1]. Swim bladder: 

Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea and western U.S. Beaufort Sea [3, 4]. Elsewhere in Alaska, in Bering Sea and Aleutian Islands from 

Attu Island to southeastern Alaska at Steamer Bay. Worldwide, Gulf of Anadyr to Commander Islands and to Seas of Okhotsk 

and Japan [1, 3].

Relative Abundance: Common in eastern Chukchi Sea [4, 7] and more common in Bering Strait and southeastern Chukchi Sea 

[6] and eastern Bering Sea [8] but rare in Sea of Japan [9] and U.S. Beaufort Sea.

Eyeshade Sculpin (Nautichthys pribilovius), 109 mm TL, 

Bering Strait, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Eyeshade Sculpin (Nautichthys pribilovius) within Arctic Outer Continental 

Shelf Planning Areas [5] based on review of published literature and specimens from historical and recent 

collections [1, 3, 4, 6]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Eyeshade Sculpin

Nautichthys pribilovius

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
336 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 11 m [4] to 422 m, typically less than 135 m in Bering Sea and Gulf of Alaska [1]. Documented at depths of 

11–53 m [4] in the U.S. Chukchi Sea and 24–53 m in the U.S. Beaufort Sea [4].

Habitats and Life History

Eggs—Size: 2.5–2.7 mm [10]. Time to hatching: Unknown. Habitat: Likely benthic based on life history of the 

Sailfin Sculpin (Nautichthys oculofasciatus) [11]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Likely pelagic based on life history of Sailfin Sculpin [11]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Unknown. Maximum Age: Unknown. Maximum size: As long as 10.9 cm 

TL [1]. Habitat: Benthic [1]. 

Substrate—Shell hash, gravel, sand, mud and rock in U.S. Chukchi Sea [6]. 

Physical/chemical—Temperature: -1.7–11.4 °C [4, 6, 12]. Prefers temperatures greater than 0 °C [10]. Salinity: 

Marine to slightly estuarine [13]. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [10]. 

Spawning season—Unknown. 

Fecundity—50 eggs [10].

3-8-2g_EyeshadeSculpin

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Some fish to 422 m

Nautichthys pribilovius

Eyeshade Sculpin

Benthic distribution

Overall depth data from outside 

U.S. Beaufort-Chukchi Seas 500

400

300

200

100

0

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Depth range of spawning is uncertain. 

Potential location of spawning and eggs 

is based on depth range of benthic individuals

Adults and juveniles 

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Larvae are pelagic.

Specific depths unknown

Depths from U.S. Beaufort-Chukchi Seas

Benthic and reproductive distribution of Eyeshade Sculpin (Nautichthys pribilovius).
Eyeshade Sculpin 337

Food and Feeding

Food items—Unknown. Small invertebrates in general for members of the Hemitripteridae family [2]. 

Trophic level—3.62 standard error 0.65 [14].

Biological Interactions

Predators—Unknown. 

Competitors—Unknown. 

1980 1990 2000 2010

Resilience

Unknown. No data. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Eyeshade Sculpin are not commercially harvested.

Potential Effects of Climate Change

A predominantly Boreal Pacific species [3], Eyeshade Sculpin are apt to become more abundant in the Chukchi 

and Beaufort Seas as water temperatures increase. A range expansion in the Beaufort Sea also is likely.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year Demersal 

recruitment, (3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and 

ontogenetic movements, (6) population studies, (7) prey, and (8) predators. 

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., avaliable from U.S. Department of Commerce, Springfield, Virginia] [10]
338 Alaska Arctic Marine Fish Ecology Catalog

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [6]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140 and Online Resource 1. [3]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., 2003, Family Hemitripteridae Gill 1872—Sea ravens or sailfin sculpins: California Academy of 

Sciences Annotated Checklists of Fishes no. 5, 6 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

7. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

8. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

9. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

10. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., avaliable from U.S. Department of Commerce, Springfield, Virginia]

11. Matarese, A.C., Kendall, J., Arthur W., Blood, D.M., and Vinter, B.M., 1989, Laboratory guide to early life history stages 

of northeast Pacific fishes: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National 

Marine Fisheries Service Technical Report NMFS 80, 652 p.

12. Cui, X., Grebmeier, J.M., Cooper, L.W., Lovvorn, J.R., North, C.A., Seaver, W.L., and Kolts, J.M., 2009, Spatial 

distributions of groundfish in the northern Bering Sea in relation to environmental variation: Marine Ecology Progress 

Series, v. 393, p. 147–160.

13. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

14. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Polar Sculpin 339

Polar Sculpin (Cottunculus microps)

Collett, 1875

Family Psychrolutidae

Note on taxonomy: Based on recent taxonomic revision Cottunculus 

sadko (Essipov, 1937) is treated as a synonym of C. microps [1, 2]. Data 

on fish originally identified as C. sadko are included here.

Note: Except for geographic range data, all information is from areas 

outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Minor in the U.S. Beaufort Sea.

Physical Description/Attributes: Tadpole shaped body with moveable skin over a clear, gelatinous layer. Body is greyish with 

three vague, dark bands. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 518 [as 

C. sadko]) [3]. Swim bladder: Absent [3]. Antifreeze glycoproteins in blood serum: Unknown.

Range: One record from continental slope off U.S. Beaufort Sea. One record outside the U.S. 200-mile limit north of the eastern 

Chukchi Sea. North to about 82°N [4]; circumpolar; Kara Sea slope in Voronin Trough, Laptev Sea slope off Bolshevik Island, 

Faroe Trough, Nansen Rise off Spitsbergen, western Barents Sea, slope off Franz Josef Land; and south to Gulf of St. Lawrence 

and New Jersey, English Channel, and North Sea [4]. 

Relative Abundance: Rare in U.S. Beaufort Sea [4]. Common eastward from Hudson Strait and Baffin Bay to Greenland and 

the Barents Sea [1, 7, 8].

Polar Sculpin (Cottunculus microps) 135 mm TL, Chukchi 

Borderland, 2009. Photograph by C.W. Mecklenburg, 

Point Stephens Research.

Geographic distribution of Polar Sculpin (Cottunculus microps) within Arctic Outer Continental Shelf Planning 

Areas [5] in the study area based on review of published literature and specimens from historical and recent 

collections [4, 6]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Polar Sculpin

Cottunculus microps

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
340 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Upper continental slopes at 159–2,476 m [6], mainly 170–400 m in Barents Sea [1].

Habitats and Life History

Eggs—Size: 3–5 mm [1]. Time to hatching: Unknown. Habitat: Benthic [9]. 

Larvae—Size at hatching: Size at juvenile transformation: Days to juvenile transformation: Habitat: Unknown. 

Juveniles—Age and size: Unknown. Habitat: Benthic, deep troughs, and upper parts of continental slope [3]. 

Adults—Age and size at first maturity: Unknown. Maximum age: 8–10 years [1]. Maximum size: As long 

as 33 cm TL (mainly 6–16 cm) and 0.7 kg [1]. Habitat: Benthic, deep troughs, and upper parts of continental 

slope [3]. 

Substrate—Mud, soft bottoms [1, 3]. 

Physical/chemical—Temperature: -0.4–1.9 °C in Baffin Bay [10]. Typically temperatures greater than 0 °C [1]. 

Salinity: Prefers high salinity (34.5–35 ppt) [1].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Adults of this family typically guard their eggs [7]. 

Schooling—Unknown. Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [7]. 

Spawning season—Summer–autumn in Barents Sea [1]. 

Fecundity—125–435 eggs [1].

Food and Feeding

Food items—Benthic invertebrates such as pantopods, amphipods, gammarids, polychaetes, mysids, mollusks, 

etc. [1, 11]. 

Trophic level—3.41 standard error 0.39 [12].

Benthic and reproductive distribution of Polar Sculpin (Cottunculus microps).

3-8-3g_PolarSculpin

Open ocean Shore

Depth, in meters

1,500

1,200

900

600

300

0

1,500

1,200

900

600

300

0

Adults and juveniles Spawning and eggs

Larvae locations unknown

Some fish to 1,458 m

Cottunculus microps

Polar Sculpin

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Reproductive distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Depth range of spawning is 

uncertain. Potential location of 

spawning and eggs is based on 

depth range of benthic individuals

Data from outside 

U.S. Beaufort-Chukchi Seas
Polar Sculpin 341

Biological Interactions

Predators—Greenland halibut in Barents Sea [13]. 

Competitors—Juveniles would likely be consumed by a range of benthic feeders, such as flatfishes, cods, 

sculpins, and poachers.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (Fecundity = 66) [12].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Polar Sculpin are not commercially harvested.

Potential Effects of Climate Change

Unknown.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators. 

References Cited

Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p. [9]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]

Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, A., 

Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, G., and Smirnov, O., 2011, 

Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p. [1]
342 Alaska Arctic Marine Fish Ecology Catalog

Bibliography

1. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

2. Byrkjedal, I., Hadler-Jacobsen, S., Rees, D., and Orlov, A., 2013, Molecular and morphological variation in ‘saddled’ 

fathead sculpins (Psychrolutidae—Cottunculus) of the northeast Atlantic: Journal of Fish Biology, v. 85, no. 3, p. 821–837.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Møller, P.R., Nielsen, J.G., Knudsen, S.W., Poulsen, J.Y., Sünksen, K., and Jørgensen, O.A., 2010, A checklist of the fish 

fauna of Greenland waters: Zootaxa, v. 2378, p. 1–84.

8. Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic marine fishes of Canada: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, no. 2674, 112 p.

9. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

10. Jørgensen, O.A., Hvingel, C., and Treble, M.A., 2011, Identification and mapping of bottom fish assemblages in northern 

Baffin Bay: Journal of Northwest Atlantic Fisheries Science, v. 43, p. 65–79.

11. Bjelland, O., Bergstad, O.A., Skjæraasen, J.E., and Meland, K., 2000, Trophic ecology of deep-water fishes associated with 

the continental slope of the eastern Norwegian Sea: Sarsia, v. 85, no. 2, p. 101–117.

12. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

13. Hovde, S.C., Albert, O.T., and Nilssen, E.M., 2002, Spatial, seasonal and ontogenetic variation in diet of Northeast Arctic 

Greenland halibut (Reinhardtius hippoglossoides): ICES Journal of Marine Science, v. 59, no. 2, p. 421–437.
Smoothcheek Sculpin 343

Smoothcheek Sculpin (Eurymen gyrinus) 

Gilbert & Burke, 1912

Family Psychrolutidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Scarcity within U.S. Chukchi Sea waters and 

absence from the U.S. Beaufort Sea suggests a minor ecological 

role.

Physical Description/Attributes: Tadpole-shaped body with 

moveable skin over a clear, gelatinous layer. Color is highly 

variable according to habitat [1]. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 

2002, p. 517) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea to 70°26’N, 164°38’W [2]. Elsewhere in Alaska, Bering Sea south to Aleutian Islands and to 

western Gulf of Alaska at Kodiak Island. Worldwide, Seas of Japan and Okhotsk and Pacific coast of Honshu, and off Japan to 

Commander Islands [1]. 

Relative Abundance: Rare in U.S. Chukchi Sea and absent from U.S. Beaufort Sea [1, 2, 4]; typically caught in shallow waters, 

but rarely sampled by research investigations. Rare in Sea of Japan [5].

Smoothcheek Sculpin (Eurymen gyrinus), 111 mm TL, northern 

Bering Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Smoothcheek Sculpin (Eurymen gyrinus) within Arctic Outer Continental Shelf 

Planning Areas [3] based on review of published literature and specimens from historical and recent 

collections [1, 2, 4].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Smoothcheek Sculpin

Eurymen gyrinus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
344 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 2–174 m [2, 6], typically 40–100 m in the Pacific Ocean off the northern Kuril Islands, Russia [7]. Found at 

depths of 17–44 m in the U.S. Chukchi Sea [2].

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Likely benthic, like the more well-known 

members of the family Psychrolutidae [8]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Unknown. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. Adults: Age and size at first maturity: Unknown. 

Maximum age: Unknown. Maximum size: As long as 38.8 cm TL [1]. Habitat: Benthic [1]. 

Substrate—Unknown. 

Physical/chemical—Temperature: Unknown. Salinity: Marine [1]. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Adults of this family typically guard their eggs [8]. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [8]. 

Spawning season—Unknown. 

Fecundity—Unknown.

Food and Feeding

Food items—Unknown. 

Trophic level—3.41 standard error 0.46 [9].

Benthic and reproductive distribution of Smoothcheek Sculpin (Eurymen gyrinus).

3-8-4g_SmoothcheekSculpin

Spawning and eggs

Open ocean Shore

Depth, in meters

500

400

300

200

100

0 Depths documented in 

U.S. Chukchi Sea

Some fish to 494 m

Eurymen gyrinus

Smoothcheek Sculpin

Benthic distribution

Overall depth data from outside

U.S. Beaufort-Chukchi Seas 500

400

300

200

100

0

Open ocean No data for this species Shore

Depth, in meters

Depth range of

larvae unknown

Reproductive distribution

Depth range of spawning is uncertain. 

Potential location of spawning and eggs is 

based on depth range of benthic individuals

Adults and juveniles 

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown
Smoothcheek Sculpin 345

Biological Interactions

Predators—Unknown. 

Competitors—Unknown.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (Preliminary K or Fecundity) [9].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Smoothcheek Sculpin are not commercially harvested.

Potential Effects of Climate Change

It is likely that warming Arctic waters would allow this species to penetrate farther into the U.S. Chukchi Sea and 

perhaps into the U.S. Beaufort Sea.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators.

References Cited

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]
346 Alaska Arctic Marine Fish Ecology Catalog

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland: American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

6. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

7. Fedorov, V.V., 2000, Species composition, distribution and habitation depths of the northern Kuril Islands fish and fish like 

species, in Koteneva, B.N., ed., Commercial and biological studies of fishes in the Pacific waters of the Kuril Islands and 

adjacent areas of the Okhotsk and Bering Seas in 1992–1998: Moscow, VNIRO Publishing, collected papers, p. 7–41. [In 

Russian.]

8. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

9. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Alligatorfish 347

Alligatorfish (Aspidophoroides monopterygius)

(Bloch, 1786)

Family Agonidae 

Note: Except for physical description, relative abundance, and 

geographic range data, all information is from areas outside of the 

study area. 

Note on taxonomy: The Alaskan population was described and 

named as a distinct species, Aspidophoroides bartoni (Gilbert, 

1896), to which the common name Aleutian Alligatorfish has been applied [1]. Recent genetic and morphological data confirm 

that they are the same species [1, 2].

Colloquial Name: None within U.S. Chukchi and Beaufort Seas. 

Ecological Role: Current information about the occurrence of this fish is limited in the U.S. Chukchi and Beaufort Seas. It is 

unlikely to represent a significant prey resource in regional food webs. 

Physical Description/Attributes: Brownish to olive green with a slender body covered with bony plates; faint dark bands on 

sides and back; and a white belly. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 

553) [1]. Swim bladder absent [3]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea northward to Point Barrow and Beaufort Sea eastward to about 71°N, 151°W [2, 4]. Elsewhere in 

Alaska, in Bering Sea and Aleutian Islands, to Prince William Sound and Gulf of Alaska [1]. Worldwide, in western Pacific from 

Sea of Japan and Sea of Okhotsk to Gulf of Anadyr, and western Atlantic Ocean from northwestern Greenland to New Jersey 

[1, 2]. 

Alligatorfish (Aspidophoroides monopterygius), 84 mm TL, 

Chukchi Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.
348 Alaska Arctic Marine Fish Ecology Catalog

Relative Abundance: Common in U.S. Chukchi Sea [2, 4, 6, 7]. In Alaska, common in eastern Bering Sea, and along the 

western end of the Alaska Peninsula [8–10].Worldwide, common in nearshore waters off Kamchatka, Russia [11].

tac14-5222_fig3-8-5m_Aspidophoroides_monopterygius

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Alligatorfish

Aspidophoroides monopterygius

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Alligatorfish (Aspidophoroides monopterygius) within Arctic Outer Continental 

Shelf Planning Areas [5] based on review of published literature and specimens from historical and recent 

collections [1, 2, 4]. 
Alligatorfish 349

Depth Range: Overall range is 8–337 m, but almost entirely less than 200 m [1]. Depth range of larvae is unknown. In general, 

poacher larvae are abundant in near-surface waters, and over continental shelf and upper slope [12–14]. Depth range of juveniles 

is unknown.

Benthic reproductive distribution of Alligatorfish (Aspidophoroides monopterygius). 

Some fish to 350 m

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Juveniles and adults

Aspidophoroides monopterygius

Alligatorfish

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

500

400

300

200

100

0

Spawning and eggs

Depth range of spawning is

uncertain. Potential location

of spawning and eggs is based

on depth range of benthic 

individuals

Overall benthic depth

range. Specific depth

range of either juveniles 

or adults is unknown

 Data from outside

U.S. Beaufort-Chukchi Seas 

Reproductive distribution

Characteristic location of poacher larvae

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Demersal and adhesive (probably on sand and 

mud) [3]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 29.0 mm in western North Atlantic 

[15]. Days to juvenile transformation: Unknown. Habitat: Pelagic although distance from shore and depth in 

water column are unknown for this species; in general poacher larvae are abundant in near-surface waters, over 

continental shelf and shallow slope waters [12–14]. 

Juveniles—Age and size: Unknown. Habitat: Demersal [1, 3, 16], although fish to at least 4.1 cm TL are 

occasionally found in midwaters [17]. 

Adults—Age and size at first maturity: Unknown. Maximum age: At least 6 years. Females grow larger and may 

have a slightly longer life span [18]. Maximum size: 17.8 cm TL [1]. Habitat: Demersal [3, 16]. 

Substrate—Juveniles and adults live on sand and mud [1, 19]. 

Physical/chemical—Temperature: Between at least -1.7–12.2 °C. [19, 20]. Salinity: All life stages live in marine 

waters. [19–21].

Behavior

Diel—Unidentified poacher larvae have migrated into slightly deeper waters at night in southeastern Alaska [13]. 

Seasonal—Off Kamchatka, found in relatively shallow waters during summer and during winters in deeper 

waters on the edge of the continental shelf [11]. 

Reproductive—Unknown. 

Schooling—Unlikely. Other poacher species are solitary [16, 22]. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.
350 Alaska Arctic Marine Fish Ecology Catalog

Reproduction

Mode—Separate sexes, oviparous [23]. 

Spawning season—Poorly known. November to January in Atlantic Ocean [24]. 

Fecundity—Poorly known. One “medium-sized” Alligatorfish in Atlantic Ocean specimen contained about 600 

eggs [19].

Food and Feeding

Food items—Benthic individuals eat mostly zoobenthos, including crustaceans (for example, amphipods, 

isopods, cumaceans) and polychaetes [25, 26], although midwater animals such as pteropods are also consumed 

[26]. Food habits of larvae are unknown. 

Trophic level—3.5 (standard error 0.43) [27].

Biological Interactions

Predators—Cannibalism has not been reported. Bering Sea, Kamchatka Peninsula and northern Kuril Island, 

Russia, predators include Flathead Sole, Great Sculpin, Plain Sculpin, Pacific Cod, Pacific halibut, and Sablefish 

[28–34]. Of these species, only great and plain sculpins and Pacific Halibut inhabit the U.S. Chukchi Sea, and 

Great Sculpin and Pacific Halibut are rarely found there. 

Competitors—Presumably other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other poachers, 

eelpouts, and sculpins. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time is 1.4–4.4 years (Preliminary K or Fecundity) [27].

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Alligatorfish are not commercially harvested.

Potential Effects of Climate Change

Alligatorfish are common in marine waters of Arctic Alaska in temperatures 1.1–2.5 °C. Warming could lead to 

increased patterns of abundance and that could lead to increased competition with other fishes that feed on small 

benthic organisms. This would potentially, include Arctic Cod, Walleye Pollock, other poachers, and sculpins.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs in the study area include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.
Alligatorfish 351

References Cited

Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and estuarine 

fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California to the 

Alaska-Yukon Border: U.S. Department of the Interior, U.S. Geological Survey, Biological Resources Division, OCS Study 

MMS 2005-030 and USGS/NBII 2005-001, 276 p. [35]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p. [19]

Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p. [3]

Tokranov, A.M., 1987b, Species composition and distribution of sea poachers (Pisces, Agonidae) in nearshore waters of 

Kamchatka: Zoologicheskii Zhurnal, v. 66, p. 385–392. [11]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

7. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

8. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

9. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1,

281 p.

10. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

11. Tokranov, A.M., 1987b, Species composition and distribution of sea poachers (Pisces, Agonidae) in nearshore waters of 

Kamchatka: Zoologicheskii Zhurnal, v. 66, p. 385–392.

12. Boehlert, G.W., Gadomski, D.M., and Mundy, B.C., 1985, Vertical distribution of ichthyoplankton off the Oregon coast in 

spring and summer months: Fishery Bulletin, v. 83, no. 4, p. 611–621.

13. Haldorson, L., Pritchett, M., Sterritt, D., and Watts, J., 1993, Abundance patterns of marine fish larvae during spring in a 

southeastern Alaskan bay: Fishery Bulletin, v. 91, no. 1, p. 36–44.
352 Alaska Arctic Marine Fish Ecology Catalog

14. Ambrose, D.A., Carter, R.L., and Manion, S.M., 2006, Ichthyoplankton and station data for surface (manta) and oblique 

(bongo) plankton tows for California Cooperative Oceanic Fisheries Investigations survey cruises in 2005: U.S. Department 

of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NOAA-TM-NMFSSWFSC-392, 125 p.

15. Fahay, M.P., 1983, Early stages of fishes in the western North Atlantic Ocean (Davis Strait, Southern Greenland and 

Flemish Cap to Cape Hatteras), Volume 2—Scorpaeniformes through Tetraodontiformes: Dartmouth, Nova Scotia, Canada, 

p. 932–1696.

16. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, nos. 1–3, p. 15–28.

17. Dragoo, D.E., 2006, Seabird, fish, marine mammals and oceanography coordinated investigations (SMMOCI) near Bluff, 

Norton Sound, Alaska, July 2002: U.S. Fish and Wildlife Service Report AMNWR 06/03, 35 p.

18. Tokranov, A.M., 1993, Sexual dimorphism in sea poachers (Agonidae) off Kamchatka: Journal of Ichthyology, v. 33, no. 3, 

p. 113–122.

19. Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p.

20. Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution in the Kronitsky Bay in relation to abiotic factors: Zoologicheskii 

Zhurnal, v. 63, p. 1,363–1,373.

21. Cui, X., 2009, Climate-driven impacts of groundfish on food webs in the northern Bering Sea: Knoxville, University of 

Tennessee, Ph.D. dissertation, 162 p.

22. Love, M.S., University of California-Santa Barbara, pers. obs., 2009.

23. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

24. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

25. Tokranov, A.M.,1992, Dietary characteristics of sea poachers (Agonidae) in the coastal waters of Kamchatka: Journal of 

Ichthyology v. 32, no. 9, p. 85–94.

26. Hutchings, J.A., 2002, Ecology and biodiversity of commercially unexploited marine fishes in the northwest Atlantic: 

Halifax, Nova Scotia, Canada, Dalhousie University, Department of Biology, unpub. report, p. 14.

27. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

28. Hunter, M.A., 1979, Food resource partitioning among demersal fishes in the vicinity of Kodiak Island, Alaska: Seattle, 

University of Washington, Master’s thesis, 120 p.

29. Tokranov, A.M., 1987a, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok

(Cuvier) (Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114.

30. Orlov, A.M., 1999, Trophic relationships of commercial fishes in the Pacific waters off southeastern Kamchatka and the 

northern Kuril Islands, in Ecosystem approaches for fisheries management 1999: Alaska Sea Grant College Program 

AK-SG-99-01, p. 231–264.

31. Pacunski, R.E., Livingston, P.A., and Miller, B.S., 1998, Food of flathead sole Hippoglossoides elassodon in the eastern 

Bering Sea: National Oceanic and Atmospheric Administration, Technical Memorandum NMFS-AFSC-90, 36 p.

32. Napazakov, V.V., and Chuchukalo, V.I., 2003, Diet and trophic status of mass species of sculpins (Cottidae) in the western 

part of the Bering Sea in autumn: Journal of Ichthyology, v. 43, no. 3, p. 236–244.

33. Yang, M.-S., 2003, Food habits of the important groundfishes in the Aleutian Islands in 1994 and 1997: National Marine 

Fisheries Service, AFSC Processed Report 2003-07, 233 p.

34. Yang, M.-S., Dodd, K., Hibpshman, R., and Whitehouse, A., 2006, Food habits of groundfishes in the Gulf of Alaska 

in 1999 and 2001: U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical 

Memorandum, NMFS-AFSC-164, 189 p.

35. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.
Arctic Alligatorfish 353

Arctic Alligatorfish (Aspidophoroides olrikii) 

Lütken, 1877

Family Agonidae

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Largely unknown. Species information regarding 

biology and ecology is not available. Although this species appears 

to be common in the U.S. Chukchi and Beaufort Seas, it does not 

school and its biomass is likely to be relatively low. Thus, this 

species is unlikely to represent a significant prey resource to birds, 

mammals, and larger fishes.

Physical Description/Attributes: Brownish to olive green on back 

and sides and white on belly. Caudal fin has a white margin and center [1]. Males have a chalky white blotch on dorsal fin [2]. 

For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 552) [1]. Swim bladder: Absent 

[3]. Antifreeze glycoproteins in blood serum: Unknown.

Range: In U.S. Beaufort Sea and northward to at least 71°55’N, 175°18’W in U.S. Chukchi Sea [4]. Elsewhere in Alaska, south 

through the Bering Sea to northern Gulf of Alaska in Prince William Sound [5]. Worldwide, practically circumpolar in Arctic 

Ocean, from southern Barents Sea and White Sea eastward to west Greenland but not Canadian high Arctic archipelago; south in 

western Atlantic Ocean to Newfoundland [1, 2, 4, 5]. 

Relative Abundance: Common throughout U.S. Chukchi Sea and U.S. Beaufort Seas [1, 2, 4, 7, 8]. Reports of presence 

between Herschel Island and Franklin Bay, Canada [9–11] and numerous unpublished museum specimens from Alaska eastward 

to Hudson Bay indicate this species is common in Arctic Canadian waters [2].

Arctic Alligatorfish (Aspidophoroides olrikii), 61 mm TL, 

Chukchi Sea, 2004. Photograph by B.A. Sheiko and C.W. 

Mecklenburg, Russian Academy of Sciences and Point 

Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Alligatorfish

Aspidophoroides olrikii

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Arctic Alligatorfish (Aspidophoroides olrikii) within Arctic Outer Continental 

Shelf Planning Areas [6] based on review of published literature and specimens from historical and recent 

collections [1, 2, 5].
354 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: As deep as 400 m in U.S. Chukchi and Beaufort Seas [7], usually less than 100 m [1, 4, 5, 8, 12]. Elsewhere, 

overall depth range is 3–520 m [2]. Depth range of pelagic juveniles and larvae is poorly known. One pelagic larva documented 

between 48 m and surface in the U.S. Chukchi Sea [13]. Off Oregon and California, poacher larvae are abundant in near-surface 

waters, over the continental shelf and upper slope waters [14–16].

Benthic and reproductive distribution of Arctic Alligatorfish (Aspidophoroides olrikii).

Habitats and Life History

Eggs—Size: 0.75–1.5 mm [17]. Time to hatching: Unknown. Habitat: Demersal and adhesive (probably on sand 

and mud) [3]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 2.5–6.2 cm TL [7, 12, 18]. Days to 

juvenile transformation: 2–3 months [3]. Habitat: Pelagic although distance from shore and depth in water 

column are unknown for this species, in general poacher larvae are abundant in near-surface waters, over 

continental shelf and shallow slope waters [14–16]. 

Juveniles—Age and size: Unknown. Habitat: Demersal [3, 19]. 

Adults—Age and size at first maturity: Age unknown. About 62 mm off Russia [12]. Maximum age: Unknown. 

Maximum size: 10 cm TL [2]. Habitat: Demersal [1, 3, 19]. 

Substrate—Sand and mud bottoms; also lives around rocks [4, 12, 20]. 

Physical/chemical—Temperature: Juveniles and adults at least -1.8–7.9 °C [2, 21–23], prefers around 0 °C [24]. 

Salinity: Marine and estuarine waters as brackish as 20 ppt [12, 25], prefers 33–35 ppt [24].

Behavior

Diel—Unidentified poacher larvae have migrated into slightly deeper waters at night in southeastern Alaska [15]. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unlikely. Other poacher species are solitary [19, 26]. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous [27]. 

Spawning season—Unknown. 

Fecundity—At least 110–260 eggs [7, 12].

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Aspidophoroides olrikii

Arctic Alligatorfish

Benthic distribution

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Reproductive distribution

Juveniles and adults

Spawning and eggs

Characteristic location of poacher larvae 

Some fish to 520 m

outside study area

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Data from U.S. Beaufort-Chukchi Seas

One pelagic larvae documented

in U.S. Chukchi Sea between 0–48 m 

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 
Arctic Alligatorfish 355

Food and Feeding

Food items—A few fish examined in the U.S. Chukchi and Beaufort seas had eaten gammarid amphipods, 

polychaetes, and clam siphons [7, 28], whereas fish in the Russian Arctic consumed amphipods, isopods, 

ostracods, and nemerteans [12]. Food habits of larvae are unknown. 

Trophic level—3.3 standard error 0.49 [29].

Biological Interactions

Predators—Unknown. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other 

poachers, eelpouts, and sculpins. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (assuming t

m=2–4; Fecundity <2,000) [21]

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Arctic Alligatorfish are not commercially harvested.

Potential Effects of Climate Change

This is a predominantly Arctic species; thus, the distribution of Arctic Alligatorfish would be expected to shift 

northwards with warming temperatures. 

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs in the study area include: 

(1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movement,; (6) population studies, (7) prey, and (8) predators.

References Cited

Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western Beaufort 

Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA Technical Report 

NMFS-SSRF-764, 22 p. [7]
356 Alaska Arctic Marine Fish Ecology Catalog

Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and estuarine 

fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California to the 

Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological Resources 

Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p. [30]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi Sea: 

Deep-Sea Research II, v. 57, no. 1–2, p. 57–70. [13]

Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p. [3]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

3. Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p.

4. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

5. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140 and Online Resource 1.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western 

Beaufort Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA 

Technical Report NMFS-SSRF-764, 22 p.

8. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

9. McAllister, D.E., 1963, Fishes of the 1960 “Salvelinus” program from western Arctic Canada: National Museum of Canada 

Bulletin, no.185, p. 17–39.

10. Arctic Laboratories Limited, 1987, Beaufort ocean dumpsite characterization: Prepared for Environment Protection, 

Conservation and Protection, Environment Canada, Yellowknife, Northwest Territories, by Arctic Laboratories Limited and 

LGL Limited.

11. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

12. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

13. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.
Arctic Alligatorfish 357

14. Boehlert, G.W., Gadomski, D.M., and Mundy, B.C., 1985, Vertical distribution of ichthyoplankton off the Oregon coast in 

spring and summer months: Fishery Bulletin, v. 83, no. 4, p. 611–621.

15. Haldorson, L., Pritchett, M., Sterritt, D., and Watts, J., 1993, Abundance patterns of marine fish larvae during spring in a 

southeastern Alaskan bay: Fishery Bulletin, v. 91, no. 1, p. 36–44.

16. Ambrose, D.A., Carter, R.L., and Manion, S.M., 2006, Ichthyoplankton and station data for surface (manta) and oblique 

(bongo) plankton tows for California Cooperative Oceanic Fisheries Investigations survey cruises in 2005: U.S. Department 

of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NOAA-TM-NMFSSWFSC-392, 125 p.

17. Fahay, M.P., 2007, Agonidae, in Fahay, M.P., Early stages of fishes in the western North Atlantic Ocean, volume 2: 

Dartmouth, Nova Scotia, Canada, Northwest Atlantic Fisheries Organization, p. 982–986.

18. Busby, M.S., 1998, Guide to the identification of larval and early juvenile poachers (Scorpaeniformes: Agonidae) from the 

northeastern Pacific Ocean and Bering Sea: National Oceanic and Atmospheric Administration Technical Report 137, 88 p.

19. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, nos. 1–3, p. 15–28.

20. Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p.

21. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

22. Cui, X., 2009, Climate-driven impacts of groundfish on food webs in the northern Bering Sea: Knoxville, University of 

Tennessee, Ph.D. dissertation, 162 p.

23. Andriashev, A.P., 1986, Agonidae, in Whitehead, P.J.P., ed., Fishes of the north-eastern Atlantic and the Mediterranean: 

Paris, United Nations Educational, Scientific, and Cultural Organization, p. 1,265–1,268.

24. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

25. Schneider-Vieira, F., Baker, R., and Lawrence, M., 1993, The estuaries of Hudson Bay—A case study of the physical and 

biological characteristics of selected sites: Report prepared for the Hudson Bay Program by North-South Consultants Inc., 

26 p.

26. Love, M.S., University of California-Santa Barbara, pers. obs., 2009.

27. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

28. Atkinson, E.G., and Percy, J.A., 1992, Diet comparison among demersal marine fish from the Canadian Arctic: Polar 

Biology, v. 11, no. 8, p. 567–573.

29. Bureau of Offshore Energy Management, Table of population and behavioral ecology characteristics: Alaska OCS Region, 

unpub. data, 2010.

30. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.
358 Alaska Arctic Marine Fish Ecology Catalog

Fourhorn Poacher (Hypsagonus quadricornis) 

(Valenciennes, 1829)

Family Agonidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area.

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Current information about the occurrence of this 

fish is limited in the U.S. Chukchi Sea. Its rare occurrence suggests 

it is not important in local or regional food webs.

Physical Description/Attributes: Reddish brown and bony-plated 

with cryptic coloration of white, yellow, red, and brown bands and 

blotches, and a dark vertical band along margin of caudal fin. For 

specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 531) [1]. Swim bladder: Absent [2]. 

Antifreeze glycoproteins in blood serum: Unknown.

Range: From Bering Strait to northeastern Chukchi Sea west of Point Lay [3]. Found in the U.S. Chukchi Sea for the first time 

in 2007 in a region, which had been extensively sampled, this boreal species could be new to the Arctic [3, 4]. Elsewhere, from 

Commander–Aleutian Islands chain and southern Bering Sea southward to Puget Sound, Washington, and to Sea of Okhotsk and 

northern Sea of Japan [1, 3–7].

Relative Abundance: Rare in U.S. Chukchi Sea [3]. Elsewhere in Alaska, rare north of 57°N in Bering Sea [3]. Common 

around the Aleutian Islands, Pribilof Islands, and eastward to Cook Inlet, Gulf of Alaska, southward to Puget Sound, 

Washington, and around the Kuril Islands and Kamchatka Peninsula, Russia.

Fourhorn Poacher (Hypsagonus quadricornis), 103 mm TL, 

Bering Strait, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Fourhorn Poacher

Hypsagonus quadricornis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Fourhorn Poacher (Hypsagonus quadricornis) within Arctic Outer Continental 

Shelf Planning Areas [8] based on review of published literature and specimens from historical and recent 

collections [3, 4]. 
Fourhorn Poacher 359

Depth Range: From intertidal zone to 452 m [1], but most abundant at less than 200 m [1, 5, 9, 10]. Occurs at depths less than 

110 m in northern Bering Sea and U.S. Chukchi Sea [4]. In the Pacific Ocean off the northern Kuril Islands, spawning occurs at 

depths of 110–150 m [10]. Depth range of juveniles is unknown. Depth range of larvae is unknown. In general, poacher larvae 

are abundant in near-surface waters, over continental shelf and upper slope [11–13].

Some fish to 452 m

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Hypsagonus quadricornis

Fourhorn Poacher

Benthic distribution

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Reproductive distribution

Juveniles and adults

Depth documented in

U.S. Beaufort-Chukchi Seas

Spawning and eggs

Characteristic location of poacher larvae

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Specific depth of eggs is uncertain.

Potential location is based on

depth range of spawning individuals 

Data from outstide U.S. Beaufort-Chukchi Seas Data from outstide U.S. Beaufort-Chukchi Seas

Benthic and reproductive distribution of Fourhorn Poacher (Hypsagonus quadricornis).

Habitats and Life History

Eggs—Size: 1.5–3.1 mm [10]. Time to hatching: Unknown. Habitat: Demersal and adhesive [2]. 

Larvae—Size at hatching: 6.4 mm [14]. Size at juvenile transformation: 14 mm [14]. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [2]. In general, poacher larvae are abundant in near-surface waters, 

over continental shelf and shallow slope waters [11–13]. 

Juveniles—Age and size: Unknown. Habitat: Demersal and live over both high and low relief sea floors [2, 7, 

10, 15]. 

Adults—Age and size at first maturity: Off the Kuril Islands, most fish mature by 3 years at 6.5 to 7.5 cm TL, 

although some males may mature a year earlier [10]. Males and females grow to same length and have similar 

life spans [10]. Maximum age: At least 7 years old. Maximum size: 12.0 cm TL [10]. Habitat: Demersal and live 

over both high and low relief sea floors [2, 7, 10, 15]. 

Substrate—On silty, rocky sand, gravel, and pebble bottoms [1, 4]. 

Physical/chemical—Temperature: -1.2 °C [4] to 10 °C. Salinity: Marine (or occasionally estuarine [7, 15].

Behavior

Diel—Unidentified poacher larvae have migrated into slightly deeper waters at night in southeastern Alaska [12]. 

Seasonal—Off Kamchatka, found in relatively shallow waters during summer; winters deeper on the edge of the 

continental shelf [5]. 

Reproductive—Unknown. 

Schooling—Unlikely. Other poacher species are solitary [16, 17]. 

Feeding—Can use pectoral fins to lift up rocks and shells to look for prey [14].

Populations or Stocks

There have been no studies.
360 Alaska Arctic Marine Fish Ecology Catalog

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [2]. 

Spawning season—Poorly known. Spawning occurs at least during July and August [10]. 

Fecundity—Poorly known. 57–921 eggs, spawned in a single batch [10].

Food and Feeding

Food items—In general, poachers feed on crustaceans and polychaetes [2]. Off the Kuril Islands, polychaetes, 

gammarid, and caprellid amphipods are the most important foods. However, in fishes over 10 cm (4.0 in.) long, 

shrimps, limpets, and small fishes (mainly sculpins) are of increasing importance [10]. Food habits of larvae 

are unknown. 

Trophic level—3.19 (standard error: 0.37) [18].

Biological Interactions

Predators—Cannibalism has not been reported. Predators include Pacific Cod and Whiteblotched Skates 

[19, 20]. 

Competitors—Presumably other zoobenthos feeders such as Arctic Cod, Walleye Pollock, other poachers, 

eelpouts, and sculpins.

1980 1990 2000 2010

Resilience

High, minimum population doubling time: less than 15 months (Preliminary K or Fecundity) [18].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Fourhorn Poacher are not commercially harvested.

Potential Effects of Climate Change

Fourhorn Poacher are found primarily in the southeastern Chukchi Sea. Warming, associated with climate 

change, could increase this species abundance in conjunction with a range extension to the north. Potential 

effects include changes in competition with other marine fishes that feed on small benthic organisms, including 

Arctic Cod, Walleye Pollock, other poachers, flatfishes, and sculpins.

Areas for Future Research [B]

Little is known about the ecology and life history of this species in the Chukchi Sea. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators. 
Fourhorn Poacher 361

References Cited 

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., avaliable from U.S. Department of Commerce, Springfield, Virginia] [15]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p. [2]

Tokranov, A.M., 1987b, Species composition and distribution of sea poachers (Pisces, Agonidae) in nearshore waters of 

Kamchatka: Zoologicheskii Zhurnal, v. 66, p. 385–392. [5]

Tokranov, A.M., and Orlov, A.M., 2004, Some aspects of the biology of the northern four-horned poacher Hypsagonus 

quadricornis (Agonidae) in Pacific waters off the northern Kuril Islands: Journal of Ichthyology, v. 44, no. 7, p. 508–514. [10]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Tokranov, A.M., 1987b, Species composition and distribution of sea poachers (Pisces, Agonidae) in nearshore waters of 

Kamchatka: Zoologicheskii Zhurnal, v. 66, p. 385–392.

6. Dragoo, D.E., 2006, Seabird, fish, marine mammals and oceanography coordinated investigations (SMMOCI) near Bluff, 

Norton Sound, Alaska, July 2002: U.S. Fish and Wildlife Service Report AMNWR 06/03, 35 p.

7. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

8. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p.

9. Clemens, W.G., and Wilby, G.V., 1946, Fishes of the Pacific Coast of Canada: Fisheries Research Board of Canada 

Bulletin 68, 368 p.

10. Tokranov, A.M., and Orlov, A.M., 2004, Some aspects of the biology of the northern four-horned poacher Hypsagonus 

quadricornis (Agonidae) in Pacific waters off the northern Kuril Islands: Journal of Ichthyology, v. 44, no. 7, p. 508–514.

11. Boehlert, G.W., Gadomski, D.M., and Mundy, B.C., 1985, Vertical distribution of ichthyoplankton off the Oregon coast in 

spring and summer months: Fishery Bulletin, v. 83, no. 4, p. 611–621.

12. Haldorson, L., Pritchett, M., Sterritt, D., and Watts, J., 1993, Abundance patterns of marine fish larvae during spring in a 

southeastern Alaskan bay: Fishery Bulletin, v. 91, no. 1, p. 36–44.
362 Alaska Arctic Marine Fish Ecology Catalog

13. Ambrose, D.A., Carter, R.L., and Manion, S.M., 2006, Ichthyoplankton and station data for surface (manta) and oblique 

(bongo) plankton tows for California Cooperative Oceanic Fisheries Investigations survey cruises in 2005: U.S. Department 

of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NOAA-TM-NMFSSWFSC-392, 125 p.

14. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

15. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

16. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, nos. 1–3, p. 15–28.

17. Love, M.S., University of California-Santa Barbara, pers. obs., 2009.

18. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

19. Hunter, M.A., 1979, Food resource partitioning among demersal fishes in the vicinity of Kodiak Island, Alaska: Seattle, 

University of Washington, Master’s thesis, 120 p.

20. Yang, M.-S., 2003, Food habits of the important groundfishes in the Aleutian Islands in 1994 and 1997: National Marine 

Fisheries Service, AFSC Processed Report 2003-07, 233 p.
Atlantic Poacher 363

Atlantic Poacher (Leptagonus decagonus) 

(Bloch & Schneider, 1801)

Family Agonidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Current information about occurrence is limited 

in the U.S. Chukchi and Beaufort Seas. The Atlantic Poacher is 

unlikely to represent a significant prey resource to higher-level 

organisms or role in regional food webs.

Physical Description/Attributes: A yellowish gray fish with vague grayish brown bands and patches. Pectoral and caudal fins 

are brownish black toward tips. For species diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, 

p. 537) [1]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [1, 3]. Elsewhere in Alaska, south to Bristol Bay in Bering Sea. Worldwide, nearly 

circumpolar from Laptev Sea east to western North Atlantic; in Atlantic Ocean to Newfoundland; and in Pacific Ocean there is 

an isolated population in Sea of Okhotsk [1, 4, 5]. 

Relative Abundance: Uncommon in U.S. Chukchi and Beaufort Seas [1, 3]. Elsewhere, common at least in western Barents Sea 

[8] and rare around much of Greenland [9].

Atlantic Poacher (Leptagonus decagonus), 150 mm TL, off 

northeastern Sakhalin Island, Russia, Sea of Okhotsk, 2003. 

Photograph by B.A. Sheiko, Russian Academy of Sciences.

Geographic distribution of Atlantic Poacher (Leptagonus decagonus) within Arctic Outer Continental 

Shelf Planning Areas [6] based on review of published literature and specimens from historical and recent 

collections [1, 3, 7].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Atlantic Poacher

Leptagonus decagonus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
364 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Overall range is 24–930 m, but usually between 100–200 m [1, 3]. Documented in U.S. Chukchi Sea at 44–59 m 

[1, 5]. Depth range of juveniles is unknown. Depth range of larvae is unknown. In general, abundant in near-surface waters, over 

continental shelf and shallow slope waters [10–12].

Habitats and Life History

Eggs—Size: 1.5–2.0 mm [13]. Time to hatching: Unknown. Habitat: Demersal and adhesive (probably on sand 

and mud) [2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: About 28 mm TL [14]. Days to juvenile 

transformation: 2–3 months for poachers in general [2]. Habitat: Pelagic [2], although distance from shore and 

depth in water column are unknown for this species, in general poacher larvae are abundant in near-surface 

waters, over continental shelf and shallow slope waters [10–12]. 

Juveniles—Age and size: Unknown. Habitat: Demersal [1, 5]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 22.6 cm TL [1, 15]. 

Habitat: Demersal [1, 5]. 

Substrate—Both juveniles and adults live on sand and mud [1, 5]. 

Physical/chemical—Temperature: Documented -1.7–7.4 °C [5, 13–15]. Salinity: Mainly marine but occasionally 

found at 27.4 ppt [1, 5]. In U.S. Chukchi Sea, recorded at 32.58 psu [5].

Behavior

Diel—Unidentified poacher larvae migrated into slightly deeper waters at night in southeastern Alaska [11]. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unlikely. Other poacher species are solitary [16, 17]. 

Feeding—Unknown. 

Feeding Behavior—Unknown.

Populations or Stocks

There have been no studies.

Benthic and reproductive distribution of Atlantic Poacher (Leptagonus decagonus). 

3-8-8g_AtlanticPoacher

Some fish to 930 m

Open ocean Shore

Depth, in meters 1000

800

600

400

200

0

1000

800

600

400

200

0

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Recorded at 44 m in U.S. Chukchi Sea

Overall depth data from outside 

U.S. Beaufort-Chukchi Seas

Leptagonus decagonus

Atlantic Poacher

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Juveniles and adults

Characteristic location of poacher larvae

Depth range of spawning is

uncertain. Potential location of

spawning and eggs is based on

depth range of benthic individuals 
Atlantic Poacher 365

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [2]. 

Spawning season—Poorly known. At least from May to July [13]. 

Fecundity—Poorly known. 480–1,750 eggs [13].

Food and Feeding

Food items—Food habits of larvae are unknown. Adults and juveniles feed on water column crustaceans (such 

as, copepods), benthic crustaceans, and polychaetes [13]. 

Trophic level—3.21 standard error 0.36. Based on food items [18].

Biological Interactions

Predators—Cannibalism has not been reported and is unlikely. Predators are poorly known. In the Canadian 

Arctic, they are occasionally eaten by bearded seals [19]. 

Competitors—Presumably other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other poachers, 

eelpouts, flatfish, and sculpins. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Fecundity = 1,088) [18].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Atlantic Poacher are not commercially harvested.

Potential Effects of Climate Change

This species is widely distributed in the Chukchi and Beaufort Seas, as well as southward to the eastern 

Bering Sea and eastward to the North Atlantic. Climate effects will likely result in changes in abundance and 

competition with other marine fishes. 

Areas for Future Research [B]

Little is known about the ecology and life history from the region. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults (4) spawning season (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators. 
366 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., avaliable from U.S. Department of Commerce, Springfield, Virginia] [13]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [5]

Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p. [15]

Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

5. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p.

7. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

8. Byrkjedal, I., and Høines, Å., 2007, Distribution of demersal fish in the south-western Barents Sea: Polar Research, v. 26, 

no. 2, p. 135–151.

9. Rätz, H.-J., 1999, Structures and changes of the demersal fish assemblage off Greenland, 1982–96: Northwest Atlantic 

Fisheries Organization Science Council Studies, v. 32, p. 1–15.

10. Boehlert, G.W., Gadomski, D.M., and Mundy, B.C., 1985, Vertical distribution of ichthyoplankton off the Oregon coast in 

spring and summer months: Fishery Bulletin, v. 83, no. 4, p. 611–621.

11. Haldorson, L., Pritchett, M., Sterritt, D., and Watts, J., 1993, Abundance patterns of marine fish larvae during spring in a 

southeastern Alaskan bay: Fishery Bulletin, v. 91, no. 1, p. 36–44.

12. Ambrose, D.A., Carter, R.L., and Manion, S.M., 2006, Ichthyoplankton and station data for surface (manta) and oblique 

(bongo) plankton tows for California Cooperative Oceanic Fisheries Investigations survey cruises in 2005: U.S. Department 

of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NOAA-TM-NMFSSWFSC-392, 125 p.
Atlantic Poacher 367

13. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

14. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

15. Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p.

16. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, nos. 1–3, p. 15–28.

17. Love, M.S., University of California-Santa Barbara, pers. obs., 2009.

18. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

19. Finley, K.J., and Evans, C.R., 1983, Summer diet of the bearded seal (Erignathus barbatus) in the Canadian High Arctic: 

Arctic, v. 36, no. 1, p. 82–89.
368 Alaska Arctic Marine Fish Ecology Catalog

Bering Poacher (Occella dodecaedron) 

(Tilesius, 1813)

Family Agonidae 

Note: Except for geographic range data, all information is from areas 

outside of the study area.

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Current information about this species occurrence 

in the U.S. Chukchi and Beaufort Seas is very limited. The Bering 

Poacher is unlikely to represent a significant prey resource to other fish or higher trophic level organisms. 

Physical Description/Attributes: Brownish olive on the back and sides, and light tan or ivory white on the underside. Males 

have orange pectoral fins and those of females are translucent; pelvic fin membranes in males are black and in females are white 

[1]. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 536) [2]. Swim bladder: 

Absent [3]. Antifreeze glycoproteins in blood serum: Unknown.

Range: Eastern Chukchi Sea [4]. Elsewhere in Alaska, from Port Clarence (northeastern Bering Sea near Bering Strait) [4] to 

western Gulf of Alaska [2]. Worldwide, Sea of Japan northward from Peter the Great Bay, Sea of Okhotsk, and Kuril Islands to 

Pacific Ocean off Kamchatka Peninsula, Russia, and western Bering Sea at Gulf of Anadyr [2]. 

Relative Abundance: Rare in U.S. Chukchi Sea [2, 4, 6, 7]. Elsewhere in Alaska, common in southeastern [8, 9] but rare in 

northern Bering Sea [4]. There appears to be a northward shift in the southeastern Bering Sea [10].Worldwide, common at least 

in Sea of Japan [11] and around Kamchatka Peninsula, Russia [12]. 

Bering Poacher (Occella dodecaedron), 107 mm TL, Norton 

Sound, Bering Sea, 2002. Photograph by C.W. Mecklenburg, 

Point Stephens Research.

Geographic distribution of Bering Poacher (Occella dodecaedron) within Arctic Outer Continental Shelf 

Planning Areas [5] based on review of published literature and specimens from historical and recent collections 

[4, 6].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Bering Poacher

Occella dodecaedron

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Bering Poacher 369

Depth Range: Overall range is 5–92 m, mainly less than 50 m [12–14]. A record from 375 m [13] is likely in error. Juveniles 

may live in less than 1 m [15]. Depth range of larvae unknown. In general, poacher larvae are abundant in near-surface waters, 

over continental shelf and shallow slope waters [16–18].

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic and adhesive (probably on sand and 

mud) [3]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: 2–3 months [3]. Habitat: Pelagic, although distance from shore and depth in water column are 

unknown for this species, in general, poacher larvae are abundant in near-surface waters, over continental shelf 

and shallow slope waters [16–18]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [3, 19]. 

Adults—Age and size at first maturity: Unknown. Maximum age: At least 9 years. Females are larger at age 

than males [1]. Maximum size: As long as 23 cm TL [20]. Habitat: Benthic [3, 19]. Substrate: Both juveniles and 

adults live on sand and mud [21]. 

Physical/chemical—Temperature: -1.5–15 °C [6, 12, 22]. In southeastern Bering Sea, mainly 2.2–5.5 °C [22]. 

Salinity: All life stages live in marine waters [12, 22] and occasionally estuaries [6].

Behavior

Diel—Unidentified poacher larvae have migrated into slightly deeper waters at night in southeastern Alaska [17]. 

Seasonal—Off Kamchatka Peninsula, Russia, found in relatively shallow waters during the summer; during 

winters found deeper on the edge of the continental shelf [12]. 

Reproductive—Unknown for this family [3]. 

Schooling—Unlikely. Other poacher species are solitary [19, 23]. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Benthic and reproductive distribution of Bering Poacher (Occella dodecaedron).

Some fish 

to 92 m

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

Adults

Overall benthic 

depth range 

Data from outside U.S. Beaufort-Chukchi Seas

Occella dodecaedron

Bering Poacher

Benthic distribution

Juveniles in <1 m

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

Spawning and eggs

Reproductive distribution

Characteristic location of poacher larvae

No data for this species

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 
370 Alaska Arctic Marine Fish Ecology Catalog

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [3]. 

Spawning season—Poorly known. Spawning occurs in the spring [1, 21]. 

Fecundity—400–4,575 eggs [1, 21].

Food and Feeding

Food items—In eastern Bering Sea, off Kamchatka Peninsula, Russia, and in Sea of Japan, small epibenthic 

crustaceans (for example, mysids, amphipods, isopods, and shrimps) and polychaetes [1, 20, 24]. Food habits of 

larvae are unknown. 

Trophic level—4.0 [10].

Biological Interactions

Predators—Pacific Cod and Plain Sculpin in eastern Bering Sea and off Kamchatka Peninsula, Russia. 

Cannibalism has not been reported and is unlikely [25, 26]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other 

poachers, eelpouts, and sculpins. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [27]. 

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Bering Poacher are not commercially harvested.

Potential Effects of Climate Change

Bering Poacher is a Boreal Pacific species [4] and may be expected to expand its distribution northward and 

become more abundant in the Chukchi Sea as the climate continues to warm. It already appears to be moving 

northward in the eastern Bering Sea, perhaps in response to warming water temperatures [10]. This may increase 

competition with other fishes that feed on small benthic organisms, including Arctic Cod, Walleye Pollock, other 

poachers, and sculpins.

Areas for Future Research [B]

Little is known about the biology and ecology of this species. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators.
Bering Poacher 371

References Cited 

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [21]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]

Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p. [3]

Smith, R.L., and Gillispie, J.G., 1988, Notes on the biology of the Bering Poacher, Occella dodecaedron (Tilesius), and the 

Sturgeon Poacher, Agonus acipenserinus Tilesius in the southeast Bering Sea: Copeia, v. 1988, no. 2, p. 454–460. [1]

Tokranov, A.M., 1987b, Species composition and distribution of sea poachers (Pisces, Agonidae) in nearshore waters of 

Kamchatka: Zoologicheskii Zhurnal, v. 66, p. 385–392. [12]

Bibliography

1. Smith, R.L., and Gillispie, J.G., 1988, Notes on the biology of the Bering Poacher, Occella dodecaedron (Tilesius), and the 

Sturgeon Poacher, Agonus acipenserinus Tilesius in the southeast Bering Sea: Copeia, v. 1988, no. 2, p. 454–460.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

8. Acuna, E.I., Goddard, P., and Kotwicki, S., 2003, 2002 bottom trawl survey of the eastern Bering Sea continental shelf: 

National Marine Fisheries Service, Alaska Fisheries Science Center, Processed Report 2003-01.

9. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

10. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.

11. Kolpakov, N.V., 2005, Diversity and seasonal dynamics of ichthyocenosis of the Circumlittoral of Russkaya Bight 

(Northern Primor’s): Journal of Ichthyology, v. 45, no. 9, p. 744–753.
372 Alaska Arctic Marine Fish Ecology Catalog

12. Tokranov, A.M., 1987b, Species composition and distribution of sea poachers (Pisces, Agonidae) in nearshore waters of 

Kamchatka: Zoologicheskii Zhurnal, v. 66, p. 385–392.

13. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

14. Fedorov, V.V., 2000, Species composition, distribution and habitation depths of the northern Kuril Islands fish and fish like 

species, in Koteneva, B.N., ed., Commercial and biological studies of fishes in the Pacific waters of the Kuril Islands and 

adjacent areas of the Okhotsk and Bering Seas in 1992–1998: Moscow, VNIRO Publishing, collected papers, p. 7–41. 

[In Russian.]

15. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

16. Boehlert, G.W., Gadomski, D.M., and Mundy, B.C., 1985, Vertical distribution of ichthyoplankton off the Oregon coast in 

spring and summer months: Fishery Bulletin, v. 83, no. 4, p. 611–621.

17. Haldorson, L., Pritchett, M., Sterritt, D., and Watts, J., 1993, Abundance patterns of marine fish larvae during spring in a 

southeastern Alaskan bay: Fishery Bulletin, v. 91, no. 1, p. 36–44.

18. Ambrose, D.A., Carter, R.L., and Manion, S.M., 2006, Ichthyoplankton and station data for surface (manta) and oblique 

(bongo) plankton tows for California Cooperative Oceanic Fisheries Investigations survey cruises in 2005: U.S. Department 

of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NOAA-TM-NMFSSWFSC-392, 125 p.

19. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, nos. 1–3, p. 15–28.

20. Tokranov, A.M.,1992, Dietary characteristics of sea poachers (Agonidae) in the coastal waters of Kamchatka: Journal of 

Ichthyology, v. 32, no. 9, p. 85–94.

21. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., avaliable from U.S. Department of Commerce, Springfield, Virginia]

22. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

23. Love, M.S., University of California-Santa Barbara, pers. obs., 2009.

24. Dolganova, N.T., Kolpakov, N.V., and Chuchukalo, V.I., 2006, Feeding habits and trophic relations of fish in coastal waters 

off northern Primorye: Izvestiya TINRO, v. 144, p. 140–179.

25. Tokranov, A.M., 1987a, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok

(Cuvier) (Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114.

26. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-158, 249 p.

27. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Tubenose Poacher 373

Tubenose Poacher (Pallasina barbata) 

(Steindachner, 1876)

Family Agonidae 

Note on taxonomy: The eastern Pacific form is treated by Russian 

authors as a separate species, Pallasina aix (Starks, 1896), although 

other taxonomists have given evidence for it being the same as

P. barbata [1]. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas. 

Ecological Role: Largely unknown. Current information about the occurrence of this fish in the U.S. Chukchi and Beaufort Seas 

is limited. The Tubenose Poacher is unlikely to represent a significant prey resource for many higher level organisms.

Physical Description/Attributes: An elongate, slender fish with gray to brownish back and sides and white underside. For 

specific diagnostic characteristics, see Fishes of Alaska, (Mecklenburg and others, 2002, p. 533) [1]. Swim bladder: Absent [2]. 

Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea as far northward to Point Barrow, Alaska [3]. Elsewhere in Alaska, along coast southward from 

Bering Strait [1]. Worldwide, from Sea of Japan off Korea, to Sea of Okhotsk, and Pacific Ocean off Kuril Islands south to 

Bodega Bay, central California [1, 4–8].

Relative Abundance: Rare in U.S. Chukchi Sea [3, 10, 11]. Elsewhere in Alaska, common in Bering Sea from Norton Sound 

southwards [10]. Worldwide, common in Pacific Ocean from Sea of Japan and Sea of Okhotsk, and southwards to Puget Sound 

[4–8].

Tubenose Poacher (Pallasina barbata), 138 mm TL, eastern 

Chukchi Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Tubenose Poacher (Pallasina barbata) within Arctic Outer Continental Shelf 

Planning Areas [9] based on review of published literature and specimens from historical and recent 

collections [1, 3, 10].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Tubenose Poacher

Pallasina barbata

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
374 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Intertidal zone, including tide pools, to 128 m [10]. Most abundant in shallow nearshore waters down to 60 m [4, 

8, 10, 12–14]. Depth range of juveniles is unknown. Depth range of larvae is unknown. In general, poacher larvae are abundant 

in near-surface waters, over continental shelf and upper slope [15–17]. Overall spawning depth is unknown but has been 

observed in shallow, nearshore waters [12, 13].

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown, though larvae have been caught in Barkley Sound, British 

Columbia, in April [18]. Habitat: Benthic and adhesive (probably on sand and mud) [2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 2–3 cm TL [4, 18, 19]. Days to juvenile 

transformation: 2–3 months [2]. Habitat: Pelagic, although distance from shore and depth in water column are 

unknown for this species, in general poacher larvae are abundant in near-surface waters, over continental shelf 

and shallow slope waters [15–17]. 

Juveniles—Age: Unknown. Size: 2–3 cm TL [4, 18, 19] to 9 cm TL [13]. Habitat: Demersal and shallow (often 

intertidal) [4, 10, 18, 19]. 

Adults—Age and size at first maturity: Age unknown. May mature at about 9 cm TL [13]. Maximum age: At 

least 5 years [20]. Maximum size: 20.8 cm TL [1]. Habitat: Benthic and shallow (often intertidal) [1, 4, 10, 18, 

19]. Generally, a structure-oriented fish [12, 21]. Common in eelgrass beds, although they also inhabit kelp 

stands, rocky outcrops, and shell hash sea floors [8, 11–13]. 

Substrate—Shell hash, sand and gravel bottoms [1, 11]. 

Physical/chemical—Temperature: Documented off Kamchatka Peninsula, Russia, at 0–12 °C [14], and in 

Bering Strait at 10.5 °C [11]. Salinity: Marine and estuarine [14], found at 30.62 salinity units in U.S. Bering 

Strait [11].

Behavior

Diel—Unidentified poacher larvae migrated into slightly deeper waters at night in southeastern Alaska [16]. 

Seasonal—Off Kamchatka Peninsula, Russia, found in relatively shallow waters during summer and during 

winter in deeper waters on the edge of continental shelf [14]. 

Reproductive—During summer months, adults move into shallow waters to spawn [12, 13]. 

Schooling—Unlikely. Other poacher species are solitary [22, 23]. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Some fish 

to 128 m

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

Juveniles and adults

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

Pallasina barbata

Tubenose Poacher

Benthic distribution

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

Reproductive distribution

Data from outside U.S. Beaufort-Chukchi Seas

Characteristic location of poacher larvae

No data for this species

Spawning and eggs

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Benthic and reproductive distribution of Tubenose Poacher (Pallasina barbata).
Tubenose Poacher 375

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [2]. 

Spawning season—Spawning occurs during the summer [12, 13]. 

Fecundity—Unknown. 

Food and Feeding

Food items—Small benthic and epibenthic crustaceans dominate the diet. Mysids are important, as are 

euphausiids, caprellid amphipods, and copepods. Polychaetes and shrimps also are consumed [5, 12, 13, 20, 24]. 

Food habits of larvae are unknown. 

Trophic level—3.23 (standard error 0.41) [25].

Biological Interactions

Predators—Cannibalism has not been reported. Off Kamchatka Peninsula, this species is eaten by great sculpin 

[26]. 

Competitors—Presumably a wide range of other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other 

poachers, eelpouts, and sculpins. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [25].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Tubenose Poacher are not commercially harvested.

Potential Effects of Climate Change

The expected result of climate warming would be an increased abundance of this species in the Chukchi Sea. 

An eventually expansion of range into the Beaufort Sea also would be expected. These changes would result in 

increased interactions and competition with other fishes that feed on small benthic organisms. Major competitors 

would potentially include: Arctic Cod, Walleye Pollock, other poachers, and sculpins.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.
376 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., avaliable from U.S. Department of Commerce, Springfield, Virginia] [12] 

Marliave, J.B., 1975, The behavioral transformation from the planktonic larval stage of some marine fishes reared in the 

laboratory: Vancouver, University of British Columbia, Ph.D. dissertation. [18]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p. [2]

Tack, S.L., 1970, The summer distribution and standing stock of the fishes of Izembek Lagoon, Alaska: Fairbanks, University of 

Alaska, Master’s thesis. [13]

Tokranov, A.M., 1987b, Species composition and distribution of sea poachers (Pisces, Agonidae) in nearshore waters of 

Kamchatka: Zoologicheskii Zhurnal, v. 66, p. 385–392. [14]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p.

3. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

4. Barton, L.H., 1978, Finfish resource surveys in Norton Sound and Kotzebue Sound: Alaska Department of Fish and Game, 

Commercial Fisheries Division, p. 75–313.

5. Cross, J.N., Fresh, K., Miller, B.S., Simenstad, C.A., Steinfort, S.N., and Fegley, J.C., 1978, Nearshore fish and 

invertebrates assemblages along the Strait of Juan de Fuca including food habits of the common inshore fishes: U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, Environmental Research Laboratories, 

NOAA Technical Memorandum ERL MESA-32.

6. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

7. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

8. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

9. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 
Tubenose Poacher 377

10. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

11. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

12. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., avaliable from U.S. Department of Commerce, Springfield, Virginia]

13. Tack, S.L., 1970, The summer distribution and standing stock of the fishes of Izembek Lagoon, Alaska: Fairbanks, 

University of Alaska, Master’s thesis.

14. Tokranov, A.M., 1987b, Species composition and distribution of sea poachers (Pisces, Agonidae) in nearshore waters of 

Kamchatka: Zoologicheskii Zhurnal, v. 66, p. 385–392.

15. Boehlert, G.W., Gadomski, D.M., and Mundy, B.C., 1985, Vertical distribution of ichthyoplankton off the Oregon coast in 

spring and summer months: Fishery Bulletin, v. 83, no. 4, p. 611–621.

16. Haldorson, L., Pritchett, M., Sterritt, D., and Watts, J., 1993, Abundance patterns of marine fish larvae during spring in a 

southeastern Alaskan bay: Fishery Bulletin, v. 91, no. 1, p. 36–44.

17. Ambrose, D.A., Carter, R.L., and Manion, S.M., 2006, Ichthyoplankton and station data for surface (manta) and oblique 

(bongo) plankton tows for California Cooperative Oceanic Fisheries Investigations survey cruises in 2005: U.S. Department 

of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NOAA-TM-NMFSSWFSC-392, 125 p.

18. Marliave, J.B., 1975, The behavioral transformation from the planktonic larval stage of some marine fishes reared in the 

laboratory: Vancouver, University of British Columbia, Ph.D. dissertation.

19. Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, Fisheries Research Institute, FRI-UW-7719.

20. Tokranov, A.M., 1992, Dietary characteristics of sea poachers (Agonidae) in the coastal waters of Kamchatka: Journal of 

Ichthyology v. 32, no. 9, p. 85–94.

21. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

22. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, nos. 1–3, p. 15–28.

23. Love, M.S., University of California-Santa Barbara, pers. obs., 2009.

24. McConnaughey, T., and McRoy, C.P., 1979, 13C label identifies eelgrass (Zostera marina) carbon in an Alaskan estuarine 

food web: Marine Biology, v. 53, no. 3, p. 263–269.

25. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

26. Tokranov, A.M., 1987a, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok

(Cuvier)(Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114.
378 Alaska Arctic Marine Fish Ecology Catalog

Veteran Poacher (Podothecus veternus) 

Jordan & Starks, 1895

Family Agonidae 

Note on taxonomy: Because of historical confusion of P. veternus 

with P. accipenserinus, geographic and bathymetric ranges for the 

species are not well known [1].

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Largely unknown. Current information about 

the occurrence of this fish is limited but the species is common in 

coastal waters near Point Barrow. Species information regarding the biology and ecology is not available. The veteran poacher is 

unlikely to represent a significant prey resource to higher-level organisms but may be locally and seasonally important.

Physical Description/Attributes: Brown back and sides marked by dark vertical bands, white on chest and belly. For specific 

diagnostic characteristics, see Fishes of Alaska (Meckenburg and others, 2002, p. 542) [1]. Swim bladder: Absent [2]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: Eastern Chukchi Sea northward to 71°23’N, 160°15’W [3] eastward to western Beaufort Sea at Cooper Island [4]. 

One unconfirmed record from the U.S. Beaufort Sea [3]. Elsewhere in Alaska, to eastern Bering Sea at Norton Sound and near 

St. Matthew Island. Worldwide, Seas of Okhotsk and Japan [1].

Relative Abundance: Common in U.S. Chukchi Sea [3]. Elsewhere in Alaska, common in northern Bering Sea, absent from 

southern Bering Sea [1]. Rare in Sea of Japan [7].

Veteran Poacher (Podothecus veternus), 172 mm TL, 

northeastern Bering Sea, 2006. Photograph by C.W. 

Mecklenburg, Point Stephens Research.

Geographic distribution of Veteran Poacher (Podothecus veternus) within Arctic Outer Continental Shelf Planning 

Areas [5] based on review of published literature and specimens from historical and recent collections [3, 6].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Veteran Poacher

Podothecus veternus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Veteran Poacher 379

Depth Range: Documented in U.S. Chukchi 41–48 m [8]. Elsewhere, overall range is intertidal to 240 m [6]. Depth range of 

juveniles is unknown. Depth range of larvae is unknown. In general, poacher larvae are abundant in near-surface waters, over 

continental shelf and shallow slope waters [9–11].

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Demersal and adhesive (probably on sand and 

mud) [2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: For poachers in general, 2–3 months [2]. Habitat: Pelagic. Although distance from shore and 

depth in water column are unknown for this species, in general poacher, larvae are abundant in near-surface 

waters, over continental shelf and shallow slope waters [9–11]. 

Juveniles—Age and size: Unknown. Habitat: Demersal [2]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 28.5 cm TL [1]. 

Habitat: Demersal [2]. 

Substrate—Shell hash and mud [8]. 

Physical/chemical—Temperature: -1.8–10.5 °C [8, 12] Salinity: All life stages live in marine waters [2].

Behavior

Diel—Unidentified poacher larvae have migrated into slightly deeper waters at night in southeastern Alaska [10]. 

Seasonal—Unknown. 

Reproductive—Unknown for this family [2]. 

Schooling—Unlikely. Other poacher species are solitary [13, 14]. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [2]. 

Spawning season—Unknown. 

Fecundity—Unknown. 

Benthic and reproductive distribution of Veteran Poacher (Podothecus veternus).

Open ocean Shore

Depth, in meters

250

200

150

100

50

0

250

200

150

100

50

0

Juveniles 

and adults

Podothecus veternus

Veteran Poacher

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs 

Depth range documented in

U.S. Chukchi and Beaufort Seas

Reproductive distribution

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

Characteristic location of poacher larvae

Overall depth data from outside U.S. Beaufort-Chukchi Seas

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 
380 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Unknown. 

Trophic level—3.31 standard error 0.46. Based on size and trophics of closest relatives [15].

Biological Interactions

Predators—Unknown. 

Competitors—Presumably other zoobenthic feeders such as Arctic Cod, Walleye Pollock, other poachers, 

eelpouts, and sculpins. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [15].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Veteran Poacher are not commercially harvested.

Potential Effects of Climate Change

Climate change would likely increase the abundance of this species in the study area. This may increase 

competition with other fishes that feed on small benthic organisms, including Arctic Cod, Walleye Pollock, other 

poachers, and sculpins.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators. The veteran poacher may be an important prey of migratory birds and 

mammals in the ice lead along northwest Alaska during spring.
Veteran Poacher 381

References Cited

Haldorson, L., Pritchett, M., Sterritt, D., and Watts, J., 1993, Abundance patterns of marine fish larvae during spring in a 

southeastern Alaskan bay: Fishery Bulletin, v. 91, no. 1, p. 36–44. [10]

Johnson, S.W., Thedinga, J.F., Neff, A.D., and Hoffman, C.A., 2010, Fish fauna in nearshore waters of a Barrier Island in the 

western Beaufort Sea, Alaska: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, Technical 

Memorandum, NMFSAFSC-210, 28 p. [4]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3] 

Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes no. 30, 27 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Johnson, S.W., Thedinga, J.F., Neff, A.D., and Hoffman, C.A., 2010, Fish fauna in nearshore waters of a Barrier Island 

in the western Beaufort Sea, Alaska: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, 

Technical Memorandum, NMFSAFSC-210, 28 p. [4]

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p.

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

8. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

9. Boehlert, G.W., Gadomski, D.M., and Mundy, B.C., 1985, Vertical distribution of ichthyoplankton off the Oregon coast in 

spring and summer months: Fishery Bulletin, v. 83, no. 4, p. 611–621.

10. Haldorson, L., Pritchett, M., Sterritt, D., and Watts, J., 1993, Abundance patterns of marine fish larvae during spring in a 

southeastern Alaskan bay: Fishery Bulletin, v. 91, no. 1, p. 36–44.

11. Ambrose, D.A., Carter, R.L., and Manion, S.M., 2006, Ichthyoplankton and station data for surface (manta) and oblique 

(bongo) plankton tows for California Cooperative Oceanic Fisheries Investigations survey cruises in 2005: U.S. Department 

of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NOAA-TM-NMFSSWFSC-392, 125 p.
382 Alaska Arctic Marine Fish Ecology Catalog

12. Cui, X., Grebmeier, J.M., Cooper, L.W., Lovvorn, J.R., North, C.A., Seaver, W.L., and Kolts, J.M., 2009, Spatial 

distributions of groundfish in the northern Bering Sea in relation to environmental variation: Marine Ecology Progress 

Series, v. 393, p. 147–160.

13. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, nos. 1–3, p. 15–28.

14. Love, M.S., University of California-Santa Barbara, pers. obs., 2009.

15. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Pimpled Lumpsucker 383

Pimpled Lumpsucker (Eumicrotremus andriashevi) 

Perminov, 1936

Family Cyclopteridae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: A rare species that likely is of little ecological 

significance in the U.S. Chukchi Sea.

Physical Description/Attributes: Globose body with short tails 

and a disk on the underside, formed by modified pelvic fins, for 

clinging to rocks and other objects. Head and body covered in 

spiny tubercles. Color in life not reported. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, 

p. 568) [1]. Swim bladder: Absent [2]. Antifreeze glycoproteins in 

blood serum: Unknown.

Range: U.S. Chukchi Sea. Elsewhere in Alaska, eastern Bering Sea as far southward as St. Matthew Island. Worldwide, western 

Bering Sea southward to Karaginskiy Bay [3]. 

Relative Abundance: Uncommon in U.S. Chukchi Sea [1, 5, 6].

Pimpled Lumpsucker (Eumicrotremus andriashevi), 56 mm 

TL, Chukchi Sea, 1991 (preserved specimen, University of 

Alaska Museum of the North 4592). Photograph by C.W. 

Mecklenburg, Point Stephens Research.

Geographic distribution of Pimpled Lumpsucker (Eumicrotremus andriashevi) within Arctic Outer Continental 

Shelf Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [1, 3, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Pimpled Lumpsucker

Eumicrotremus andriashevi

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
384 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Benthic, at depths of 20–93 m [5]. Specific spawning depth unknown. In general, most lumpsuckers spawn in 

shallow coastal waters [2].

 Benthic and reproductive distribution of Pimpled Lumpsucker (Eumicrotremus andriashevi).

3-8-12g_PimpledLumpsucker

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Juveniles and adults

Some fish

to 83 m

Eumicrotremus andriashevi

Pimpled Lumpsucker

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning, eggs, and larvae

Reproductive distribution

Overall benthic depth range.

Specific depth range of either 

juveniles or adults is unknown 

Depth range of spawning is uncertain. 

Potential locations of spawning, eggs and larvae

are based on depth range of benthic individuals

Data from outside U.S. Beaufort-Chukchi Seas

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [1, 2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Likely benthic [2]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [2]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: As long as 9.7 cm 

TL [1]. Habitat: Benthic [1, 2]. 

Substrate—Mud, sand, and pebble bottoms [1]. 

Physical/chemical—Temperature: -1.6–2.8 °C [5]. Salinity: Unknown. 

Behavior

Unknown for pimpled lumpsucker. All behavior given is for lumpsuckers in general. 

Diel—Known to inflate their bodies by swallowing air or water in what is likely a defensive reaction. 

Seasonal—Unknown. 

Reproductive—After spawning male lumpsuckers guard the eggs [2]. 

Schooling—Unknown. 

Feeding—Feeds on slow moving prey [2].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous. 

Spawning season—Unknown. 

Fecundity—Unknown. 
Pimpled Lumpsucker 385

Food and Feeding

Food items—Unknown. 

Trophic level—3.15 standard error 0.31 [7].

Biological Interactions

Predators—Unknown. 

Competitors—Likely sculpins, poachers, and flatfishes.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (assuming Fecundity <1,000) [7].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Pimpled Lumpsucker are not commercially harvested.

Potential Effects of Climate Change

Unknown.

Areas for Future Research [B]

Little is known about the ecology of this species. Research needs include: (1) depth and location of pelagic 

larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth ranges for 

juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population studies, 

(7) prey, and (8) predators.
386 Alaska Arctic Marine Fish Ecology Catalog

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., and Sheiko, B.A., 2003, Family Cyclopteridae Bonaparte 1831—Lumpsuckers: California Academy of 

Sciences Annotated Checklists of Fishes no. 6, 17 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

7. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Leatherfin Lumpsucker 387

Leatherfin Lumpsucker (Eumicrotremus derjugini) 

Popov, 1926 

Family Cyclopteridae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: An uncommon species in the U.S. Beaufort Sea. 

Its ecological significance is not known but believed to be of minor 

consequence.

Physical Description/Attributes: Globose body with short 

tail and an adhesive disk on the underside, formed by modified 

pelvic fins, for clinging to rocks and other objects. Head and 

body covered in spiny tubercles. Colored blackish brown on 

head, back, and sides fading to nearly white on belly. For specific 

diagnostic characteristics, see Fishes of Alaska, (Mecklenburg and 

others, 2002, p. 566) [1]. Swim bladder: Absent [2]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: Eastern Chukchi Sea and Alaskan Beaufort Sea [5]. Worldwide, circumpolar in Arctic seas; northward in Barents Sea to 

81–82°N; isolated population in northern Sea of Okhotsk [3].

Relative Abundance: Common in U.S. Beaufort Sea, not reported from U.S. Chukchi Sea [1, 5]. Common off northwest and 

northeast Greenland [6].

Leatherfin Lumpsucker (Eumicrotremus derjugini), 101 mm 

TL, Beaufort Sea, 2011. Photograph by C.W. Mecklenburg, 

Point Stephens Research.

Geographic distribution of Leather Lumpsucker (Eumicrotremus derjugini), within Arctic Outer Continental 

Shelf Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [1, 3, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Leatherfin Lumpsucker

Eumicrotremus derjugini

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
388 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 50–930 m, typically less than 275 m [1]. Documented in U.S. Beaufort Sea from 50–110 m [7]. A juvenile was 

taken at 41 m in U.S. Beaufort Sea [3]. Specific spawning depth unknown. In general, most lumpsuckers spawn in shallow 

coastal waters [2].

Habitats and Life History

Eggs—Size: 4.0–5.0 mm [8]. Females can contain two size classes of eggs at one time [7]. Time to hatching: 

Unknown. In Barents Sea, eggs laid in autumn hatch the following summer [9]. Habitat: Benthic [1, 2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Unknown, likely benthic [2]. 

Juveniles—Age and size: Unknown. One juvenile from U.S. Beaufort Sea was 10.1 mm SL, 14.2 mm TL [3]. 

Habitat: Benthic [2]. In Barents Sea, they remain in coastal areas until about 1 year old and up to 3–4 cm [2, 9]. 

Adults—Age and size at first maturity: Unknown. Over 6.5 cm TL for females in U.S. Beaufort Sea [7]. 

Maximum age: Unknown. Maximum size: 12.7 cm TL [1]. Habitat: Benthic [2, 10, 11]. Substrate—Mud, gravel, 

stony bottoms [1, 10]. 

Physical/chemical—Temperature: Poorly documented; -2–0 °F [11]. Salinity: Marine [10]. 

Behavior

Diel—Unknown. In general, lumpsuckers are known to inflate their bodies by swallowing air or water in what is 

likely a defensive reaction. 

Seasonal—Unknown. 

Reproductive—Unknown. In general, after spawning male lumpsuckers guard the eggs [2]. 

Schooling—Unknown. 

Feeding—Feeds on slow moving prey [2].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous. 

Spawning season—Unknown. Autumn in Barents Sea [9]. Females taken off Russia during August and 

September contained large eggs [10]. 

Fecundity—Unknown.

Benthic and reproductive distribution of Leatherfin Lumpsucker (Eumicrotremus derjugini).

3-8-13g_LeatherfinLumpsucker

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

1,000

800

600

400

200

0

Adults and juveniles 

Some fish to 930 m

Eumicrotremus derjugini

Leatherfin Lumpsucker

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning, eggs, and larvae

Reproductive distribution

Depths documented in U.S. 

Beaufort-Chukchi Seas

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Depth range of spawning is uncertain. 

Potential locations of spawning, eggs 

and larvae are based on depth range 

of benthic individuals

Overall data from outside 

U.S. Beaufort-Chukchi Seas
Leatherfin Lumpsucker 389

Food and Feeding

Food items—In U.S. Beaufort Sea, primarily hyperiid amphipods and gammarid amphipods, mysids, and 

polychaetes [7]. 

Trophic level—3.25 standard error 0.34 [12].

Biological Interactions

Predators—Thick-billed Murres in the Canadian Arctic [13]. 

Competitors—Likely sculpins, poachers, and flatfishes.

1980 1990 2000 2010

Resilience

High, minimum population doubling time is less than 15 months (Preliminary K or Fecundity) [12].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Leatherfin Lumpsucker are not commercially harvested.

Potential Effects of Climate Change

Unknown.

Areas for Future Research [B]

Little is known about the ecology of this species. Research needs include: (1) depth and location of pelagic 

larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth ranges for 

juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population studies, 

(7) prey, and (8) predators.
390 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [10]

Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western Beaufort 

Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA Technical Report 

NMFS-SSRF-764, 22 p. [7]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Mecklenburg, C.W., and Sheiko, B.A., 2003, Family Cyclopteridae Bonaparte 1831—Lumpsuckers: California Academy of 

Sciences Annotated Checklists of Fishes no. 6, 17 p. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., and Sheiko, B.A., 2003, Family Cyclopteridae Bonaparte 1831—Lumpsuckers: California Academy of 

Sciences Annotated Checklists of Fishes no. 6, 17 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Møller, P.R., Nielsen, J.G., Knudsen, S.W., Poulsen, J.Y., Sünksen, K., and Jørgensen, O.A., 2010, A checklist of the fish 

fauna of Greenland waters: Zootaxa, v. 2378, p. 1–84.

7. Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western 

Beaufort Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA 

Technical Report NMFS-SSRF-764, 22 p.

8. Able, K.W., Markle, D.F., and Fahay, M.P., 1984, Cyclopteridae—Development, in Moser, H.G. and others, eds., Ontogeny 

and systematics of fishes: Lawrence, Kansas, Allen Press, American Society of Ichthyology and Herpetology Special 

Publication 1, p. 428–437.

9. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

10. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., avaliable from U.S. Department of Commerce, Springfield, Virginia]

11. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

12. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

13. Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p.
Sea Tadpole 391

Sea Tadpole to Polar Eelpout

Sea Tadpole (Careproctus reinhardti) 

(Krøyer, 1862)

Family Liparidae

Note: Except for geographic range data, all information is from 

areas outside of the study area.

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown, as its abundance in the Arctic is poorly 

understood.

Physical Description/Attributes: A gelatinous, pink-translucent 

snailfish with a small, recessed, pelvic adhesive disk. For diagnostic 

details see Nozères and others (2010, p. 88) [1]. Swim bladder: 

Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: Chukchi Borderland and upper continental slope of Chukchi and Beaufort Seas (to as far eastward as 71°15’N, 

150°06’W) [5]. Worldwide, found in Arctic Ocean north of Alaska, in Baffin Bay and Davis Strait, off eastern Greenland, and 

Kara and Laptev Seas [5].

Sea Tadpole (Careproctus reinhardti), 170 mm TL, Chukchi 

Borderland, 2009. Photograph by C.W. Mecklenburg, Point 

Stephens Research.
392 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_fig3-9-1m_Careproctus_reinhardti

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Sea Tadpole

Careproctus reinhardti

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Sea Tadpole (Careproctus reinhardti) within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [3, 5]. 

Relative Abundance: Not documented in U.S. Chukchi Sea. Rare in U.S. Beaufort Sea [3]. Has been reported to be abundant in 

Gulf of St. Lawrence [1, 6].
Sea Tadpole 393

Depth Range: 100–1,840 m [3], usually 180–350 m [7]. Taken in U.S. Beaufort Sea at 183–223 m, and north of 

U.S. Chukchi Sea, outside the 200-mile limit, at 227–236 m [3, 5]. 

Benthic and reproductive distribution for Sea Tadpole (Careproctus reinhardti).

3-9-1g_SeaTadpole

Open ocean

Depth, in meters

2,000

1,600

1,200

800

400

0

Adults and juveniles

Careproctus reinhardti

Sea Tadpole

Benthic distribution

Some fish to 1,840 m

Overall benthic depth range. 

Depth, in meters

Specific depth range of either 

juveniles and adults is known

Open ocean No data for this species

Depths documented in U.S.

Beaufort-Chukchi Seas

Shore

Spawning and eggs

Reproductive distribution

Depth range of spawning is uncertain. 

Potential location of spawning and eggs is

based on depth range of benthic individuals

No data for larvae

Overall depth data from 

outside U.S. Beaufort-Chukchi Seas 2,000

1,600

400

800

1,200

0

Habitats and Life History

Eggs—Size: 4.2–4.7 mm [7, 8]. Time to hatching: Unknown. Habitat: Unknown. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Unknown. 

Juveniles—Age and size: Unknown. Habitat: Unknown. 

Adults—Age and size at first maturity: Unknown. About 120 mm TL in Atlantic Ocean [6]. Maximum age: 7 

years [9]. Maximum size: 30 cm TL [10]. Habitat: Benthic and benthopelagic [7]. 

Substrate—Muddy bottoms [7]. Physical/chemical—Temperature: -2 to 4 °C [5, 7]. Spawning at 2–4 °C [11]. 

Salinity: Marine [7].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous [12]. 

Spawning season—Perhaps in spring and (or) summer in Canadian Arctic [13], in November in Barents Sea [8]. 

Fecundity—About 40–300 eggs [6, 7].

Food and Feeding

Food items—Smaller fish consumed more hyperiid and gammarid amphipods; larger fish consumed more 

pandalid shrimps. In general, amphipods and shrimps are major prey. Fishes and copepods also are reported in 

the diet [7, 10]. Trophic level—3.75 standard error 0.58 [14].
394 Alaska Arctic Marine Fish Ecology Catalog

Biological Interactions

Predators—Ribbon seals eat snailfishes in the Bering Sea [16]. 

Competitors—Likely a range of benthic fish species, including various flatfishes, sculpins, poachers, and 

eelpouts.

1980 1990 2000 2010

Resilience

High, minimum population doubling time: less than 15 months (Preliminary K or Fecundity) [14].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Sea Tadpole are not commercially harvested.

Potential Effects of Climate Change

The Sea Tadpole are an Arctic species and its distribution would be expected to shift northward with warming 

temperatures. 

Areas for Future Research [B]

Little information is available regarding the biology and ecology of this species in the U.S. Arctic marine 

environment. Research needs include: (1) depth and location of larvae, (2) depth, location, and timing of youngof-the-year benthic recruitment, (3) preferred depth ranges for juveniles and adults, (4) spawning season, 

(5) seasonal and ontogenetic movements, (6) population studies, (7) prey, and (8) predators. 

Remarks

This species may be a species complex and genetic confirmation of phylogenetic relationships is needed [3].

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [7]

Chernova, N.V., 1991, Aquarium observations of the snail fishes Careproctus reinhardti and Liparis liparis: Journal of 

Ichthyology, v. 31, no. 9, p. 65–70. [8]

Falk-Petersen, I.-B., Frivoll, V., Gulliksen, B., Haug, T., and Vader, W., 1988, Age/size relations and food of two snailfishes, 

Liparis gibbus and Careproctus reinhardtii (Teleostei, Liparididae) from Spitsbergen coastal waters: Polar Biology, v. 8, no. 5, 

p. 353–358. [10]
Sea Tadpole 395

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Nozères, C., Archambault, D., Chouinard, P.-M., Gauthier, J., Miller, R., Parent, E., Schwab, P., Savard, L., and Dutil, J.-D., 

2010, Identification guide for marine fishes of the estuary and northern Gulf of St. Lawrence and sampling protocols used 

during trawl surveys between 2004 and 2008: Canadian Technical Report of Fisheries and Aquatic Sciences no. 2866. [1]

Bibliography

1. Nozères, C., Archambault, D., Chouinard, P.-M., Gauthier, J., Miller, R., Parent, E., Schwab, P., Savard, L., and Dutil, J.-D., 

2010, Identification guide for marine fishes of the estuary and northern Gulf of St. Lawrence and sampling protocols used 

during trawl surveys between 2004 and 2008: Canadian Technical Report of Fisheries and Aquatic Sciences no. 2866.

2. Eastman, J.T., Hikida, R.S., and DeVries, A.L., 1994, Buoyancy studies and microscopy of skin and subdermal extracellular 

matrix of the Antarctic snailfish, Paraliparis devriesi: Journal of Morphology, v. 220, no. 1, p. 85–101.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Able, K.W., and Irion, W., 1985, Distribution and reproductive seasonality of snailfishes and lumpfishes in 

the St. Lawrence River estuary and the Gulf of St. Lawrence: Canadian Journal of Zoology, v. 63, no. 7, 

p. 1,622–1,628.

7. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

8. Chernova, N.V., 1991, Aquarium observations of the snail fishes Careproctus reinhardti and Liparis liparis: Journal of 

Ichthyology, v. 31, no. 9, p. 65–70.

9. Falk-Petersen, I.-B., Frivoll, V., Gulliksen, B., Haug, T., and Vader, W., 1988, Age/size relations and food of two snailfishes, 

Liparis gibbus and Careproctus reinhardtii (Teleostei, Liparididae) from Spitsbergen coastal waters: Polar Biology, v. 8, 

no. 5, p. 353–358.

11. Muus, B.J., Nielsen, J.G., Dahlstrom, P., and Nystrom, B.O., 1999, Sea fish: Hedehusene, Denmark, Blackwell Science 

Limited, 340 p.

12. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

13. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

14. Able, K.W., and McAllister, D.E., 1980, Revision of the snailfish genus Liparis from Arctic Canada: Ottawa, Department of 

Fisheries and Oceans, Canadian Bulletin of Fisheries and Aquatic Sciences 208.

15. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

16. Frost, K.J., and Lowry, L.F., 1980, Feeding of ribbon seals (Phoca fasciata) in the Bering Sea in spring: Canadian Journal 

of Zoology, v. 58, no. 9, p. 1,601–1,607.
396 Alaska Arctic Marine Fish Ecology Catalog

Nebulous Snailfish (Liparis bathyarcticus) 

Parr, 1931

Family Liparidae 

Note on taxonomy: Historical confusion with L. gibbus (Bean, 

1881) makes distribution uncertain [1].

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: The ecological significance of this species in 

benthic food webs is unknown, but given its widespread distribution 

and abundance could be regionally important.

Physical Description/Attributes: Head and body with brown 

mottling and metallic gold sheen; belly and underside of head white; and dorsal, anal, and caudal fins with distinct or indistinct 

black bands [2]. For specific diagnostic characteristics, see Parr (1931) and Chernova (2008) [3, 4]. Swim bladder: Absent [5]. 

Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [1]. Elsewhere in Alaska, to southeastern Bering Sea [3]. Worldwide, predominantly 

Arctic, circumpolar from west Greenland to Labrador, Greenland, Norwegian, Barents, White, and Kara Seas, and East Siberian 

and Chukchi Seas [1].

Relative Abundance: Common in U.S. Chukchi and Beaufort Seas but historically confused with L. gibbus and other Liparis 

species [2, 7].

Nebulous Snailfish (Liparis bathyarcticus), 184 mm, Bering 

Strait, 2009. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Nebulous Snailfish (Liparis bathyarcticus) within Arctic Outer Continental Shelf 

Planning Areas [6], based on review of published literature and specimens from historical and recent 

collections [1, 2]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Nebulous Snailfish

Liparis bathyarcticus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Nebulous Snailfish 397

Depth Range: 8–510 m [2], typically 50–300 m [1].

Benthic and reproductive distribution of Nebulous Snailfish (Liparis bathyarcticus).

3-9-2g_ArcticSnailfish

Open ocean Shore

Depth, in meters 1000

800

600

400

200

0

1000

800

600

400

200

0

Adults and juveniles

Some fish to 647 m

Liparis bathyarcticus

Nebulous Snailfish

Benthic distribution

Open ocean

No data for larvae

Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Depth range of spawning is uncertain. 

Potential location of eggs is based on 

depth range of benthic individuals

Data from outside U.S. Beaufort-Chukchi Seas

No data for this species

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Likely benthic, like other Liparis species [1, 9]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Likely pelagic, like other Liparis species [1, 9]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1, 9]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 27 cm [8]. Habitat: 

Benthic [1, 9]. 

Substrate—Unknown. 

Physical/chemical—Temperature: -1.6–3.7 °C [2]. Salinity: Marine [2]. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous [10]. 

Spawning season—Unknown. 

Fecundity—Unknown. 

Food and Feeding

Food items—Plankton and benthic crustaceans (euphausiids, shrimps, gammarids) for juveniles. Fishes and 

large decapods for adults [8]. 

Trophic level—3.29 standard error 0.46 (based on information for L. gibbus) [11].
398 Alaska Arctic Marine Fish Ecology Catalog

Biological Interactions

Predators—Snailfishes outside U.S. Chukchi and Beaufort Seas eaten by such predators as larger fishes and 

seals [12–15]. 

Competitors—Likely a range of benthic fish species, including various flatfishes, sculpins, poachers, and 

eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Fecundity=4,000-12,000) (based on information for 

L. gibbus) [11].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Nebulous Snailfish are not commercially harvested.

Potential Effects of Climate Change

As the Nebulous Snailfish are a predominantly Arctic species [1], a northward shift is distribution due to climate 

warming would be expected. 

Areas for Future Research [B]

Little is known about the ecology and life history of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators. 

References Cited

Chernova, N.V., 2008, Systematics and phylogeny of fish of the genus Liparis (Liparidae, Scorpaeniformes): Journal of 

Ichthyology, v. 48, no. 10, p. 831–852. [3]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [1]

Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, A., 

Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 2011, 

Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p. [8]
Nebulous Snailfish 399

Bibliography

1. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

2. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

3. Chernova, N.V., 2008, Systematics and phylogeny of fish of the genus Liparis (Liparidae, Scorpaeniformes): Journal of 

Ichthyology, v. 48, no. 10, p. 831–852. 

4. Parr, A.E., 1931 A study of subspecies and racial variations in Liparis liparis and Liparis koefoedi, n. sp., in northern 

Europe and the European Arctic waters: Bergens Museums Bibliothek, v. 6, p. 1–53.

5. Eastman, J.T., Hikida, R.S., and DeVries, A.L., 1994, Buoyancy studies and microscopy of skin and subdermal extracellular 

matrix of the Antarctic snailfish, Paraliparis devriesi: Journal of Morphology, v. 220, no. 1, p. 85–101.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

8. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

9. Able, K.W., and McAllister, D.E., 1980, Revision of the snailfish genus Liparis from Arctic Canada: Ottawa, Department of 

Fisheries and Oceans, Canadian Bulletin of Fisheries and Aquatic Sciences 208.

10. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

11. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

12. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.

13. Wakefield, W.W., 1984, Feeding relationships within assemblages of nearshore and mid-continental shelf benthic fishes off 

Oregon: Corvallis, Oregon State University, Master’s thesis, 102 p.

14. Frost, K.J., and Lowry, L.F., 1980, Feeding of ribbon seals (Phoca fasciata) in the Bering Sea in spring: Canadian Journal 

of Zoology, v. 58, no. 9, p. 1,601–1,607.

15. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990.
400 Alaska Arctic Marine Fish Ecology Catalog

Gelatinous Seasnail (Liparis fabricii) 

Krøyer, 1847

Family Liparidae 

Note: Liparis fabricii was proposed to be a species complex [1], but 

molecular analysis of specimens caught in the U.S. Chukchi and 

Beaufort Seas to Davis Strait in the Atlantic indicates they are the 

same species [2].

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown, as its abundance in the Arctic is poorly 

understood.

Physical Description/Attributes: Tadpole shaped, soft and 

covered with gelatinous tissue. Head and body uniformly brown, 

gray, or black with a black peritoneum. For specific diagnostic characteristics, see Fishes of Alaska (Meckenburg and others, 

2002, p. 592) [3]. Swim bladder: Absent [4]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas, as far north as the continental slope of the Chukchi Sea outside the U.S. 200-mile 

limit [2]. Elsewhere in Alaska, southward in Bering Sea to a point south of Bering Strait west of Cape Rodney and well north 

of St. Lawrence Island at 64°30’N, 170°26’W. Worldwide, circumpolar in the Arctic, northward to northern Ellesmere Island, 

southward to Georges Bank, Canada, and Iceland [2].

Relative Abundance: Common, usually caught at one to several stations offshore on research cruises in the U.S. Chukchi and 

Beaufort Seas but not in great numbers [2, 3, 7]; relatively large catches have been made well offshore of Point Barrow [8] as 

well as off Herschel Island, Canada [9].

Gelatinous Seasnail (Liparis fabricii), 94 mm TL, western 

Chukchi Sea, 2004. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Gelatinous Seasnail

Liparis fabricii

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Gelatinous Seasnail (Liparis fabricii) within Arctic Outer Continental Shelf Planning 

Areas [5] based on review of published literature and specimens from historical and recent collections [2, 3, 6, 7].
Gelatinous Seasnail 401

Depth Range: Less than 13 m [10] to 1,880 m, typically at 100–300 m [2, 3]. In 2004, one adult was taken at 72 m in the 

northwestern (Russian) Chukchi Sea [7] and a few larvae were taken from surface to 81 m in the western and eastern Chukchi 

Sea [11]. Adults have been taken between 40–500 m in U.S. Beaufort Sea [12].

2,000

1,600

1,200

800

400

0

2,000

1,600

1,200

800

400

0

Open ocean

Depth, in meters

Juveniles and adults

Liparis fabricii

Gelatinous Seasnail

Benthic distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Some fish to 1,800 m

Shore Open ocean Shore Depth, in meters

Spawning and eggs

Larvae

Reproductive distribution

Depths documented in 

U.S. Beaufort Sea 

Overall data from outside 

U.S. Beaufort-Chukchi Seas

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Depths documented in 

U.S. Beaufort Sea 

Benthic and reproductive distribution of Gelatinous Seasnail (Liparis fabricii).

Habitats and Life History

Eggs—Size: 2.1–2.7 mm for mature eggs [13]. Ovaries can contain three sizes of eggs [13]. Time to hatching: 

Unknown. Habitat: Likely benthic, like other Liparis species [2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: At least greater than 38 mm SL (42 mm 

TL) in Canadian Beaufort Sea [14]. Days to juvenile transformation: About one year [15]. Habitat: Pelagic [14].

A few larvae were taken in midwaters in U.S. Chukchi Sea [11]. 

Juveniles—Age and size: Unknown. Habitat: Benthic and pelagic [3]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 21 cm TL [15]. 

Habitat: Benthic and water-column species, often found swimming among ice floes [2, 16, 17]. Nearshore to 

beneath pack ice over bottom depths of more than 2 km. Usually more than 20 km offshore [18]. Attaches to 

laminarian fronds [10]. 

Substrate—Mud or mixed mud, sand, or detritus bottom [16]. 

Physical/chemical—Temperature: -1.9–3.7 °C [6, 10, 19], prefers less than 0 °C [15]. Salinity: Marine and 

estuarine [16, 17], but prefers 30–34 ppt [15].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Swims and feeds up to 1 m above sea bottom [10].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous; sexes are separate [20]. 

Spawning season—September and October in Russian Arctic [13]. 

Fecundity—485–735 large and medium-sized eggs per female [13].
402 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Benthic and pelagic crustaceans (for example, mysids, euphausiids, amphipods, and shrimps), 

polychaetes, and fishes [13, 15, 16]. 

Trophic level—3.33 standard error 0.42 [21].

Biological Interactions

Predators—Snailfishes outside U.S. Chukchi and Beaufort Seas eaten by such predators as larger fishes and 

seals [22–25]. 

Competitors—Likely a range of benthic fish species, including various flatfishes, sculpins, poachers, and 

eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Assuming t

max>3) [21].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Gelatinous Seasnail are not commercially harvested.

Potential Effects of Climate Change

Because the Gelatinous Seasnail are an Arctic species [2] climate warming might result in a northward shift in 

distribution and potential decreased abundance.

Areas for Future Research [B]

Little is known about the ecology and life history of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.
Gelatinous Seasnail 403

References Cited

Able, K.W., and McAllister, D.E., 1980, Revision of the snailfish genus Liparis from Arctic Canada: Ottawa, Department of 

Fisheries and Oceans, Canadian Bulletin of Fisheries and Aquatic Sciences 208. [16]

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [13]

Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86. [10]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [7]

Bibliography

1. Chernova, N.V., 2008, Systematics and phylogeny of fish of the genus Liparis (Liparidae, Scorpaeniformes): Journal of 

Ichthyology, v. 48, no. 10, p. 831–852.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Eastman, J.T., Hikida, R.S., and DeVries, A.L., 1994, Buoyancy studies and microscopy of skin and subdermal extracellular 

matrix of the Antarctic snailfish, Paraliparis devriesi: Journal of Morphology, v. 220, no. 1, 

p. 85–101.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

8. Rand, K.M., and Logerwell, E.A., 2011, The first demersal trawl survey of benthic fish and invertebrates in the Beaufort Sea 

since the late 1970s: Polar Biology, v. 34, no. 4, p. 475–488.

9. McAllister, D.E., 1963, Fishes of the 1960 “Salvelinus” program from western Arctic Canada: National Museum of Canada 

Bulletin, no.185, p. 17–39.

10. Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86.
404 Alaska Arctic Marine Fish Ecology Catalog

11. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, nos. 1–2, p. 57–70.

12. Logerwell, E.A., and Rand, K.M., 2010, Beaufort Sea marine fish monitoring 2008—Pilot survey and test of hypotheses: 

Seattle, Washington, National Oceanic and Atmospheric Administration, National Marine Fisheries Service, Alaska 

Fisheries Science Center, Final Report, BOEMRE 2010-048, 262 p.

13. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

14. Majewski, A.R., Sareault, J.E., and Reist, J., 2006, Fish catch data from offshore sites in the Mackenzie River estuary and 

Beaufort Sea during the open water season, August 2004 aboard the CCGS Nahidik: Winnipeg, Manitoba, Fisheries and 

Oceans Canada, no. 2771, 42 p.

15. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

16. Able, K.W., and McAllister, D.E., 1980, Revision of the snailfish genus Liparis from Arctic Canada: Ottawa, Department of 

Fisheries and Oceans, Canadian Bulletin of Fisheries and Aquatic Sciences 208.

17. Morin, R., and Dodson, J.J., 1986, The ecology of fishes in James Bay, Hudson Bay and Hudson Strait, in Martini, I.P., ed., 

Canadian Inland Seas: Amsterdam, Elsevier Science Publishers B.V., p. 293–323.

18. Mecklenburg, C.W., and Mecklenburg, T.A., 2011, Gelatinous Seasnail—Liparis fabricii Krøyer, 1847: Arctic Ocean 

Diversity Web site, accessed May 2011, at http://www.arcodiv.org/Fish/Liparis_fabricii.html.

19. Christiansen, J.S., ed., 2003, TUNU-1 Expedition—The fish fauna of the NE Greenland fjord systems—Technical report: 

Tromsø, Norway, University of Tromsø, Norwegian College of Fishery Science, Institute of Aquatic Resources, 33 p.

20. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

21. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

22. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.

23. Wakefield, W.W., 1984, Feeding relationships within assemblages of nearshore and mid-continental shelf benthic fishes off 

Oregon: Corvallis, Oregon State University, Master’s thesis, 102 p.

24. Frost, K.J., and Lowry, L.F., 1980, Feeding of ribbon seals (Phoca fasciata) in the Bering Sea in spring: Canadian Journal 

of Zoology, v. 58, no. 9, p. 1,601–1,607.

25. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990.
Variegated Snailfish 405

Variegated Snailfish (Liparis gibbus) 

Bean, 1881

Family Liparidae 

Note on taxonomy: Snailfishes in the genus Liparis 

can be difficult to distinguish from one another and 

L. gibbus has often been confused with other species, including

L. bathyarcticus and L. tunicatus, making much of the available 

information concerning these species uncertain [1].

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: It is particularly difficult to differentiate among 

snailfish species when found as prey. However, snailfishes are generally common and are likely of some ecological importance in 

the Arctic. Larval stages may be seasonally important in coastal food webs.

Physical Description/Attributes: Body tadpole shaped, soft, and covered with gelatinous tissue; and pink, with tan stripes, 

tan with black fin margins, or brownish with blackish spots. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 585) [2]. Swim bladder: Absent [3]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [1]. Elsewhere in Alaska, in eastern Bering Sea, along Aleutian Islands, and in Gulf of 

Alaska south to northern British Columbia, Canada. Worldwide, circumpolar in Arctic Ocean. There are unconfirmed reports 

from southeastern Kamchatka Peninsula, northern Kuril Islands, Russia, and Commander Islands [1]. 

Relative Abundance: Reported to be common in northeastern U.S. Chukchi Sea [6], but this is uncertain because of confusion 

with L. bathyarcticus, which may actually be the more abundant species in the study area [5]. Common in eastern Bering Sea [7].

Variegated Snailfish (Liparis gibbus), 183 mm TL, Bering 

Strait 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Variegated Snailfish (Liparis gibbus) within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [1, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Variegated Snailfish

Liparis gibbus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
406 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 25–364 m, mainly 150 m or less [1, 8]. Three adults were taken between 48–54 m in area between Bering 

Strait and southern Chukchi Sea [9]. Larvae and young juveniles were taken in midwaters of central and northern Chukchi Sea 

between 22–81 m and possibly as shallow as the surface [9].

Benthic and reproductive distribution of Variegated Snailfish (Liparis gibbus).

Some fish to 364 m

Open ocean Shore

Depth, in meters

375

300

225

150

75

0

375

300

225

150

75

0

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Liparis gibbus

Variegated Snailfish

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning and eggs

Depths from U.S. Chukchi Sea

Reproductive distribution

Larvae and young juveniles

(U.S. Chukchi Sea)

Overall data from outside U.S. Beaufort-Chukchi Seas

Older juveniles and adults

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Likely benthic, like other Liparis species [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 15.2 mm SL [10]. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [10]. 

Juveniles—Age and size: Unknown. Habitat: Pelagic to benthic [1, 10]. 

Adults—Age and size at first maturity: Unknown. Females appear to be larger at age than males [11]. Maximum 

age: About 6 years for both males and females. A 250 mm TL fish (much smaller than this species’ maximum 

length) was 6 years [11]. Maximum size: 52.4 cm TL [10], but because the authors included L. bathyacrticus in 

the synonymy of L. gibbus, this measurement could instead pertain to L. bathyarcticus. 

Habitat: Primarily benthic [10, 12] and possibly benthopelagic in that some large fish have been taken in 

midwater trawls [10]. 

Substrate—Soft sea floors and occasionally over rocks [10, 12]. 

Physical/chemical—Temperature: 0.8–10.5 °C [12, 13]. Salinity: Marine [10, 12].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous [14]. 

Spawning season—Perhaps summer based on capture of small larvae in summer in St. Lawrence 

estuary. However, no females taken in Canadian Arctic in spring or summer had mature ovaries [10]. 

Fecundity—Unknown. 
Variegated Snailfish 407

Food and Feeding

Food items—Both benthic and epibenthic, and occasionally water column prey. Crustaceans (for example, 

gammarid and caprellid amphipods, copepods, shrimps, and crabs), polychaetes, pteropods, and small fishes [11, 

15, 16]. 

Trophic level—3.29 standard error 0.46 [17].

Biological Interactions

Predators—Atlantic Cod (Gadus morhua) in Hudson Bay [10]. Snailfishes outside U.S. Chukchi and Beaufort 

Seas eaten by such predators as larger fishes and seals (18–21]. 

Competitors—Likely a range of benthic fish species, including various flatfishes, sculpins, poachers, and 

eelpouts. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Fecundity=4,000–12,000) [17].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Variegated Snailfish are not commercially harvested.

Potential Effects of Climate Change

The Variegated Snailfish are a predominantly Arctic species [1] and it is possible that climate warming could 

cause a retraction northward of its southern populations. If industrial fishing operations ever develop in the 

offshore marine, the species would be vulnerable to bycatch. Changes in energy transfers through decoupling of 

benthic-pelagic energy pathways could affect the availability of benthic foods.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators. It is common enough in most habitats to be considered as a potential indicator 

species in long term monitoring.

References Cited

Able, K.W., and McAllister, D.E., 1980, Revision of the snailfish genus Liparis from Arctic Canada: Ottawa, Department of 

Fisheries and Oceans, Canadian Bulletin of Fisheries and Aquatic Sciences 208. [10]

Falk-Petersen, I.-B., Frivoll, V., Gulliksen, B., Haug, T., and Vader, W., 1988, Age/size relations and food of two snailfishes, 

Liparis gibbus and Careproctus reinhardtii (Teleostei, Liparididae) from Spitsbergen coastal waters: Polar Biology, v. 8, no. 5, 

p. 353–358. [11]
408 Alaska Arctic Marine Fish Ecology Catalog

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [1] 

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [12]

Bibliography

1. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Eastman, J.T., Hikida, R.S., and DeVries, A.L., 1994, Buoyancy studies and microscopy of skin and subdermal extracellular 

matrix of the Antarctic snailfish, Paraliparis devriesi: Journal of Morphology, v. 220, no. 1, 

p. 85–101.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2 p. 195–209.

7. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

8. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

9. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

10. Able, K.W., and McAllister, D.E., 1980, Revision of the snailfish genus Liparis from Arctic Canada: Ottawa, Department of 

Fisheries and Oceans, Canadian Bulletin of Fisheries and Aquatic Sciences 208.

11. Falk-Petersen, I.-B., Frivoll, V., Gulliksen, B., Haug, T., and Vader, W., 1988, Age/size relations and food of two snailfishes, 

Liparis gibbus and Careproctus reinhardtii (Teleostei, Liparididae) from Spitsbergen coastal waters: Polar Biology, v. 8, 

no. 5, p. 353–358.

12. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

13. Christiansen, J.S., ed., 2003, TUNU-1 Expedition—The fish fauna of the NE Greenland fjord systems—Technical report: 

Tromsø, Norway, University of Tromsø, Norwegian College of Fishery Science, Institute of Aquatic Resources, 33 p.

14. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

15. Atkinson, E.G., and Percy, J.A., 1992, Diet comparison among demersal marine fish from the Canadian Arctic: Polar 

Biology, v. 11, no. 8, p. 567–573.
Variegated Snailfish 409

16. Yang, M.-S., Dodd, K., Hibpshman, R., and Whitehouse, A., 2006, Food habits of groundfishes in the Gulf of Alaska 

in 1999 and 2001: U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical 

Memorandum, NMFS-AFSC-164, 189 p.

17. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

18. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.

19. Wakefield, W.W., 1984, Feeding relationships within assemblages of nearshore and mid-continental shelf benthic fishes off 

Oregon: Corvallis, Oregon State University, Master’s thesis, 102 p.

20. Frost, K.J., and Lowry, L.F., 1980, Feeding of ribbon seals (Phoca fasciata) in the Bering Sea in spring: Canadian Journal 

of Zoology, v. 58, no. 9, p. 1,601–1,607.

21. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990.
410 Alaska Arctic Marine Fish Ecology Catalog

Kelp Snailfish (Liparis tunicatus) 

(Reinhardt, 1836)

Family Liparidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown. The life history and ecology of this 

species and its roles in regional ecosystems and food webs are 

poorly understood. 

Physical Description/Attributes: Body tadpole shaped, soft, and 

covered with gelatinous tissue. Yellowish brown, plain or patterned 

with dark brown blotches and bars or stripes, and usually with dark 

bars on caudal fin. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 589) [1]. Swim 

bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas (as far north as 73°1’N, 174°7’W [3]. Elsewhere in Alaska, to northern Bering Sea 

south of St. Lawrence Island. Worldwide, circumpolar, to as far north as 82°29’N, 62°15’W at Ellesmere Island, Canada [4]; and 

in Gulf of Anadyr, Russia [1].

Relative Abundance: Patchily abundant (common) in U.S. Chukchi and Beaufort Seas [6–8].

Kelp Snailfish (Liparis tunicatus), 76 mm TL, western Chukchi 

Sea, 2004. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Kelp Snailfish

Liparis tunicatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Kelp Snailfish (Liparis tunicatus) within Arctic Outer Continental Shelf Planning 

Areas [5] based on review of published literature and specimens from historical and recent collections [3, 4].
Kelp Snailfish 411

Depth Range: Intertidal, including tide pools, to 415 m [4] and perhaps to 600 m [9]. Typically shallower than 50 m [4]. In both 

Russian and U.S. Chukchi Sea, 18 adults and large juveniles (68–110 mm TL) and 5 small juveniles (31–35 mm TL) were taken 

on the bottom between 34–72 m [10] and a juvenile was taken in midwaters between 51 m and the surface [11].

Juveniles and adults

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

Overall data from outside 

U.S. Beaufort-Chukchi Seas

Characteristic of Liparis sp. larvae

Spawning and eggs

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Liparis tunicatus

Kelp Snailfish

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

500

400

300

200

100

0

Reproductive distribution

Some fish perhaps to 600 m

Depths from U.S. Chukchi Sea

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Benthic and reproductive distribution of Kelp Snailfish (Liparis tunicatus).

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. 

Habitat: Likely benthic, like other Liparis species [4]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: At least 36 mm TL [12]. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [12]. 

Juveniles—Age and size: Unknown. Habitat: Pelagic to benthopelagic [12]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 14.3 cm TL [3]. 

Habitat: Benthic to benthopelagic [1, 13]. Commonly found attached to algal blades and occasionally among 

rocks [14–16]. 

Substrate—Rock, sand, and mud [1, 10]. 

Physical/chemical—Temperature: -1.9–2.8 °C [10, 14]. Salinity: Marine [10, 14]; 32.41–33.52 salinity in the 

Chukchi Sea [10].

Behavior

Diel—Both juveniles and adults swim into surface waters at night [13]. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Despite their association with kelp, they have been observed feeding in the water column as much as 1 

m above the substrate [15].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Unknown. 

Spawning season—Unknown. 

Fecundity—Unknown.
412 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Almost entirely benthic and epibenthic crustaceans, including gammarid and lysianassid 

amphipods, cumaceans, cyclopoid copepods, and mysids in Canadian Arctic [17]. Trophic level—3.5 standard 

error 0.50 [18].

Biological Interactions

Predators—Seals in the Canadian Arctic [19]. Unknown. Snailfishes outside U.S. Chukchi and Beaufort Seas 

eaten by such predators as larger fishes and seals (20–23]. 

Competitors—Likely a range of benthic fish species, including various flatfishes, sculpins, poachers, and 

eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Assuming t

max = 3–10) [18].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Kelp Snailfish are not commercially harvested.

Potential Effects of Climate Change

The Kelp Snailfish are a predominantly Arctic species [4] and climate warming could be expected to shift 

distributions to the north and extirpate more southern populations.

Areas for Future Research [B]

Little is known about the ecology of this species from the region. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators.
Kelp Snailfish 413

References Cited

Byers, T., and Prach, R.W., 1988, Diet of the kelp snailfish, Liparis tunicatus, in Jones Sound, Canadian High Arctic: The 

Canadian Field-Naturalist, v. 102, no. 2, p. 242–245. [17]

Majewski, A.R., Sareault, J.E., and Reist, J., 2006, Fish catch data from offshore sites in the Mackenzie River estuary and 

Beaufort Sea during the open water season, August 2004 aboard the CCGS Nahidik: Winnipeg, Manitoba, Fisheries and 

Oceans Canada, no. 2771, 42 p. [12]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [10]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Eastman, J.T., Hikida, R.S., and DeVries, A.L., 1994, Buoyancy studies and microscopy of skin and subdermal extracellular 

matrix of the Antarctic snailfish, Paraliparis devriesi: Journal of Morphology, v. 220, no. 1, 

p. 85–101.

3. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. McAllister, D.E., 1963, Fishes of the 1960 “Salvelinus” program from western Arctic Canada: National Museum of Canada 

Bulletin, no.185, p. 17–39.

7. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

8. Wiswar, D.W., and Frugé, D.J., 2006, Fisheries investigations in western Camden Bay, Arctic National Wildlife Refuge, 

Alaska, 1987: Alaska Fisheries Data Series, U.S. Fish and Wildlife Service, 2006-10, 49 p.

9. Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic marine fishes of Canada: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, no. 2674, 112 p.

10. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

11. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.
414 Alaska Arctic Marine Fish Ecology Catalog

12. Majewski, A.R., Sareault, J.E., and Reist, J., 2006, Fish catch data from offshore sites in the Mackenzie River estuary and 

Beaufort Sea during the open water season, August 2004 aboard the CCGS Nahidik: Winnipeg, Manitoba, Fisheries and 

Oceans Canada, no. 2771, 42 p.

13. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.

14. Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86.

15. McAllister, D.E., 1975, Ecology of the marine fishes of Arctic Canada, in Proceedings of the Circumpolar Conference on 

Northern Ecology, September 15–18, 1975: Ottawa, National Research Council of Canada, 

p. II-49–II-65.

16. Dunton, K.H., and Schonberg, S.V., 2000, The benthic faunal assemblage of the boulder patch kelp community, in Truett, 

J.C., and Johnson, S.R., eds., The natural history of an Arctic oil field—Development and the biota: San Diego, Academic 

Press, p. 371–397.

17. Byers, T., and Prach, R.W., 1988, Diet of the kelp snailfish, Liparis tunicatus, in Jones Sound, Canadian High Arctic: The 

Canadian Field-Naturalist, v. 102, no. 2, p. 242–245.

18. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

19. Able, K.W., and McAllister, D.E., 1980, Revision of the snailfish genus Liparis from Arctic Canada: Ottawa, Department of 

Fisheries and Oceans, Canadian Bulletin of Fisheries and Aquatic Sciences 208.

20. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.

21. Wakefield, W.W., 1984, Feeding relationships within assemblages of nearshore and mid-continental shelf benthic fishes off 

Oregon: Corvallis, Oregon State University, Master’s thesis, 102 p.

22. Frost, K.J., and Lowry, L.F., 1980, Feeding of ribbon seals (Phoca fasciata) in the Bering Sea in spring: Canadian Journal 

of Zoology, v. 58, no. 9, p. 1,601–1,607.

23. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990.
Black Seasnail 415

Black Seasnail (Paraliparis bathybius) 

(Collett, 1879)

Family Liparidae 

Note: Except for geographic range data, all information is from 

areas outside of the Chukchi and Beaufort Seas. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Not important in shelf ecosystems of the U.S. 

Chukchi and Beaufort Seas. Appears to be a deeper water species, 

of low abundance, and limited significance to regional food webs 

and energy flows.

Physical Description/Attributes: Elongate, soft-bodied snailfish without adhesive pelvic disk, and with smooth loose skin 

and brownish black body and fins [1]. For specific diagnostic characteristics, see Coad and others, (1995, page 111) [1]. Swim 

bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: One record from the U.S. Chukchi Sea [3]. Deep continental slope and basin waters offshore of U.S. Chukchi and 

Beaufort Seas. Worldwide, in Norwegian and Greenland Seas, Baffin Bay, central Arctic Basin, Canada Basin north of Alaska [3].

Relative Abundance: Probably common in deep slope and basin waters adjacent to the U.S. Chukchi and Beaufort Seas. 

Elsewhere, common off Greenland and Jan Mayen [3, 6]. Occurrence is rare in shelf habitats in the U.S. Chukchi and Beaufort 

Seas. 

Black Seasnail (Paraliparis bathybius), 198 mm TL, Canada 

Basin, 2005. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution within 2008–09 lease areas [4] of Black Seasnail (Paraliparis bathybius) based on 

review of published literature and specimens from historical and recent collections [3, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Black Seasnail

Paraliparis bathybius

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
416 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 545 m [3] to about 2,824 m [5], common at 545–1,600 m off northern Greenland [3].

3-9-6g_BlackSeasnail

Open ocean

Depth, in meters

4,000

3,200

2,400

1,600

800

0

Adults and juveniles

Paraliparis bathybius

Black Seasnail

Benthic distribution

Some fish to 4,009 m

Depth, in meters

Overall benthic depth range. 

Specific depth range of either 

juveniles and adults is known

Open ocean No data for this species Shore

Spawning and eggs

Reproductive distribution

Depth range of spawning is 

uncertain. Potential location of 

spawning and eggs is based on 

depth range of benthic individuals

Larvae maybe pelagic

but specific depths unknown

Data from outside U.S. Beaufort-Chukchi Seas 3,000

2,400

600

1,200

1,800

0

Benthic and reproductive distribution of Black Seasnail (Paraliparis bathybius). 

Habitats and Life History

Eggs—Size: 4 mm [6]. Time to hatching: Unknown. Habitat: Likely benthic, like those of other Paraliparis

species [7]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic, like those of other Paraliparis species [7]. 

Juveniles—Age and size: Unknown. Habitat: Benthic to benthopelagic [3, 7]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 27 cm [8]. Habitat: 

Benthic to benthopelagic [3, 7]. 

Substrate—Unknown. 

Physical/chemical—Temperature: -0.1–0.9 °C [5]. Salinity: Marine [7].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—May form large schools to spawn [7]. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous; sexes are separate [9]. 

Spawning season—June through August off Russia [7]. 

Fecundity—100–434 eggs [7].
Black Seasnail 417

Food and Feeding

Food items—Mostly pelagic hyperiid amphipods and mysids as well as benthic gastropods [7, 10]. 

Trophic level—3.28 standard error 0.50 [11].

Biological Interactions

Predators—Unknown. Snailfishes outside U.S. Chukchi and Beaufort Seas eaten by such predators as larger 

fishes and seals [12–15]. 

Competitors—Likely a range of deep water benthic fish species, including various flatfishes, sculpins, poachers, 

and eelpouts.

1980 1990 2000 2010

Resilience

High, minimum population doubling time: less than 15 months (Preliminary K or Fecundity) [11]. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Black Seasnail are not commercially harvested.

Potential Effects of Climate Change

The Black Seasnail are an endemic Arctic species found in deeper and less studied waters of the region [3]. There 

is not enough information to make an informed assessment about potential climate effects.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators. 
418 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [8]

Coad, B.W., Waszczuk, H., and Labignan, I., 1995, Encyclopedia of Canadian fishes: Canadian Museum of Nature and Canadian 

Sportfishing Productions, Inc., 928 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, A., 

Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 2011, 

Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p. [6]

Bibliography

1. Coad, B.W., Waszczuk, H., and Labignan, I., 1995, Encyclopedia of Canadian fishes: Canadian Museum of Nature and 

Canadian Sportfishing Productions, Inc., 928 p.

2. Eastman, J.T., Hikida, R.S., and DeVries, A.L., 1994, Buoyancy studies and microscopy of skin and subdermal extracellular 

matrix of the Antarctic snailfish, Paraliparis devriesi: Journal of Morphology, v. 220, no. 1, 

p. 85–101.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

7. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

8. Wienerroither, R., Nedreaas, K., Uiblein, F., Christiansen, J., Byrkjedal, I., and Karamushko, O., 2011, The marine fishes of 

Jan Mayen Island, NE Atlantic—Past and present: Marine Biodiversity, v. 41, no. 3, p. 395–411

9. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

10. Bjelland, O., Bergstad, O.A., Skjæraasen, J.E., and Meland, K., 2000, Trophic ecology of deep-water fishes associated with 

the continental slope of the eastern Norwegian Sea: Sarsia, v. 85, no. 2, p. 101–117.

11. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

12. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.
Black Seasnail 419

13. Wakefield, W.W., 1984, Feeding relationships within assemblages of nearshore and mid-continental shelf benthic fishes off 

Oregon: Corvallis, Oregon State University, Master’s thesis, 102 p.

14. Frost, K.J., and Lowry, L.F., 1980, Feeding of ribbon seals (Phoca fasciata) in the Bering Sea in spring: Canadian Journal 

of Zoology, v. 58, no. 9, p. 1601–1607.

15. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990.
420 Alaska Arctic Marine Fish Ecology Catalog

Halfbarred Pout (Gymnelus hemifasciatus) 

Andriashev, 1937

Family Zoarcidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: The Halfbarred Pout is likely of relatively small 

ecological importance in the U.S. Chukchi and Beaufort Seas. 

Physical Description/Attributes: An elongate body with dark 

bands down to lateral line or almost to anal fin on females. Dark 

orange bands reach anal fin on males, and anal fin changes to black [1]. For specific diagnostic characteristics, see Fishes 

of Alaska, Mecklenburg and others, 2002, p. 689) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: 

Unknown.

Range: U.S. Chukchi and Beaufort Seas [2]. Elsewhere in Alaska, Bering Sea to Prince William Sound, northern Gulf of Alaska 

[2]. Worldwide, Sea of Okhotsk, Commander Islands, east coast of Kamchatka Peninsula and Barents Sea [1]. 

Relative Abundance: Common in U.S. Chukchi and Beaufort Seas [4].

Halfbarred Pout (Gymnelus hemifasciatus), male, 142 mm TL, 

Chukchi Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Halfbarred Pout (Gymnelus hemifasciatus) within Arctic Outer Continental 

Shelf Planning Areas [3] based on review of published literature and specimens from historical and recent 

collections [1, 2, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Halfbarred Pout

Gymnelus hemifasciatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Halfbarred Pout 421

Depth Range: Intertidal to 216 m, typically at 40–80 m on inner and outer shelf [2]. Taken in U.S. Chukchi Sea off 

Point Lay at 29 m [5]. In general, eelpout spawning and larvae are at same depths that adults inhabit [1].

Open ocean Shore

Depth, in meters

250

200

150

100

50

0

250

200

150

100

50

0

Juveniles and adults

Gymnelus hemifasciatus

Halfbarred Pout

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning, 

eggs, and larvae

Documented in U.S. Chukchi Sea at 29 m

Reproductive distribution

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

Overall data from outside U.S. Beaufort-Chukchi Seas

Some fish to 

216 m

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

Benthic and reproduction distribution of Halfbarred Pout (Gymnelus hemifasciatus).

Habitats and Life History

Eggs—Size: 2.5–4.5 mm [6]. Time to hatching: Unknown. Habitat: Benthic [1]. Larvae: Size at hatching: 

Unknown. Size at juvenile transformation: Unknown. Days to juvenile transformation: Unknown. Habitat: 

Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Age is unknown. Females mature at 70–80 mm SL [1]. Maximum age: 

Unknown. Maximum size: As long as 18 cm TL [4]. Habitat: Benthic [1]. 

Substrate—Mud and gravel [1]. 

Physical/chemical—Temperature: -1.8–10.5 °C [4]. Salinity: Unknown. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies. An analysis of DNA barcode and morphological data confirmed the validity of 

this species in the Pacific Arctic Region and suggested the possibility of two morphotypes based on ecological 

preferences [8].

Reproduction

Mode—Separate sexes. Oviparous. 

Spawning season—Late August to September. 

Fecundity—8–26 eggs [6].
422 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Gammarid amphipods, polychaetes, bivalves, other benthic invertebrates [6]. 

Trophic level—3.11 standard error 0.30 [7].

Biological Interactions

Predators—Seals, seabirds, cods and other fishes [6]. 

Competitors—Likely competitors include a range of small benthic-feeding fishes, including snailfishes, 

sculpins, flatfishes, and other eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Assuming t

max = 3–10) [7].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, the Halfbarred Pout are not commercially harvested.

Potential Effects of Climate Change

Unknown. Halfbarred Pout occur in Arctic and Boreal Pacific waters. Climate warming would have little or no 

effect on its distribution but may affect abundance patterns.

Areas for Future Research [B]

This is a little-known species. Research needs include: (1) depth and location of pelagic larvae, (2) depth, 

location, and timing of young-of-the-year benthic recruitment, (3) preferred depth ranges for juveniles and adults, 

(4) spawning season, (5) seasonal and ontogenetic movements, (6) population studies, (7) prey, and (8) predators.
Halfbarred Pout 423

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia] [6]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140 and Online Resource 1. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

6. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

7. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

8. Mecklenburg, C.W., and Anderson, M.E., 2015, Reassessment of multiple species of Gymnelus (Teleostei: Zoarcidae) in 

Pacific Arctic and boreal regions: Zootaxa, v. 3948, no. 2, p. 263–278.
424 Alaska Arctic Marine Fish Ecology Catalog

Fish Doctor (Gymnelus viridis) 

(Fabricius, 1780)

Family Zoarcidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown, although likely of minor importance. 

Physical Description/Attributes: Coloration grayish to greenish 

brown, varying from nearly monotone to mottled overall or with 

broad, mottled or solid orange or reddish brown to bluish black 

bands [1, 2]. In mature males, anal fin is black [3, 4]. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg 

and others, 2002, p. 690) [3]. Swim bladder: Absent [3]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [5]. Elsewhere in Alaska, southward to eastern Bering Sea and Aleutian Islands [3]. 

Worldwide, circumpolar in Arctic Ocean and southward to Nova Scotia, Gulf of Anadyr, and southern Bering Sea [5].

Relative Abundance: Common with patchy distributions in both U.S. Chukchi and Beaufort Seas [7, 8].

Fish Doctor (Gymnelus viridis), male, 174 mm TL, Chukchi 

Sea, 2007. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Fish Doctor (Gymnelus viridis) within Arctic Outer Continental Shelf Planning Areas 

[6] based on review of published literature and specimens from historical and recent collections [4, 5].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Fish Doctor

Gymnelus viridis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Fish Doctor 425

Depth Range: Intertidal to 318 m, typically 50 m or less [1, 3, 9, 10]. In general, eelpouts produce eggs and larvae at same 

depths adults inhabit [3].

Benthic and reproductive distribution of Fish Doctor (Gymnelus viridis).

398FihD

Juveniles and adults

Overall benthic depth range. 

Specific depth range of either

juveniles or adults is unknown

Data from outside U.S. Beaufort-Chukchi Seas

Spawning, eggs, and larvae

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Gymnelus viridis

Fish Doctor

Benthic distribution

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Reproductive distribution

Some fish to 318 m

No data for this species

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

Habitats and Life History

Eggs—Size: 1.6–5.2 mm [1]. Time to hatching: Unknown. Habitat: Benthic and adhesive [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 3.0 cm TL [1]. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1], nearshore, under debris, kelp, rocks, and other cover 

[2]. 

Adults—Age and size at first maturity: Age is unknown. Females mature at about 9–10 cm TL [1, 7]. Maximum 

age: In Northwest Territories, at least 12 years [1]. Maximum size: 26.0 cm SL, more than 26.6 cm TL. Males 

grow larger than females [1]. Habitat: Benthic [1], nearshore, under debris, kelp, rocks, and other cover [2]. 

Substrate—Boulders, cobble, and soft sea floors [1, 9–12]. 

Physical/chemical—Temperature: -1.9–10.5 °C [4, 12, 13]. Salinity: Mainly marine but tolerates brackish 

conditions with salinity as low as 27 ppt [11].

Behavior

Diel—Shelters under some form of cover, such as rocks, algae, or wood, and individuals rarely remain out in the 

open [1]. 

Seasonal—Unknown. 

Reproductive—Little is known. During late summer or autumn, eggs are laid in nests and may be guarded by 

one or both parents [1, 14]. 

Schooling—Unknown. 

Feeding—In Canadian Arctic, feeding activity declines in the winter [15].

Populations or Stocks

There have been no studies. An analysis of DNA barcodes and morphological data confirmed the validity of this 

species in the Pacific Arctic Region [19].

Reproduction

Mode—Separate sexes. Oviparous. 

Spawning season—Late summer and early autumn [1, 11, 14]. 

Fecundity—20–106 eggs. Females may not spawn every year [1, 11, 14].
426 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Mainly gammarid amphipods, as well as caprellid amphipods, mysids, polychaetes, and copepods 

[7]. In Canadian Arctic, a similar diet as well as clam siphons. Males tended to focus on amphipods and females 

on polychaetes. 

Trophic level—3.06 standard error 0.26 [16].

Biological Interactions

Predators—In central and eastern Canadian Arctic, cods, Thick-billed Murres, and bearded seals [17, 18]. 

Competitors—A range of bottom-feeding microcarnivores including sculpins, flatfishes, snailfishes, and other 

eelpouts.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (t

max = 12) [16].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Fish Doctor are not commercially harvested.

Potential Effects of Climate Change

As the Fish Doctor are predominantly an Arctic species it would be expected that climate warming would shift 

the distribution of the species northward. 

Areas for Future Research [B]

Little is known about this species in U.S. Arctic marine environment. Research needs include: (1) depth and 

location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred 

depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, 

(6) population studies, (7) prey, and (8) predators. 
Fish Doctor 427

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [11]

Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western Beaufort 

Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA Technical Report 

NMFS-SSRF-764, 22 p. [7]

Green, J.M., and Mitchell, L.R., 1997, Biology of the fish doctor, an eelpout, from Cornwallis Island, Northwest Territories, 

Canada: American Fisheries Society Symposium 19, p. 140–174. [1]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [5]

Bibliography

1. Green, J.M., and Mitchell, L.R., 1997, Biology of the fish doctor, an eelpout, from Cornwallis Island, Northwest Territories, 

Canada: American Fisheries Society Symposium 19, p. 140–174.

2. Mecklenburg, C.W., and Mecklenburg, T.A., 2011, Fish Doctor—Gymnelus viridis (Fabricius, 1780): Arctic Ocean 

Diversity Web site, accessed June 2011 at http://www.arcodiv.org/Fish/Gymnelus_viridis.html.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,=116 p.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western 

Beaufort Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA 

Technical Report NMFS-SSRF-764, 22 p.

8. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

9. Ellis, D.V., 1962, Observations on the distribution and ecology of some Arctic fish: Arctic, v. 15, no. 3, 

p. 179–189.

10. McAllister, D.E., 1975, Ecology of the marine fishes of Arctic Canada, in Proceedings of the Circumpolar Conference on 

Northern Ecology, September 15–18, 1975: Ottawa, National Research Council of Canada, 

p. II-49–II-65.

11. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]
428 Alaska Arctic Marine Fish Ecology Catalog

12. Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86.

13. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

14. Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p.

15. Green, J.M., 1983, Observations on the food of marine fishes from Resolute Bay, Cornwallis Island, Northwest Territories: 

Astarte, v. 12, no. 2, p. 63–68.

16. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

17. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.

18. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.

19. Mecklenburg, C.W., and Anderson, M.E., 2015, Reassessment of multiple species of Gymnelus (Teleostei: Zoarcidae) in 

Pacific Arctic and boreal regions: Zootaxa, v. 3948, no. 2, p. 263–278.
Adolf’s Eelpout 429

Adolf’s Eelpout (Lycodes adolfi) 

Nielsen & Fosså, 1993

Family Zoarcidae 

Note: Except for physical description and geographic 

range data, all information is from areas outside of 

the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: This species has been reported from slope waters of the U.S. Beaufort Sea. It is rare with an unknown but 

suspected minimal role in regional food webs.

Physical Description/Attributes: Brown, grading from nearly black on the head to gray-brown toward the tail, without 

distinctive markings [1]. For specific diagnostic characteristics, see Nielsen and Fosså (1993) [2]. Swim bladder: Absent [3]. 

Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Beaufort Sea [1]. Arctic Ocean north of the Chukchi Sea (north of the 200-mile limit); Baffin Bay off Canada and 

Greenland, Greenland and Norwegian Seas, and Arctic Ocean north of Spitsbergen [4].

Relative Abundance: Common in U.S. Beaufort Sea in deep water on the continental slope, based on a few records of number 

of specimens per haul (as many as 36 or more) [1]. Not found in Chukchi Sea; found far north of the Chukchi Sea, north of 

the 200-mile limit. Elsewhere, common in Baffin Bay [6], eastern and western sides of Greenland, eastern slope of the Yermak 

Plateau north of Spitsbergen, Norway [7], and Norwegian Sea [6].

Adolf’s Eelpout, (Lycodes adolfi), 193 mm, Chukchi Borderland, 

2009. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Adolf’s Eelpout (Lycodes adolfi) within Arctic Outer Continental Shelf Planning 

Areas [5] based on review of published literature and specimens from historical and recent collections [1, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Adolf’s Eelpout

Lycodes adolfi

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
430 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Benthic, 386–2,150 m [6, 8]. Taken on U.S. Beaufort Sea slope at 500 m and 1,000 m [1]. In general, eelpout 

spawning and larvae occur at the same depths that adults inhabit [9].

Benthic and reproductive distribution of Adolf’s Eelpout (Lycodes adolfi).

3-9-9gAdolfsEelpout

2,000

1,600

1,200

800

400

0

2,000

1,600

1,200

800

400

0

Open ocean

Depth, in meters

Juveniles and adults

Lycodes adolfi

Adolf’s Eelpout

Benthic distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Some fish to 2,150 m

Shore Open ocean Shore Depth, in meters

Spawning, eggs, and larvae

Reproductive distribution

Overall data from outside 

U.S. Beaufort-Chukchi Seas

Depth range of spawning is

uncertain. Potential location of 

spawning, eggs and larvae is based

on depth range of benthic individuals 

Depths documented in 

U.S. Beaufort Sea 

No data for this species

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [3]. Larvae: Size at hatching: Unknown. 

Size at juvenile transformation: Unknown. Days to juvenile transformation: Unknown. Habitat: Benthic [3]. 

Juveniles—Age and size: Unknown. Immature fish from Arctic Ocean north of Chukchi Sea were 130–193 mm 

TL [1]. Habitat: Benthic [3]. 

Adults—Age and size at first maturity: Unknown. Mature fish off Greenland were 205–238 mm TL [1]. 

Maximum age: Unknown. Maximum size: 28.6 cm TL [12]. Habitat: Benthic, in deep waters [4]. 

Substrate—Mixture of mud, gravel, and rock [1]. 

Physical/chemical—Temperature: -0.9–3.7 °C off Greenland [1]. Taken at 0.8–0.9 °C in Arctic Ocean far north 

Chukchi Sea [1]. Salinity: Marine, taken at 34.86 ppt in the Arctic Ocean far north of Chukchi Sea [1]. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [3]. 

Spawning season—Unknown. 

Fecundity—Unknown.
Adolf’s Eelpout 431

Food and Feeding

Food items—Crustaceans [10]. 

Trophic level—3.5 standard error ±0.50 [11].

Biological Interactions

Predators—Unknown. 

Competitors—Likely other benthic microcarnivores including sculpins, flatfishes, snailfishes, and other 

eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [11]. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Adolf’s Eelpout are not commercially harvested.

Potential Effects of Climate Change

Because Adolf’s Eelpout are a strictly deep-water, slope-dwelling Arctic species and little is known about climate 

change at slope depths in the Arctic, potential effects on this species has not been estimated.

Areas for Future Research [B]

Little is known about this species’ ecology and life history. Research needs include: (1) depth and location of 

pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators. 
432 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, Iceland, 

Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at http://caff.is/

monitoring-series/370-pacific-arcticmarine-fishes.

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]

Nielsen, J.G., and Fosså, S.A., 1993, Lycodes adolfi, a new species of eelpout (Zoarcidae) from Greenland: Cybium, v. 17, 

p. 39–44. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

2. Nielsen, J.G., and Fosså, S.A., 1993, Lycodes adolfi, a new species of eelpout (Zoarcidae) from Greenland: Cybium, v. 17, 

p. 39–44.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Byrkjedal, I., Brattegard, T., and Møller, P.R., 2009, Lycodes adolfi Nielsen & Fosså, 1993 (Teleostei: Zoarcidae) recorded 

near Jan Mayen and in the eastern part of the Norwegian Sea: Fauna Norvegica, v. 28, p. 1–3.

7. Byrkjedal, I., Langhelle, G., Wenneck, T.d.L., and Wienerroither, R., 2011, Lycodes adolfi Nielsen and Fosså, 1993 

(Teleostei: Zoarcidae) found in the Arctic Ocean: Polar Biology, v. 34, no. 3, p. 465–467.

8. Møller, P.D., and Gravelund, P., 2003, Phylogeny of the eelpout genus Lycodes (Pisces, Zoarcidae) as inferred from 

mitochondrial cyctochrome b and 12S rDNA: Molecular and Phylogenetic Evolution, v. 26, no. 3, 

p. 369–388.

9. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

10. Coad, B.W., Waszczuk, H., and Labignan, I., 1995, Encyclopedia of Canadian fishes: Canadian Museum of Nature and 

Canadian Sportfishing Productions, Inc., 928 p.

11. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

12. California Academy of Science, Catalog of fishes: California Academy of Sciences online database, accessed June 16, 2016, 

at http://www.calacademy.org/scientists/projects/catalog-of-fishes.
Doubleline Eelpout 433

Doubleline Eelpout (Lycodes eudipleurostictus) 

Jensen, 1902

Family Zoarcidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown, but based on occurrence, most likely 

of minimal importance in regional food webs.

Physical Description/Attributes: Dark brown elongate body with narrow light bands passing onto dorsal fin and posteriorly 

onto anal fin. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 

p. 723) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Beaufort Sea; three records from deep water [4]. Elsewhere, from eastern Canadian Arctic, including Baffin Bay, to 

western and eastern Greenland, Iceland, and to Kara Sea, northward to Svalbard Archipelago, and north of Severnaya Zemlya, 

Russia [1, 2].

Relative Abundance: At least occasional in U.S. Beaufort Sea [4].Common off West Greenland [5].

Doubleline Eelpout, (Lycodes eudipleurostictus), about 

255 mm TL, Svalbard Archipelago, 2011. Photograph by Arve 

Lynghammar, University of Tromsø, Norway. 

Location within Arctic Outer Continental Shelf Planning Areas [3] of the one existing record of Doubleline 

Eelpout (Lycodes eudipleurostictus) [1, 2, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Doubleline Eelpout

Lycodes eudipleurostictus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
434 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 54–1,280 m [2, 4], rarely less than 250 m [1]. In general, eelpout spawning and larvae occur at same depths that 

adults inhabit [1].

Benthic and reproductive distribution of Doubleline Eelpout (Lycodes eudipleurostictus). 

3910DblliElt

Some fish to 1,187 m

Open ocean Shore

Depth, in meters

1,250

1,000

750

500

250

0

1,250

1,000

750

500

250

0

 Data from outside 

U.S. Beaufort-Chukchi Seas 

Lycodes eudipleurostictus

Doubleline Eelpout

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

No data for this species

Juveniles and adults Spawning, eggs, and larvae

Depth range of spawning

is uncertain. Potential location

of spawning, eggs, and larvae

is based on depth range of

benthic individuals 

Habitats and Life History

Eggs—Size: 2.7–8 mm [5]. Time to hatching: Unknown. Habitat: Benthic [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Age unknown. More than 23 cm TL for females and 29 cm TL for males. 

Males grow larger than females. [5]. Maximum age: 9 years [6]. Maximum size: To 45 cm TL [1, 5]. Habitat: 

Benthic [1]. 

Substrate—Muddy bottoms [1, 6]. 

Physical/chemical—Temperature: -0.6–4.9 °C [5, 7], typically less than 2.5 °C [8]. Salinity: 

Marine [1, 9].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes. Oviparous. 

Spawning season—Unknown. In Russia, almost ripe eggs were taken in August [8]. 

Fecundity—120–300 orange eggs [5, 8].
Doubleline Eelpout 435

Food and Feeding

Food items—Unknown. In general, smaller eelpouts eat mainly infaunal prey; larger individuals feed more on 

epibenthic prey [10]. 

Trophic level—3.54 standard error 0.47 [10].

Biological Interactions

Predators—Unknown. 

Competitors—Perhaps other eelpouts, as well as such other benthic feeding fishes as sculpins and flatfishes.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (Preliminary K or Fecundity) [10].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Doubleline Eelpout are not commercially harvested.

Potential Effects of Climate Change

Unknown.

Areas for Future Research [B]

Little is known about the life history and ecology of this species. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators.
436 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140 and Online Resource 1. [2]

Møller, P.R., and Jørgensen, O.A., 2000, Distribution and abundance of eelpouts (Pisces, Zoarcidae) off West Greenland: Sarsia, 

v. 85, no. 1, p. 23–48. [5]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Møller, P.R., and Jørgensen, O.A., 2000, Distribution and abundance of eelpouts (Pisces, Zoarcidae) off West Greenland: 

Sarsia, v. 85, no. 1, p. 23–48.

6. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

7. Christiansen, J.S., ed., 2003, TUNU-1 Expedition—The fish fauna of the NE Greenland fjord systems—Technical report: 

Tromsø, Norway, University of Tromsø, Norwegian College of Fishery Science, Institute of Aquatic Resources, 33 p.

8. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

9. Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic marine fishes of Canada: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, no. 2674, 112 p.

10. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Glacial Eelpout 437

Glacial Eelpout (Lycodes frigidus) 

Collett, 1879

Family Zoarcidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown, although likely of minor importance 

in the study area. This is possibly the most abundant eelpout in the 

Arctic Basin and thus is ecologically significant there.

Physical Description/Attributes: An elongate, tapering body 

with a large head (22–28 percent of body) and all ages uniformly colored dark brownish gray, brown, or gray-violet. For 

specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 726) [1]. Swim bladder: Absent [1]. 

Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [4]. Circumpolar on Arctic Ocean continental rises and basins, including Makarov 

Basin north of eastern Siberia, Canada Basin north of Alaska, Eurasian Arctic Basin, and Norwegian and Greenland Seas [2].

Relative Abundance: Unknown, but may be common in very deep waters of U.S. Chukchi and Beaufort Seas [2]. Common in 

deep part (>1,500 m) of Norwegian Sea [2].

Glacial Eelpout, (Lycodes frigidus), juvenile, 76 mm, Canada 

Basin, 2005. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Glacial Eelpout (Lycodes frigidus) within Arctic Outer Continental Shelf Planning 

Areas [3] based on review of published literature and specimens from historical and recent collections [1, 2, 4].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Glacial Eelpout

Lycodes frigidus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
438 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 475–3,600 m [2, 5], rarely at less than 1,000 m [2, 6]. Taken in U.S. Chukchi Sea north-northwest of Point 

Barrow at 2,500 m [2]. In general, eelpout spawning and larvae occur at the same depths that adults inhabit [6].

Benthic and reproductive distribution of Glacial Eelpout (Lycodes frigidus).

Open ocean Shore

Depth, in meters

4,000

3,200

2,400

1,600

800

0

3,000

2,400

1,800

1,200

600

0

Some fish to 3,600 m

Lycodes frigidus

Glacial Eelpout

Benthic distribution

Data from outside U.S. Beaufort-Chukchi Seas

Open ocean No data for this species Shore

Depth, in meters

 Adults and juveniles Spawning, eggs, and larvae

Reproductive distribution

Depth range of spawning

is uncertain. Potential location of

spawning, eggs and larvae is based 

on depth range of benthic individuals 

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Habitats and Life History

Eggs—Size: 7 mm [6]. Time to hatching: Unknown. Habitat: Benthic [2, 6]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [2, 6]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [2, 6]. 

Adults—Age and size at first maturity: Unknown. One ripe female was 50 cm long TL [6]. Maximum age: 33 

years [7]. Maximum size: 69.0 cm TL [1]. Habitat: Benthic [2, 6]. 

Substrate—Mud bottoms [1]. 

Physical/chemical—Temperature: Off Russia, found almost always at -0.6–1.6 °C [6]. Spawns from -2 to 

-0.6 °C [8, 9]. Salinity: Marine.

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Moves along bottom, stirring up small prey [10].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes. Oviparous [11]. 

Spawning season—Perhaps autumn or winter [7]. 

Fecundity—One female taken near the Faroe Islands contained 500 ripe eggs [6].
Glacial Eelpout 439

Food and Feeding

Food items—Eats various crustaceans including amphipods and shrimps, as well as fishes, brittle stars, 

cephalopods, sipunculids [6]. 

Trophic level—3.84 standard error 0.51 [12].

Biological Interactions

Predators—Unknown. 

Competitors—Unknown, but likely other deeper water eelpouts, as well as flatfishes.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (Preliminary K or Fecundity) [12].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Glacial Eelpout are not commercially harvested.

Potential Effects of Climate Change

As Glacial Eelpout are predominantly Arctic species, climate effects might result in a northward shift in its 

distribution. 

Areas for Future Research [B]

There is very little known about this species biology or ecology species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.
440 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [6]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Bergmann, M., Dannheim, J., Bauerfeind, E., and Klages, M., 2009, Trophic relationships along a bathymetric gradient at 

the deep-sea observatory HAUSGARTEN: Deep-Sea Research I, v. 56, no. 3, p. 408–424.

6. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

7. Hildebrandt, N., Bergmann, M., and Knust, R., 2011, Longevity and growth efficiency of two deep-dwelling Arctic 

zoarcids and comparison with eight other zoarcid species from different climatic regions: Polar Biology, v. 34, no. 10, 

p. 1,523–1,533.

8. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

9. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

10. Mecklenburg, C.W., and Mecklenburg, T.A., 2011, Glacial Eelpout—Lycodes frigidus Collett, 1879: Arctic Ocean Diversity 

Web site, accessed June 2011, at http://www.arcodiv.org/Fish/Lycodes_frigidus.html.

11. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

12. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Shulupaoluk 441

Shulupaoluk (Lycodes jugoricus) 

Knipowitsch, 1906

Family Zoarcidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Rarely observed in U.S. Beaufort Sea. Of potential 

ecological importance in brackish shallow waters of and near river 

mouths. 

Physical Description/Attributes: Slender, elongate body, white ventrally and yellowish laterally, with wedge-shaped 

dark bands widening dorsally and extending onto dorsal fins. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, p. 706) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Beaufort Sea [2]. Worldwide, in White, Kara, Laptev, and East Siberian Seas; and Canadian Beaufort Sea to 

Boothia Peninsula, Canada [1, 3].

Relative Abundance: Possibly common in freshened nearshore waters of the U.S. Beaufort Sea but generally overlooked 

because of confusion with similar looking species [2].Common in Canadian Beaufort Sea at Tuktoyaktuk Harbor [5].

Shulupaoluk, (Lycodes jugoricus),156 mm, Beaufort Sea. 

Photograph by C.W. Mecklenburg, Point Stephens Research.

Geographic distribution of Shulupaoluk (Lycodes jugoricus) within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [1–3]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Shulupaoluk

Lycodes jugoricus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

Occurrence records

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
442 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 3–90 m [3], typically 7–15 m [1, 5]. In general, eelpout spawning and larvae are at same depths where adults 

reside [1].

Benthic and reproductive distribution of Shulupaoluk (Lycodes jugoricus). 

3-9-12gShulupaolukEelpout

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Juveniles and adults

Lycodes jugoricus

Shulupaoluk

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning, eggs, and larvae

Reproductive distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Data from outside U.S. Chukchi and Beaufort Seas

Some fish

to 90 m

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic, nearshore [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic, nearshore [1]. 

Juveniles: Age and size: Unknown. Habitat: Benthic, nearshore [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: To 40.0 cm TL [1], 

reported without documentation to 52.1 cm TL [6]. Habitat: Benthic, nearshore [1, 3]. 

Substrate—Soft bottoms [7]. 

Physical/chemical—Temperature: Below 0 to 4.5 °C [8]. Salinity: Brackish waters of about 15–27 ppt [7, 8].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes. Oviparous [9]. 

Spawning season—Unknown. 

Fecundity—Unknown. 
Shulupaoluk 443

Food and Feeding

Food items—Isopods, mysids, polychaetes, bivalves, and fishes [6, 7, 10]. 

Trophic level—3.16 standard error 0.38 [7].

Biological Interactions

Predators—Unknown. 

Competitors—Unknown, but likely other shallow-water eelpouts, sculpins, and flatfishes.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [7].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Shlupaoluk are not commercially harvested.

Potential Effects of Climate Change

Shulupaoluk are an Arctic nearshore species inhabiting low-salinity, cold water habitats. If they are unable to 

adapt to warmer, brackish conditions, their population could be extremely vulnerable to effects associated with 

high discharges of warm freshwater to the coast. 

Areas for Future Research [B]

Little is known about the biology and ecology from the region. Research needs include: (1) depth and location of 

pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators. 
444 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [8]

Hopky, G.E., and Ratynski, R.A., 1983, Relative abundance, spatial and temporal distribution, age and growth of fishes in 

Tuktoyaktuk Harbour, N.W.T., 28 June to 5 September, 1981: Canadian Manuscript Report of Fisheries and Aquatic Sciences 

1983, 77 p. [5]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Hopky, G.E., and Ratynski, R.A., 1983, Relative abundance, spatial and temporal distribution, age and growth of fishes 

in Tuktoyaktuk Harbour, N.W.T., 28 June to 5 September, 1981: Canadian Manuscript Report of Fisheries and Aquatic 

Sciences 1983, 77 p.

6. Lacho, G., 1991, Stomach content analyses of fishes from Tuktoyaktuk Harbour, N.W.T., 1981: Winnipeg, Manitoba, 

Canadian Data Report of Fisheries and Aquatic Sciences, Central and Arctic Region, Department of Fisheries and Oceans, 

no. 853, 15 p.

7. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

8. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

9. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

10. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.
White Sea Eelpout 445

White Sea Eelpout (Lycodes marisalbi) 

Knipowitsch, 1906

Family Zoarcidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area.

Note on taxonomy: Once classified as a subspecies of Lycodes 

pallidus but recently resurrected as a valid species [1].

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: This species appears to be rare in the U.S. Arctic and thus is likely to be of limited ecological importance. 

Physical Description/Attributes: An elongate, tapering body either brownish with 6–11 white bars along each side and dorsal 

fin or uniformly brown. Peritoneum speckled. For specific diagnostic characteristics, see Møller (2000) [1] and Fishes of Alaska, 

(Mecklenburg and others, 2002, page 721, as L. pallidus) [2]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood 

serum: Unknown.

Range: Central and eastern U.S. Beaufort Sea [2, 3]. Elsewhere, in Canadian Arctic at Mould Bay, Prince Patrick Island, to 

Mackenzie Bay and eastward to Dease Strait, Bathurst Inlet; disjunct population in White Sea [1].

Relative Abundance: Occasional, although previously misidentified as L. pallidus, in U.S. Beaufort Sea [3]. Common in 

southeastern part of Canadian Beaufort Sea [1].

White Sea Eelpout, (Lycodes marisalbi), 135 mm, Beaufort 

Sea, 2011. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of White Sea Eelpout (Lycodes marisalbi) within Arctic Outer Continental Shelf 

Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [2, 3, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

White Sea Eelpout

Lycodes marisalbi

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
446 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 6–335 m [1, 3]. Mostly less than 50 m in U.S. Beaufort Sea [3]. In general, eelpout spawning and larvae occur at 

same depths that adults inhabit [2].

Benthic and reproductive distribution of White Sea Eelpout (Lycodes marisalbi).

3913WhiSEl

Juveniles and adults

Overall benthic depth range.

Specific depth range of either 

juveniles or adults is unknown 

Overall data from outside 

U.S. Beaufort-Chukchi Seas

Spawning, eggs, and larvae

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Lycodes marisalbi

White Sea Eelpout

Benthic distribution

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Reproductive distribution

Some fish to 335 m

No data for this species

Depth range of spawning is uncertain. 

Potential location of spawning,eggs and 

larvae is based on depth rangeof benthic 

individuals 

Depths from U.S. Beaufort Sea

Habitats and Life History

Eggs—Size: 2–2.5 mm [1]. Time to hatching: Unknown. Habitat: Benthic [1, 3]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1, 3]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1, 3]. 

Adults—Age and size at first maturity: Age unknown. At least 11.9 cm SL in females [1]. Maximum age: 

Unknown. Maximum size: 21.3 cm SL [1]. Habitat: Benthic [1, 3], in shallow waters influenced by major Arctic 

rivers [1]. 

Substrate—Mud or clay and stone bottoms [2]. 

Physical/chemical—Temperature: -1.4–3 °C [1, 5]. Salinity: Marine to slightly brackish; 27–32 ppt [1].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes. Oviparous. 

Spawning season—Unknown. 

Fecundity—32–38 orange eggs [1].
White Sea Eelpout 447

Food and Feeding

Food items—Unknown. 

Trophic level—3.45 standard error 0.45 [6].

Biological Interactions

Predators—Unknown. 

Competitors—Unknown. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [6].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, White Sea Eelpout are not commercially harvested.

Potential Effects of Climate Change

Unknown.

Areas for Future Research [B]

Virtually nothing is known about this species from the region. Research needs include: (1) depth and location of 

pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators. 
448 Alaska Arctic Marine Fish Ecology Catalog

Bibliography

1. Møller, P.R., 2000, Restoration of the taxon Lycodes marisalbi, with notes on its disjunct Arctic distribution: Journal of Fish 

Biology, v. 57, no. 6, p. 1,404–1,415.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Saddled Eelpout 449

Saddled Eelpout (Lycodes mucosus) 

Richardson, 1855

Family Zoarcidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Due to a lack of data on this species, it is 

difficult to make an assessment of its ecological role. However, it is 

commonly encountered in the U.S. Chukchi Sea and therefore may 

be of moderate significance in marine ecosystem functioning there. 

Physical Description/Attributes: Elongate, cream-colored body 

with dark brown U- and Y-shaped marks and white underside. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 708) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [2, 3]. Elsewhere in Alaska, south in Bering Sea to near St. Matthew I. (60°N) [3]. 

Worldwide, Canadian High Arctic eastward to Hudson Strait and northwest Greenland; Russian Arctic southward to Gulf of 

Anadyr [3]. 

Relative Abundance: Common in U.S. Chukchi Sea and U.S. Beaufort Sea [3].

Saddled Eelpout (Lycodes mucosus), 229 mm TL, northeastern 

Bering Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Saddled Eelpout (Lycodes mucosus) within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections 

[1, 3, 5].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Saddled Eelpout

Lycodes mucosus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
450 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 0–180 m, typically less than 80 m [3]. Documented in U.S. Chukchi Sea 20–59 m [2, 6, 7]. In 

general, eelpout spawning and larvae occur at the same depths that adults inhabit [1].

Benthic and reproductive distribution of Saddled Eelpout (Lycodes mucosus).

3914SddldElt

Open ocean Shore

Depth, in meters

250

200

150

100

50

0

250

200

150

100

50

0

Juveniles and adults

Lycodes mucosus

Saddled Eelpout

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning, 

eggs, and larvae

Depths documented in U.S. Chukchi Sea

Reproductive distribution

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown. 

Overall depth data from outside U.S. Beaufort-Chukchi Seas

Some fish to 

 180 m

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

Habitats and Life History

Eggs—Size: 3 mm for almost ripe eggs [2, 8]. Time to hatching: Unknown. Habitat: Benthic [1]. Larvae—Size 

at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile transformation: Unknown. 

Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Age unknown. 24–35 cm TL for females [8]. Maximum age: Unknown. 

Maximum size: About 49 cm TL [1, 8]. Habitat: Benthic [1], sometimes at entrance to burrows or under debris 

[9]. Have been found in intertidal rock pools in Greenland [3]. 

Substrate—Mostly on mud, also found on shell hash, mixed gravel, sand and mud, and gravel and rock [2]. 

Physical/chemical—Temperature: -1.9–10.5 °C [2, 9]. Salinity: Marine. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes. Oviparous. 

Spawning season—In Russia, likely in the autumn [8]. 

Fecundity—Unknown. 
Saddled Eelpout 451

Food and Feeding

Food items—Mysids, copepods, ostracods, polychaetes [10]. 

Trophic level—3.92 standard error 0.64 [11].

Biological Interactions

Predators—Unknown. 

Competitors—Likely such benthic microcarnivores as flatfishes, sculpins, snailfishes, and other eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [11].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Saddled Eelpout are not commercially harvested.

Potential Effects of Climate Change

The Saddled Eelpout is an Arctic species. Shifts in range and abundance are possible effects. 

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators. 

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [8]

Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86. [9]
452 Alaska Arctic Marine Fish Ecology Catalog

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

7. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

8. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

9. Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86.

10. Green, J.M., 1983, Observations on the food of marine fishes from Resolute Bay, Cornwallis Island, Northwest Territories: 

Astarte, v. 12, no. 2, p. 63–68.

11. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Wattled Eelpout 453

Wattled Eelpout (Lycodes palearis)

Gilbert, 1896

Family Zoarcidae 

Colloquial Name: None within U.S. Chukchi and 

Beaufort Seas.

Ecological Role: Poorly understood, although based on studies 

outside the Arctic Ocean, may be of moderate importance as food 

for larger fishes. 

Physical Description/Attributes: Elongate brown or reddish brown body and fins with narrow, creamy white bands on upper 

body and dorsal fin; bands fade with growth. For specific diagnostic characteristics, see Fishes of Alaska (Meckenburg and 

others, p. 719) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea as far north as 71°17’N, 156°48’W near Barrow and western U.S. Beaufort Sea near Point Barrow 

at 71°22’N, 156°19’W [2]. Elsewhere, in Bering Sea, along the Aleutian Islands and Gulf of Alaska south to Oregon, and to 

southeastern Kamchatka Peninsula, Russia [2, 3].

Relative Abundance: Common as far north as northeastern U.S. Chukchi Sea [6]. Common at least in eastern Bering Sea [7]. 

Uncommon in southeastern Bering Sea [8] and rare in Sea of Japan [9].

Wattled Eelpout (Lycodes palearis), 152 mm TL, Chukchi Sea, 

2007. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Wattled Eelpout (Lycodes palearis) within Arctic OCS Planning Areas [4] based on 

review of published literature and specimens from historical and recent collections [1, 2, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Wattled Eelpout

Lycodes palearis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
454 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 2–925 m, almost always less than 200 m [10]. Taken in U.S. Chukchi Sea at 30–54 m [3, 6, 11]. In general, 

eelpout spawning and larvae occur at the same depths that adults inhabit [1].

Benthic and reproductive distribution of Wattled Eelpout (Lycodes palearis).

3915WldEl

Some fish to 925 m

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

1,000

800

600

400

200

0

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Overall depth data from outside

U.S. and Beaufort-Chukchi Seas

Lycodes palearis

Wattled Eelpout

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning, eggs, and larvae

Reproductive distribution

Depths documented in 

U.S. Chukchi Sea

Juveniles and adults

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 62.0 cm TL [12]. 

Habitat: Benthic [1]. 

Substrate—Mud or sand [1, 3, 13]. 

Physical/chemical—Temperature: –1.7–14.6 °C [3, 5, 13, 14]. Salinity: Marine [3, 13].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous. 

Spawning season—Unknown. 

Fecundity—Unknown.
Wattled Eelpout 455

Food and Feeding

Food items—In eastern Bering Sea and Gulf of Alaska, a wide range of benthic and epibenthic prey. Crustaceans 

(for example, gammarid amphipods, crabs, and shrimps) and fishes are often consumed, along with polychaetes, 

brittle stars, echiuroids, and clam siphons [15–17]. Larger Wattled Eelpouts feed more heavily on fishes than do 

smaller individuals [16]. 

Trophic level—3.6 [18].

Biological Interactions

Predators—Arrowtooth Flounder, Great Sculpin, Greenland Halibut, Pacific Cod, and Pacific Halibut in eastern 

Bering Sea, Gulf of Alaska, and off Kamchatka Peninsula, Russia [19–22]. 

Competitors—Likely a range of moderate-sized benthic fishes, including flatfishes, gadids, and sculpins.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (Preliminary K or Fecundity) [23].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Wattled Eelpout are not commercially harvested currently.

Potential Effects of Climate Change

The Wattled Eelpout is a predominantly Boreal Pacific species which would be expected to become more 

abundant in the Chukchi Sea and possibly expand its range farther into the Beaufort Sea.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators. 
456 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [13]

Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi Sea, 

Alaska: Fishery Bulletin, v. 95, no. 2 p. 195–209. [6]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [3]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

7. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

8. Acuna, E.I., Goddard, P., and Kotwicki, S., 2003, 2002 bottom trawl survey of the eastern Bering Sea continental shelf: 

National Marine Fisheries Service, Alaska Fisheries Science Center, Processed Report 2003-01.

9. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

10. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

11. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, nos. 1–2, p. 57–70.
Wattled Eelpout 457

12. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

13. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

14. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

15. Brodeur, R.D., and Livingston, P.A., 1988, Food habits and diet overlap of various Eastern Bering Sea fishes: National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS F/NWC-127, 76 p.

16. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

17. Yang, M.-S., Dodd, K., Hibpshman, R., and Whitehouse, A., 2006, Food habits of groundfishes in the Gulf of Alaska 

in 1999 and 2001: U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical 

Memorandum, NMFS-AFSC-164, 189 p.

18. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.

19. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.

20. Kihara, K., and Shimada, A.M., 1988, Prey-predator interactions of Pacific cod Gadus macrocephalus and water 

temperature: Nippon Suisan Gakkaishi, v. 54, no. 12, p. 2,085–2,088.

21. Lang, G.M., Livingston, P.A., Pacunski, R.E., Parkhurst, J., and Yang, M.-S., 1991, Groundfish food habits and predation on 

commercially important prey species in the eastern Bering Sea from 1984 to 1986: Seattle, Washington, U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFS F/NWC-207, 240 p.

22. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.

23. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
458 Alaska Arctic Marine Fish Ecology Catalog

Polar Eelpout (Lycodes polaris) 

(Sabine, 1824)

Family Zoarcidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Based on food habits research outside the 

U.S. Arctic [1, 2], Polar Eelpout may be of moderate ecological 

importance.

Physical Description/Attributes: Slender, elongate orange to light brown body with by 9–11 dark brown bands extending from 

sides onto dorsal fin. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 710) [3]. 

Swim bladder: Absent [3]. Antifreeze glycoproteins in blood serum: Present [4].

Range: U.S. Chukchi and Beaufort Seas (in Chukchi Sea, northward to at least 73°32’N) [5]. Elsewhere in Alaska, southward 

in eastern Bering Sea to between Hall Island and St. Lawrence Island. Worldwide, nearly circumpolar in Arctic seas; in western 

Bering Sea southward to Glubokaya Bay on the Koryak Coast, Russia (about 61°N) [5–7] and in the western North Atlantic to 

the Gulf of St. Lawrence [7].

Relative Abundance: Common in both U.S. Chukchi and Beaufort Seas [5, 10, 11], and eastward to at least Cornwallis Island, 

Nunavut, Canada [12, 13]. 

Polar Eelpout (Lycodes polaris), 180 mm TL, Chukchi Sea, 

2004. Photograph by C.W. Mecklenburg and B.A. Sheiko, Point 

Stephens Research and Russian Academy of Sciences.

Geographic distribution of Polar Eelpout (Lycodes polaris) within Arctic OCS Planning Areas [8] based on 

review of published literature and specimens from historical and recent collections [3, 7, 9]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Polar Eelpout

Lycodes polaris

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Polar Eelpout 459

Depth Range: 0–333 m [14], typically 30–150 m [7]. Taken in U.S. Chukchi and Beaufort Seas from 17 to 183 m [9]. In 

general, eelpout spawning and larvae occur at the same depths that adults inhabit [3].

Benthic and reproductive distribution of Polar Eelpout (Lycodes polaris).

3916PlElt

Juveniles and adults

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

 Overall depth data from 

outside U.S. Beaufort-Chukchi Seas

Spawning, eggs, and larvae

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Lycodes polaris

Polar Eelpout

Benthic distribution

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Reproductive distribution

Some fish to 333 m

Depths documented in 

U.S. Chukchi Sea

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

No data for this species

Habitats and Life History

Eggs—Time to hatching: Unknown. Size: Large (about 4–5 mm) [6]. Habitat: Benthic [3]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [3]. Juveniles: Age and size: Unknown. Habitat: Benthic [3]. 

Adults—Age and size at first maturity: Age unknown. Probably around 15.5 cm TL [10]. Maximum age: At least 

5 years [10]. Maximum size 33.3 cm in U.S. Beaufort Sea [9]. Habitat: Benthic [3]. 

Substrate—Mud and sandy mud [5, 6, 12]. 

Physical/chemical—Temperature:-1.9–2.5 °C [5, 12], usually 0 °C or less [6]. Salinity: Predominantly marine, 

occasionally estuarine (as low as 25.3 ppt) [6].

Behavior

Diel—Lives in burrows in upper layers of sediment and has been observed entering them tail first [6, 12]. 

Seasonal— Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous. 

Spawning season—Likely in autumn or early winter [6, 10]. 

Fecundity—66–200 eggs [1, 6, 10].
460 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Primarily gammarid amphipods and polychaetes, along with smaller amounts of cumaceans, 

caprellid amphipods, isopods, mysids, shrimps, brittle stars, and occasionally fishes [10], including Arctic Cod 

[1]. A similar diet has been reported from Russian Arctic [6]. 

Trophic level—3.32 standard error 0.38 [16].

Biological Interactions

Predators—Seabirds [1]. Commonly eaten by bearded seals in Canadian High Arctic [2]. 

Competitors—Likely other benthic and epibenthic feeding fishes, including flatfishes, sculpins, and other 

eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [16].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Polar Eelpout are not commercially harvested currently.

Potential Effects of Climate Change

The Polar Eelpout is an Arctic species inhabiting the continental shelf and upper slope. It is possible that climate 

warming could cause extirpation of local populations as distributions shift to more northern, deeper, and colder 

waters.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.

Remarks

One of the most widespread, abundant, and northerly distributed continental shelf eelpouts [1].
Polar Eelpout 461

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [6]

Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western Beaufort 

Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA Technical Report 

NMFS-SSRF-764, 22 p. [10]

Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86. [12]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [5]

Bibliography

1. Mecklenburg, C.W., and Mecklenburg, T.A., 2011, Polar Eelpout—Lycodes polaris (Sabine, 1824): Arctic Ocean Diversity 

Web site, accessed 2011, at http://www.arcodiv.org/Fish/Lycodes_polaris.html.

2. Finley, K.J., and Evans, C.R., 1983, Summer diet of the bearded seal (Erignathus barbatus) in the Canadian High Arctic: 

Arctic, v. 36, no. 1, p. 82–89.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Power, G., 1997, A review of fish ecology in Arctic North America: American Fisheries Society Symposium, no. 19, 

p. 13–39.

5. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

6. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

7. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

8. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

9. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

10. Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and invertebrates trawled in the northeastern Chukchi and western 

Beaufort Seas 1976–1977: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, AA 

Technical Report NMFS-SSRF-764, 22 p.

11. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2 p. 195–209.
462 Alaska Arctic Marine Fish Ecology Catalog

12. Green, J.M., and Steele, D.H., 1975, Observations on marine life beneath sea ice, Resolute Bay, N.W.T., in Proceedings of 

the Circumpolar Conference on Northern Ecology, September 15–18, 1975: Ottawa, Ontario, National Research Council of 

Canada, p. II-77–II-86.

13. Arctic Laboratories Limited, 1987, Beaufort ocean dumpsite characterization: Prepared for Environment Protection, 

Conservation and Protection, Environment Canada, Yellowknife, Northwest Territories, by Arctic Laboratories Limited and 

LGL Limited.

14. Christiansen, J.S., ed., 2003, TUNU-1 Expedition—The fish fauna of the NE Greenland fjord systems—Technical report: 

Tromsø, Norway, University of Tromsø, Norwegian College of Fishery Science, Institute of Aquatic Resources, 33 p.

15. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

16. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Marbled Eelpout 463

Marbled Eelpout to Banded Gunnel

Marbled Eelpout (Lycodes raridens)

Taranetz & Andriashev, 1937

Family Zoarcidae 

Colloquial Name: None within U.S. Chukchi

and Beaufort Seas.

Ecological Role: Common but not abundant, the Marbled Eelpout is 

likely of relatively small ecological importance in the U.S. Chukchi 

and Beaufort Seas.

Physical Description/Attributes: Elongate cream to tan body with 

7–9 brown or reddish brown bands extending onto dorsal and anal fins; blackish dorsal fin margin at ends of the bands; head, 

nape, and body bands becoming marbled in appearance in adults. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 711) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas. Northernmost record in U.S. Chukchi Sea is at 71°27’N, 158°02’W, and in U.S. 

Beaufort Sea is at 71°13.5’N, 152°47.9’W; easternmost record in U.S. Beaufort Sea is north of Arey Island at 70°36’N, 

143°55’W [2]. Elsewhere in Alaska, eastern Bering Sea to Bristol Bay and one record north of Near Islands, western Aleutian 

Islands [1]. Worldwide, in western Bering Sea and Commander Islands to Okhotsk Sea [1] and in East Siberian Sea [2].

Relative Abundance: Common in U.S. Chukchi and Beaufort Seas [2, 4]. Patchily abundant (common) from eastern Sakhalin 

Island, Russia through Bering Sea and to Chukchi Sea and Beaufort Seas [2, 4–9].

Marbled Eelpout (Lycodes raridens), 102 mm, Chukchi Sea, 

2007. Photograph by C.W. Mecklenburg, Point Stephens 

Research. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Marbled Eelpout

Lycodes raridens

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Marbled Eelpout (Lycodes raridens) within Arctic Outer Continental Shelf Planning 

Areas [3] based on review of published literature and specimens from historical and recent collections [1, 2, 4]. 
464 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 8–467 m, rarely deeper than 150 m [4]. Taken in U.S. Chukchi Sea between 55–59 m [9]. A maximum depth of 

525 m was reported for fish wintering off shelf in Sea of Okhotsk [10]. Generally, eelpout spawning and larvae occur at same 

depths that adults inhabit [1].

Benthic and reproductive distribution of Marbled Eelpout (Lycodes raridens). 

Juveniles and adults

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

 Overall depth data from 

outside U.S. Beaufort-Chukchi Seas

Spawning, eggs, and larvae

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Lycodes raridens

Marbled Eelpout

Benthic distribution

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Reproductive distribution

Some fish to 360 m

Winters down to 525 m

Depth documented in 

U.S. Chukchi Sea

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: 7 years in western Bering Sea [7]. Maximum 

size: To 86 cm TL [7]. Habitat: Benthic [1]. 

Substrate—Sandy mud and mud [1, 11]. 

Physical/chemical—Temperature: -1.7–7.9 °C [7, 11, 12]. Salinity: Marine, for example, 32.58 parts per 

thousand in the U.S. Chukchi Sea [9] and occasionally in brackish waters [4].

Behavior

Diel—Unknown. 

Seasonal—Migrates to deeper water in Sea of Okhotsk in winter [10]. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous. 

Spawning season—Likely in autumn or autumn-winter in western Bering Sea [7]. 

Fecundity—Unknown.
Marbled Eelpout 465

Food and Feeding

Food items—Benthic and epibenthic prey, such as gammarid amphipods, euphausiids, shrimps, polychaetes, and 

clams [7]. 

Trophic level—3.6 [13].

Biological Interactions

Predators—Unknown. 

Competitors—Likely other benthic-dwelling fishes, including flatfishes, sculpins, snailfishes, and other eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Fecundity=3,116) [14].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Marbled Eelpout are not commercially harvested.

Potential Effects of Climate Change

The Marbled Eelpout is a predominantly Boreal Pacific species. Increases in abundance and interspecific 

competition are possible outcomes of climate warming.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [11]

Balanov, A.A., Badaev, O.Z., Napazakov, V.V., and Chuchukalo, V.I., 2006, Distribution and some biological features of Lycodes 

raridens (Zoarcidae) in the western part of the Bering Sea: Journal of Ichthyology, v. 46, no. 2, p. 148–155. [7]
466 Alaska Arctic Marine Fish Ecology Catalog

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [9]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at http://

caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

6. Kim, S.T., and Shepeleva, O.N., 2001, The structure of shelf ichthyocenoses of northeastern Sakhalin and Terpeniya Bay: 

Journal of Ichthyology, v. 41, no. 9, p. 711–722.

7. Balanov, A.A., Badaev, O.Z., Napazakov, V.V., and Chuchukalo, V.I., 2006, Distribution and some biological features of 

Lycodes raridens (Zoarcidae) in the western part of the Bering Sea: Journal of Ichthyology, v. 46, no. 2, p. 148–155.

8. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

9. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

10. Kim, S.T., 2001, Winter migrations of shelf fish to the continental slope: Journal of Ichthyology, v. 41, no. 8, p. 564–574.

11. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

12. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

13. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.

14. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Arctic Eelpout 467

Arctic Eelpout (Lycodes reticulatus)

Reinhardt, 1835

Family Zoarcidae 

Note: Morphological differences between L. reticulatus 

(Reinhardt, 1835) and L. rossi are not clear, making 

identifications and geographic ranges uncertain [1]. 

Colloquial Name: None within U.S. Chukchi and Beaufort 

Seas.

Ecological Role: Because of the lack of taxonomic clarity, uncommon occurrence, and paucity of life history information, the 

ecological role of this species was not evaluated. 

Physical Description/Attributes: Brown body with dark bands that are reticulate in large individuals; light spots on upper side 

of head. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 713) [2]. Swim bladder: 

Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Beaufort Sea [3]. Elsewhere, Arctic Canada to Greenland and east to Barents, Kara, and Laptev Seas [2].

Relative Abundance: Apparently common in U.S. Beaufort Sea. Abundance estimates are unreliable because of lack of 

sampling in the offshore waters, and the species is likely common, at least locally, considering the 2012 archived voucher 

specimens from at least five stations [3]. Common in Barents Sea [5].

Arctic Eelpout (Lycodes reticulatus), 306 mm, Beaufort Sea, 2011. 

Photograph by C.W. Mecklenburg, Point Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Eelpout

Lycodes reticulatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Arctic Eelpout (Lycodes reticulatus) within Arctic Outer Continental Shelf Planning 

Areas [4], based on review of published literature and specimens from historical and recent collections [1-3]. 
468 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Taken offshore at 6–930 m in U.S. Beaufort Sea [3]. 20–930 m, usually 380 m or shallower [1, 2]. Generally, 

eelpout spawning and larvae occur at same depths that adults inhabit [2].

Benthic and reproductive distribution of Arctic Eelpout (Lycodes reticulatus).

Some fish to 930 m

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

1,000

800

600

400

200

0

Data from outside U.S. Beaufort-Chukchi Seas

Lycodes reticulatus

Arctic Eelpout

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning, eggs, and larvae

Reproductive distribution

Juveniles and adults

Juveniles

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [2]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [2]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [2]. 

Adults—Age and size at first maturity: Unknown. Maximum age: 19 years [6]. Maximum size: 76.0 cm [2]. 

Habitat: Benthic, most often on outer shelf [1]. 

Substrate—Sand to mud [2]. 

Physical/chemical—Temperature: -1.5–4.5 °C [7]. Salinity: Marine [2].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes. Oviparous [8]. 

Spawning season—Unknown. 

Fecundity—Unknown.
Arctic Eelpout 469

Food and Feeding

Food items—A wide variety of benthic invertebrates (amphipods, bivalves, brittle stars, gastropods, 

echinoderms, crustaceans, and polychaetes) and fishes and pelagic crustaceans (euphausids) [6, 9–11]. 

Trophic level—3.5 ±0.53 standard error [12].

Biological Interactions

Predators—Bearded seals in northeastern Canada [13]. 

Competitors—Likely other benthic microcarnivores including some sculpins, flatfishes, snailfishes, and other 

eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [12].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Arctic Eelpout are not commercially harvested.

Potential Effects of Climate Change

Arctic Eelpouts have fairly wide depth and temperature tolerances. Thus, it is difficult to speculate about the 

effects of climate warming. Changes in temperature, species composition of fish assemblages, and productivity 

can be expected to affect distribution and abundance patterns

Areas for Future Research [B]

Little is known about this species biology and ecology from the region. 

Research needs include: 

(1) Depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment; (3) preferred 

depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic movements; (6) population studies; 

(7) prey; and (8) predators.
470 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [1]

Bibliography

1. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at http://

caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

6. von Dorrien, C.F., 1993, Ecology and respiration of selected Arctic benthic fish species: Berichte zur Polarforschung, v. 125, 

118 p. [In German] 

7. Møller, P.R., and Jørgensen, O.A., 2000, Distribution and abundance of eelpouts (Pisces, Zoarcidae) off West Greenland: 

Sarsia, v. 85, no. 1, p. 23–48. 

8. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

9. Chambers, C.A., and Dick, T.A., 2005, Trophic structure of one deep-sea fish community in the eastern Canadian Arctic—

Application of food, parasites and multivariate analysis: Environmental Biology of Fishes, v. 74, nos. 3–4, p. 365–378.

10. Torres, P., Rodriquez-Marin, E., and Loureiro, I., 2000, Preliminary results from feeding analysis for the most abundant 

demersal fishes in Flemish Cap during summer (1993–2000): Northwest Atlantic Fisheries Organization, Serial No. N4302, 

NAFO SCR Doc. 00/60.

11. Román, E., González, C., and Ceballos, E., 2004, Food and feeding of most abundant fish species in Flemish Cap: 

Northwest Atlantic Fisheries Organization, Serial No. N5018, NAFO SCR Doc. 04/58.

12. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

13. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.
Threespot Eelpout 471

Threespot Eelpout (Lycodes rossi) 

Malmgren, 1865

Family Zoarcidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Note on taxonomy: Morphological differences 

between L. reticulatus (Reinhardt, 1835) and L. rossi are not clear, 

making identifications and geographic ranges uncertain [1]. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Uncertain. The life history and ecology of this 

species and its roles in regional ecosystems and food webs are 

poorly understood. Issues regarding the taxonomy of this species need to be resolved.

Physical Description/Attributes: Brown, with dark brown bands on body and dorsal fin; light band across top of head 

connecting gill openings, often broken into spots. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and 

others, 2002, p. 713) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Beaufort Sea, at least between 152° and 155°W [2]. Elsewhere, from Greenland and Norwegian Seas to Kara Sea, 

including Iceland, Svalbard (Norway), and Barents Sea; and Canadian Beaufort Sea eastward to Dease Strait [1, 2].

Relative Abundance: Rare in U.S. Beaufort Sea [2, 4]. Common in Barents Sea [5].

Threespot Eelpout (Lycodes rossi), Beaufort Sea, 2008. 

Photograph by E. Akuna, National Oceanic and Atmospheric 

Administration, Alaska Fisheries Science Center.

Geographic distribution of Threespot Eelpout (Lycodes rossi), within Arctic Outer Continental Shelf Planning 

Areas [3] based on review of published literature and specimens from historical and recent collections [1, 2, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Threespot Eelpout

Lycodes rossi

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
472 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Adults at 42–365 m, usually deeper than 130 m [1, 2, 6]. Juveniles as shallow as 9 m [1, 6]. Generally, eelpout 

spawning and larvae occur at same depths that adults inhabit [1].

Benthic and reproductive distribution of Threespot Eelpout (Lycodes rossi).

Some fish to 930 m

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

1,000

800

600

400

200

0

Data from outside U.S. Beaufort-Chukchi Seas

Lycodes rossi

Threespot Eelpout

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning, eggs, and larvae

Reproductive distribution

Juveniles and adults

Juveniles

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

Habitats and Life History

Eggs—Size: 3–4 mm [5]. Time to hatching: Unknown. Habitat: Benthic [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: 10 years [5]. Maximum size: 38 cm TL [7]. 

Habitat: Benthic, most often on outer shelf and upper slope [2]. 

Substrate—Muddy [1, 8]. 

Physical/chemical—Temperature: -1.5–1.1 °C [8]. Found mainly at minus temperatures [6, 8]. Salinity: 27.6–

35.0 ppt [8]; prefers high salinity, usually not found in less than 34 ppt [6].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes. Oviparous [9]. 

Spawning season—Probably winter or early spring in Barents Sea [5]. 

Fecundity—390 eggs [5]. 
Threespot Eelpout 473

Food and Feeding

Food items—Polychaetes, copepods, amphipods, cumaceans [6, 8]. 

Trophic level—3.49 ±0.53 standard error [10].

Biological Interactions

Predators—Bearded seals in Ungava Bay, Canada [11]. 

Competitors—Likely other benthic microcarnivores including some sculpins, flatfishes, snailfishes, and other 

eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [10].

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Threespot Eelpout are not commercially harvested.

Potential Effects of Climate Change

Because Threespot Eelpout are mainly a deep-water, slope-dwelling Arctic species and little is known about 

climate change at slope depths in the Arctic, potential effects on this species cannot be estimated.

Areas for Future Research [B]

Little is known about this species biology and ecology from the region. Research needs include: (1) depth and 

location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred 

depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, 

(6) population studies, (7) prey, and (8) predators. 
474 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [6]

McAllister, D.E., Anderson, M.E., and Hunter, J.G., 1981, Deep-water eelpouts, Zoarcidae, from Arctic Canada and Alaska: 

Canadian Journal of Fisheries and Aquatic Sciences, v. 38, no. 7, p. 821–839. [8]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at http://

caff.is/monitoring-series/370-pacific-arcticmarine-fishes. 

5. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

6. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

7. Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic marine fishes of Canada: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, no. 2674, 112 p.

8. McAllister, D.E., Anderson, M.E., and Hunter, J.G., 1981, Deep-water eelpouts, Zoarcidae, from Arctic Canada and Alaska: 

Canadian Journal of Fisheries and Aquatic Sciences, v. 38, no. 7, p. 821–839.

9. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

10. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

11. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.
Archer Eelpout 475

Archer Eelpout (Lycodes sagittarius)

McAllister, 1976

Family Zoarcidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Likely to be absent over shelve habitats of the 

U.S. Beaufort Sea. Survey data suggest that this species rarely 

occurs in slope and deep water habitats. The ecological role of the 

species is probably minimal.

Physical Description/Attributes: Elongate, narrow body colored 

uniformly dark brown; peritoneum black. For specific diagnostic 

characteristics see Fishes of Alaska (Mecklenburg and others, 2002, p. 717) [1]; specimens with white bars and fewer vertebrae 

illustrated therein [1] and in previous publications, (for example, McAllister and others, 1981) [2], are now known to belong to 

L. marisalbi [3]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Beaufort Sea, and Beaufort Sea of western Canada and Kara and Laptev Seas [5].

Relative Abundance: Apprentlly common in Beaufort Sea off Alaska [5].

Archer Eelpout (Lycodes sagittarius), 180 mm TL or less 

from Franklin Bay, Northwest Territories, Canada, (from 

Mecklenburg and others 2002, p. 717).

Geographic distribution of Archer Eelpout (Lycodes sagittarius), within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [3, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Archer Eelpout

Lycodes sagittarius

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
476 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 357–600 m in U.S. Beaufort Sea [2, 3] otherwise depths of 120 to at least 1,934 m [5]. Generally, eelpout 

spawning and larvae occur at same depths that adults inhabit [1].

Benthic and reproductive distribution of Archer Eelpout (Lycodes sagittarius).

Some fish to 600 m

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

1,000

800

600

400

200

0

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

Data from U.S. Beaufort Sea

Lycodes sagittarius

Archer Eelpout

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning, eggs, and larvae

Reproductive distribution

Juveniles and adults

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: To 27.8 cm TL [2]. 

Habitat: Benthic [1]. 

Substrate—Mud [2]. 

Physical/chemical—Temperature: Range unknown. Taken at –0.9 °C [2]. Salinity: Marine, taken at 34 ppt. [2].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous. 

Spawning season—Likely late summer or early autumn [2]. 

Fecundity—Unknown. 
Archer Eelpout 477

Food and Feeding

Food items—Annelids, bivalves, gastropods, and crustaceans [2]. 

Trophic level—3.22 ±0.40 standard error [6].

Biological Interactions

Predators—Unknown. 

Competitors—Likely other benthic microcarnivorous fishes, including snailfishes, sculpins, flatfishes, and other 

eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [6]. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Archer Eelpout are not commercially harvested.

Potential Effects of Climate Change

Unknown. The Archer Eelpout is an endemic species occurring rarely in deep water areas of the U.S. Beaufort 

Sea [3]. Lack of information about this species and understanding of its role in this benthic ecosystem precludes 

an informed assessment of potential climatic effects. 

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators. 
478 Alaska Arctic Marine Fish Ecology Catalog

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. McAllister, D.E., Anderson, M.E., and Hunter, J.G., 1981, Deep-water eelpouts, Zoarcidae, from Arctic Canada and Alaska: 

Canadian Journal of Fisheries and Aquatic Sciences, v. 38, no. 7, p. 821–839.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at http://

caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Longear Eelpout 479

Longear Eelpout (Lycodes seminudus) 

Reinhardt, 1837

Family Zoarcidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Because of its apparent rarity in shallow U.S. 

Arctic waters, this species is likely of little ecological importance 

in the U.S. Chukchi and Beaufort Sea. 

Physical Description/Attributes: Elongate, dark gray-brown body, sometimes with short, pale bands above lateral line and 

extending onto dorsal fin. For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 707) 

[1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: Continental slope of U.S. Chukchi and Beaufort Seas [2]. Elsewhere, in Baffin Bay off Canada and northwest 

Greenland, northeast Greenland, off northern Iceland to Faroe Islands, Svalbard Archipelago, northern Barents Sea, and Kara 

Sea [2, 3].

Relative Abundance: Rare in shelf waters of U.S. Chukchi and Beaufort Seas, but potentially more common in deeper waters 

over the continental slope [2, 5]. Common off Greenland [2].

Longear Eelpout (Lycodes seminudus), 238 mm TL, Chukchi 

Borderland, 2009. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Longear Eelpout (Lycodes seminudus) within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [1, 2, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Longear Eelpout

Lycodes seminudus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
480 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 50–1,400 m, mostly 200–600 m [2, 6]. In general, eelpout spawning and larvae occur at same depths that adults 

inhabit [1].

Benthic and reproductive distribution of Longear Eelpout (Lycodes seminudus).

Open ocean

Depth, in meters

Juveniles and adults

Lycodes seminudus

Longear Eelpout

Benthic distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Some fish to 1,400 m

Open ocean No data for this species Shore

Depth, in meters

Spawning, eggs, and larvae

Reproductive distribution

1,500

1,200

900

600

300

0

1,500

1,200

900

600

300

0

 Data from outside

U.S. Beaufort-Chukchi Seas

Depth range of spawning is

uncertain. Potential location of

spawning, eggs, and larvae is

based on depth range of benthic

individuals 

Habitats and Life History

Eggs—Time to hatching: Unknown. Size: Unknown. Habitat: Benthic [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 56 cm TL [3]. 

Habitat: Benthic [1]. 

Substrate—Mud or mud-clay [7]. 

Physical/chemical—Temperature: -1.7–4.9 °C [5]. In Russia, almost exclusively at less than 0 °C [6]. Salinity: 

Marine, high salinity [6].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous. 

Spawning season—Unknown. 

Fecundity—Unknown. 
Longear Eelpout 481

Food and Feeding

Food items—Amphipods, shrimps, isopods, polychaetes [6]. 

Trophic level—3.45 standard error 0.44 [8].

Biological Interactions

Predators—Unknown. 

Competitors—Likely other benthic microcarnivores, including flatfishes, snailfishes, and other eelpouts.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time: 4.5–14 years (Preliminary K or Fecundity) [8].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Longear Eelpout are not commercially harvested.

Potential Effects of Climate Change

Unknown. The Longear Eelpout is an endemic Arctic species occurring most commonly in deep water habitats 

over the slope [2]. Not enough is known about this species to predict climate change effects.

Areas for Future Research [B]

Little is known about biology and ecology of this species from the region. Research needs include: (1) depth and 

location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred 

depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, 

(6) population studies, (7) prey, and (8) predators. 
482 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [6]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Møller, P.R., and Jørgensen, O.A., 2000, Distribution and abundance of eelpouts (Pisces, Zoarcidae) off West Greenland: 

Sarsia, v. 85, no. 1, p. 23–48.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

7. McAllister, D.E., Anderson, M.E., and Hunter, J.G., 1981, Deep-water eelpouts, Zoarcidae, from Arctic Canada and Alaska: 

Canadian Journal of Fisheries and Aquatic Sciences, v. 38, no. 7, p. 821–839.

8. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Scalebelly Eelpout 483

Scalebelly Eelpout (Lycodes squamiventer) 

Jensen, 1904

Family Zoarcidae 

Note: Except for geographic range data, all information is from areas 

outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: This species generally does not occur on the 

continental shelves in the Arctic Ocean but likely has some ecological 

significance in the relatively unexplored, deep waters of the U.S. 

Chukchi and Beaufort Seas.

Physical Description/Attributes: Light to dark grayish brown body without bands or other marks. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 722)[1]. Swim bladder: Absent [1]. Antifreeze glycoproteins 

in blood serum: Unknown.

Range: U.S. Beaufort Sea [2]. Elsewhere, Canadian Beaufort Sea, Davis Strait off western Greenland; Greenland Sea, Norwegian 

Sea, Kara Sea, and off Faeroese-Shetland slope, Barents Sea [1, 3].

Relative Abundance: Common in U.S. Beaufort Sea and western Canada Beaufort Sea. Although known from only a few records, 

this species could be more common on the slope as has been reported elsewhere in the Arctic [2]. Common in Norwegian Sea and 

around Håkon Mosby Mud Volcano in Barents Sea [5]. 

Scalebelly Eelpout (Lycodes squamiventer), 358 mm, Beaufort 

Sea, 2012. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Scalebelly Eelpout (Lycodes squamiventer) within Arctic Outer Continental Shelf 

Planning Areas [4] based on review of literature and specimens from historical and recent collections [1-3]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Scalebelly Eelpout

Lycodes squamiventer

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
484 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Benthic, 160–1,808 m [2], typically from 1,273–1,546 m in Norwegian Sea [5]. Reported but not confirmed as 

shallow as 160 m [6]. Records from U.S. Beaufort Sea are depths of 357–500 m [2]. In general, eelpout spawning and larvae 

occur at the same depths that adults inhabit [7].

Benthic and reproductive distribution of Scalebelly Eelpout (Lycodes squamiventer). 

Open ocean

Depth, in meters

2,000

1,600

1,200

800

400

0

Adults and juveniles

Lycodes squamiventer

Scalebelly Eelpout

Benthic distribution

Some fish to 1,808 m

Overall benthic depth range. 

Depth, in meters

Specific depth range of either 

juveniles and adults is known

Open ocean No data for this species

Depths documented in

U.S. Beaufort Sea

Shore Shore

Spawning, eggs, and larvae

Reproductive distribution

Depth range of spawning is

uncertain. Potential location of

spawning, eggs, and larvae is based

on depth range of benthic individuals

Overall depth data from 

outside U.S. Beaufort-Chukchi Seas 2,000

1,600

400

800

1,200

0

Habitats and Life History

Eggs—Size: 3.5 mm [5]. Time to hatching: Unknown. Habitat: Benthic [1, 5]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Age unknown. Females at 14 cm, males at 17.9 cm [6]. Maximum age: 

21 years [5]. Maximum size: 37 cm TL [2]. Habitat: Benthic, in deep waters on the slope [3, 5, 6]. Substrate—

Muddy bottoms [1]. 

Physical/chemical—Temperature: -1.2–0.6 °C [5]. Salinity: Marine. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes. Oviparous [8]. 

Spawning season—Autumn, but ripe females have been found in June [5, 6]. 

Fecundity—50–60 eggs [5, 6].
Scalebelly Eelpout 485

Food and Feeding

Food items—Various benthic species such as pogonophores, gastropods, amphipods, polychaetes, copepods, 

ophiuroids, bivalves, and crustaceans [5, 6]. 

Trophic level—3.4 ±0.4 standard error [9].

Biological Interactions

Predators—Unknown. 

Competitors—Likely other benthic microcarnivores including some sculpins, flatfishes, snailfishes, and other 

eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [9].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Scalebelly Eelpout are not commercially harvested.

Potential Effects of Climate Change

Unknown. The Scalebelly Eelpout is a predominantly Arctic, slope and deep-water species. Not enough 

information is available to evaluate potential climatic effects. 

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.
486 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Hildebrandt, N., Bergmann, M., and Knust, R., 2011, Longevity and growth efficiency of two deep-dwelling Arctic zoarcids and 

comparison with eight other zoarcid species from different climatic regions: Polar Biology, v. 34, no. 10, p. 1,523–1,533. [5]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, A., 

Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 2011, 

Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p. [6]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Hildebrandt, N., Bergmann, M., and Knust, R., 2011, Longevity and growth efficiency of two deep-dwelling Arctic 

zoarcids and comparison with eight other zoarcid species from different climatic regions: Polar Biology, v. 34, no. 10, 

p. 1,523–1,533.

6. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

7. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

8. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

9. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Estuarine Eelpout 487

Estuarine Eelpout (Lycodes turneri) 

Bean, 1879

Family Zoarcidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown. Could be important in coastal food 

webs as this eelpout could be prey for piscivorous fishes and birds 

of the nearshore community.

Physical Description/Attributes: Elongate, body with 10–12 

bands. Color in adults is purple with bluish white bands bordered with purplish olive, or umber with creamy white bands and 

dark umber borders. Juveniles are creamy white with blackish bordered brown bands. For specific diagnostic characteristics, 

see Fishes of Alaska (Mecklenburg and others, 2002, p. 709) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood 

serum: Unknown.

Range: U.S. Chukchi Sea and western U.S. Beaufort Sea in vicinity of Point Barrow. Elsewhere in Alaska, southward in eastern 

Bering Sea to Bristol Bay. Worldwide, southward in western Bering Sea to Cape Olyutorskiy, Russia [2].

Relative Abundance: Uncommon in U.S. Chukchi and Beaufort Seas [2, 5, 6]. A record from the eastern U.S. Beaufort Sea off 

Point Franklin was recently shown to be in error because of a mistake in a museum catalog; the fish were actually captured in 

the Chukchi Sea [1].

Estuarine Eelpout (Lycodes turneri), juvenile, 145 mm TL, 

Chukchi Sea, 2009. Photograph by C.W. Mecklenburg, Point 

Stephens Research. Fish shown is a juvenile.

Geographic distribution of Estuarine Eelpout (Lycodes turneri) within Arctic Outer Continental Shelf Planning 

Areas [3] based on review of literature and specimens from historical and recent collections [1, 2, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Estuarine Eelpout

Lycodes turneri

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
488 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: 1–125 m, typically less than 50 m [2]. Generally, eelpout spawning and larvae occur at same depths that adults 

inhabit [1].

Benthic and reproductive distribution of Estuarine Eelpout (Lycodes turneri).

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

Lycodes turneri

Estuarine Eelpout

Benthic distribution

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

Reproductive distribution

Data from outside U.S. Beaufort-Chukchi Seas

Some fish

to 125 m

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Spawning, eggs, and larvae

No data for this species

Juveniles and adults

Depth range of spawning is uncertain.

Potential location of spawning, eggs,

and larvae is based on depth range of

benthic individuals 

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic [1]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Benthic [1]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 64 cm TL [7]. 

Habitat: Benthic [1], limited to inner and mid-shelf [5]. 

Substrate—Soft bottoms [1]. 

Physical/chemical—Temperature: Not reported. Salinity: Taken mostly in or near estuaries [4]. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Unknown. 

Spawning season—Unknown. 

Fecundity—Unknown.
Estuarine Eelpout 489

Food and Feeding

Food items—Unknown. 

Trophic level—3.36 standard error 0.44 [8]

Biological Interactions

Predators—Unknown. 

Competitors—Unknown. 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [8]. 

Traditional and Cultural Importance

Reported in the late 1800s to be an important food fish at Saint Michael, a village on the coast of Norton Sound 

in the Alaskan Bering Sea [9].

Commercial Fisheries

Currently, Estuarine Eelpout are not commercially harvested.

Potential Effects of Climate Change

Unknown. Range expansions are possible with expansion of brackish water conditions.

Areas for Future Research [B] Little is known about the biology and ecology of this species from the region. 

Research needs include: (1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-theyear benthic recruitment, (3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal 

and ontogenetic movements, (6) population studies, (7) prey, and (8) predators. 
490 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p. [5]. 

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

6. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

7. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

8. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

9. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.
Blackline Prickleback 491

Blackline Prickleback (Acantholumpenus mackayi) 

(Gilbert, 1896)

Family Stichaeidae

Note: Except for physical description, relative abundance, and 

geographic range data, all information is from areas outside of the 

study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Blackline Pricklebacks are uncommon in the U.S. 

Chukchi Sea and have not been reported from the U.S. Beaufort Sea. They are probably of relatively little ecological importance 

in U.S. Arctic waters.

Physical Description/Attributes: Elongate, compressed, slightly eel-like body colored yellow or brown with a dark line 

on back at base of dorsal fin and two dark broken lines below. Caudal fin is dark and unbanded. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 761) [1]. Swim bladder: Absent [2]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: Southern U.S. Chukchi Sea, documented by one record from Kotzebue Sound and one from northern Bering Strait [1, 

3]. No other records off Arctic Alaska, but presence is assumed from occurrence in Canadian Beaufort Sea off the Mackenzie 

Delta. Elsewhere in Alaska, in eastern Bering Sea, Aleutian Islands, and Gulf of Alaska. Worldwide, in Sea of Japan and Sea 

of Okhotsk, along Pacific coast of Hokkaido, Japan, and southeastern Kamchatka Peninsula, Russia; in Canadian Beaufort Sea 

between Phillips Bay, Yukon Territory, and Wood Bay, Northwest Territories, Canada [3, 4]. 

Blackline Prickleback (Acantholumpenus mackayi), 125 mm TL, 

Norton Sound, Bering Sea, 2004. Photograph by 

C.W. Mecklenburg, Point Stephens Research.
492 Alaska Arctic Marine Fish Ecology Catalog

Relative Abundance: Uncommon in U.S. Chukchi Sea with patchy distribution mainly in vicinity of river mouths and deltas 

[1]. Common in Sea of Japan and Sea of Okhotsk off Sakhalin Island, Russia [6, 7]. Common in Tuktoyaktuk Harbor and other 

brackish nearshore waters off the Mackenzie River Delta, Canadian Beaufort Sea [8].

Geographic distribution of Blackline Prickleback (Acantholumpenus mackayi) within Arctic Outer Continental 

Shelf Planning Area [5] based on review of published literature and specimens from historical and recent 

collections [1, 3, 4]. 

tac14-5222_fig3-10-8m_Acantholumpenus_mackayi

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Blackline Prickleback

Acantholumpenus mackayi

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Blackline Prickleback 493

Depth Range: 0.5–100 m, typically less than 50 m in Sea of Okhotsk [4]. In Alaska, documented from shallow water nearshore 

to depths of 66 m [4].

Benthic and reproductive distribution of Blackline Prickleback (Acantholumpenus mackayi).

3108BlkliPiklbk

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Juveniles and adults

Acantholumpenus mackayi

Blackline Prickleback

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown 

Larvae are pelagic. Specific depths unknown

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Data from outside U.S. Chukchi and Beaufort Seas

Some fish

to 100 m

Habitats and Life History

Eggs—Size: 1.0–1.4 mm in diameter [9]. Time to hatching: Unknown. Habitat: Benthic and adhesive [6, 10]. 

Larvae—Size at hatching: 15.8 mm [11]. Size at juvenile transformation: 21.5 mm [11]. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [12]. 

Juveniles—Age and size: Age unknown. From 21.5 to 30–40 cm TL [6, 11]. Habitat: Benthic [12]. 

Adults—Age and size at first maturity: Off Sakhalin Island in Sea of Japan, both males and females mature 

between 30 and 40 cm TL [6] and, assuming fishes in Northwest Territories, Canada, have similar growth rates, at 

around 6 years. Males may be larger at age than females; however, females may be heavier at length than males 

[11]. Maximum age: 6 years for males and 14 years for females in Northwest Territories [11]. Maximum size: 70 

cm SL. Habitat: Benthic [1–3, 6, 8]. 

Substrate—Sand, silt, or mud [1, 6, 8]. 

Physical/chemical—Temperature: 2.4–15.0 °C [4]. Salinity: Marine and brackish water (as low as 8 ppt) [6, 8].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Mature fish may move into shallow waters to spawn [6]. In general, prickleback adults brood 

their eggs [10]. 

Schooling—A non-schooling species [6]. 

Feeding—Feeds on bottom and occasionally in water column [13]. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [10]. 

Spawning season—Likely September off the Northwest Territories, Canada [13]. In Sea of Japan off Sakhalin 

Island, Russia, fish in post-spawning condition were observed June and July [6]. 

Fecundity—Unknown. 
494 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Primarily amphipods, oligochaetes, and polychaetes, and the occasional clam, copepod, mysids, 

snail, fish egg, and fish in Canadian Arctic [14, 15]. 

Trophic level—3.1 standard error 0.31 [16].

Biological Interactions

Predators—Off Kamchatka Peninsula, Russia, they are eaten by Great and Plain Sculpins [17]. 

Competitors—Unknown. Although likely other benthic species, such as smaller sculpins, eelpouts, and 

flatfishes.

1980 1990 2000 2010

Resilience

Very low, minimum population doubling time: more than 14 years (Preliminary K or Fecundity) [16]. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Blackline Prickleback are not commercially harvested.

Potential Effects of Climate Change

Blackline Prickleback are a predominantly Boreal species with an affinity for brackish waters, and could become 

more abundant in or expand into the U.S. Chukchi and Beaufort Seas as Arctic Ocean temperatures increase and 

the water freshens from increased ice melting.

Areas for Future Research [B]

Little is known about the ecology of this species. In particular, basic life history and habitat information is 

lacking; however, the species distribution and abundance in the region is limited, thus a need for directed studies 

is unwarranted at the present time.

References Cited

Houston, J., and McAllister, D.E., 1990, Status of the blackline prickleback, Acantholumpenus mackayi, in Canada: The 

Canadian Field-Naturalist, v. 104, no. 1, p. 24–28. [13]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]
Blackline Prickleback 495

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbor, N.W.T.: Canadian Technical Report of 

Fisheries and Aquatic Sciences, no.1218, 21 p. [11]

Shchetinnikov, A.S., 1983, Nutrition of Acantholumpenus mackayi (Stichaeidae) in the Gulf of Terpenium (Sakhalin Island): 

Journal of Ichthyology, v. 23, no. 6, p. 155–158. [6]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Shchetinnikov, A.S., 1983, Nutrition of Acantholumpenus mackayi (Stichaeidae) in the Gulf of Terpenium (Sakhalin Island): 

Journal of Ichthyology, v. 23, no. 6, p. 155–158.

7. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

8. Bond, W.A., and Erickson, R.N., 1993, Fisheries investigations in coastal waters of Liverpool Bay, Northwest Territories: 

Winnipeg, Manitoba, Canada Department of Fisheries and Oceans, Central and Arctic Region, Canadian Manuscript Report 

of Fisheries and Aquatic Sciences No. 2204, 59 p.

9. McAllister, D.E., 1975, Ecology of the marine fishes of Arctic Canada, in Proceedings of the Circumpolar Conference on 

Northern Ecology, September 15–18, 1975: Ottawa, National Research Council of Canada, p. II-49–II-65.

10. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

11. Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbor, N.W.T.: Canadian Technical 

Report of Fisheries and Aquatic Sciences, no.1218, 21 p.

12. Tokuya, K., and Amaoka, K., 1980, Studies on larval and juvenile blennies in the coastal waters of the southern Hokkaido 

(Pisces: Blennioidei): Bulletin of the Faculty of Fisheries, Hokkaido University, v. 31, no. 1, p. 16–49.

13. Houston, J., and McAllister, D.E., 1990, Status of the blackline prickleback, Acantholumpenus mackayi, in Canada: The 

Canadian Field-Naturalist, v. 104, no. 1, p. 24–28.

14. Lacho, G., 1991, Stomach content analyses of fishes from Tuktoyaktuk Harbour, N.W.T., 1981: Winnipeg, Manitoba, 

Canadian Data Report of Fisheries and Aquatic Sciences, Central and Arctic Region, Department of Fisheries and Oceans, 

no. 853, 15 p. 

15. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.
496 Alaska Arctic Marine Fish Ecology Catalog

16. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

17. Tokranov, A.M., 1981, Distribution of sculpins (Pisces, Cottidae) on the west Kamchatka shelf in summer: Zoologicheskii 

Zhurnal, v. 60, no. 2, p. 229–237.
Stout Eelblenny 497

Stout Eelblenny (Anisarchus medius) 

(Reinhardt, 1837)

Family Stichaeidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Although this species has not been reported in the 

stomach contents of other organisms, its abundance in the U.S. 

Chukchi Sea and food habits observations elsewhere (for example, 

Black Guillemots in Hudson Bay) suggests this species could be of 

modest importance in regional food webs. 

Physical Description/Attributes: Elongate, compressed, slightly eel-like body colored creamy white, yellowish or reddish, 

marked with darker spots. Dorsal fin has oblique brownish orange bars and caudal fin is finely banded. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 758) [1]. Swim bladder: Absent [2]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea and Beaufort Seas, northward to the shelf edge and a little beyond [3–5]. Elsewhere in Alaska, 

in Bering Sea and Gulf of Alaska. Worldwide, nearly circumpolar in the Arctic Ocean, also found from southern Greenland 

to the Gulf of St. Lawrence, the Barents Sea and along Siberian coasts to the Tatar Strait (northern Sea of Japan) and Sea of 

Okhotsk [4].

Relative Abundance: Common in western Chukchi Sea [3, 7] and U.S. Chukchi Sea in some years [5]. Abundance in U.S. 

Beaufort Sea is unknown. Common in eastern Bering Sea [8] but rare in Sea of Japan [9].

Stout Eelblenny (Anisarchus medius), 129 mm TL, Chukchi 

Sea, 2004. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Stout Eelblenny (Anisarchus medius) within Arctic Outer Continental Shelf Planning 

Areas [6] based on review of published literature and specimens from historical and recent collections [1, 3–5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Stout Eelblenny

Anisarchus medius

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
498 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: From nearshore to 150 m, typically less than 100 m in Chukchi and Beaufort Seas [1, 3, 5, 7, 10]. Intertidal to 

265 m in the northern Sea of Okhotsk [11]. Off Kodiak Island, Gulf of Alaska, larvae were found in 10–90 m of water during 

day and night [12].

Benthic and reproductive distribution of Stout Eelblenny (Anisarchus medius). 

Some fish to 265 m

Open ocean Shore

Depth, in meters

375

300

225

150

75

0

375

300

225

150

75

0

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Anisarchus medius

Stout Eelblenny

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Larvae

Overall data from outside U.S. Beaufort-Chukchi Seas Larvae data from outside U.S. Beaufort-Chukchi Seas

Juveniles and adults

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Depths in U.S. Chukchi Sea

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic and adhesive [13]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Off Kodiak Island, larvae were taken from March to July with densities peaking in 

April [12]. Off Kamchatka Peninsula, Russia, larvae were taken in June [14]. Habitat: Pelagic [12]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [15]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 18 cm TL [1]. 

Habitat: Benthic, coastal species [5]

Substrate—Sand and mud [3, 7, 15]. 

Physical/chemical—Temperature: -1.8–7.9 °C [3, 16], but may prefer temperatures near 0 °C [7, 15]. Salinity: 

Marine or brackish-water [3, 7, 15].

Behavior

Diel—Off Kodiak Island larvae were found in water column during day and night [12]. 

Seasonal—Unknown. 

Reproductive—Unknown. In general, Stout Eelblenny adults brood their eggs [13]. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous. 

Spawning season—Unknown. 

Fecundity—Unknown. 
Stout Eelblenny 499

Food and Feeding

Food items—Primarily benthic prey such as polychaetes, bivalves, cumaceans, and amphipods [7]. 

Trophic level—3.22 standard error 0.36 [17].

Biological Interactions

Predators—In Hudson Bay, Canada, they are eaten by Black Guillemots [18]. 

Competitors—Unknown, but likely include other small benthic fishes, such as snailfishes, flatfishes, sculpins, 

and other pricklebacks.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [17]. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Stout Eelblenny are not commercially harvested.

Potential Effects of Climate Change

Stout Eelblenny are widely distributed in Boreal and Arctic waters. It is unclear how populations may shift in 

response to climate change or respond to marine ecosystem changes.

Areas for Future Research [B]

Little is known about the ecology of this species. In particular, research needs for this species in the study area 

include: (1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic 

recruitment, (3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and 

ontogenetic movements, (6) population studies, (7) prey, and (8) predators. 
500 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [7]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [3]

Rogers, D.E., Rabin, D.J., Rogers, B.J., Garrison, K.J., and Wangerin, M.E., 1979, Seasonal composition and food web 

relationships of marine organisms in the nearshore zone of Kodiak Island—including ichthyoplankton, meroplankton 

(shellfish), zooplankton, and fish: Seattle, University of Washington College of Fisheries, Fisheries Research Institute, 

Technical Report FRI-UW-7906, 123 p. [12]

Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p. [15]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p.

3. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arctic-marine-fishes.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

8. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

9. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

10. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

11. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.
Stout Eelblenny 501

12. Rogers, D.E., Rabin, D.J., Rogers, B.J., Garrison, K.J., and Wangerin, M.E., 1979, Seasonal composition and food web 

relationships of marine organisms in the nearshore zone of Kodiak Island—including ichthyoplankton, meroplankton 

(shellfish), zooplankton, and fish: Seattle, University of Washington College of Fisheries, Fisheries Research Institute, 

Technical Report FRI-UW-7906, 123 p.

13. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

14. Grigor’yev, S.S., 1992, Larvae of three species of lumpenids (Anisarchus medius, Lumpenus fabricii, Leptoclinus 

maculatus) from Kamchatka: Journal of Ichthyology, v. 32, no. 6, p. 131–157.

15. Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p.

16. Cui, X., Grebmeier, J.M., Cooper, L.W., Lovvorn, J.R., North, C.A., Seaver, W.L., and Kolts, J.M., 2009, Spatial 

distributions of groundfish in the northern Bering Sea in relation to environmental variation: Marine Ecology Progress 

Series, v. 393, p. 147–160.

17. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

18. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.
502 Alaska Arctic Marine Fish Ecology Catalog

Bearded Warbonnet (Chirolophis snyderi) 

(Taranetz, 1938)

Family Stichaeidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and 

Beaufort Seas.

Ecological Role: Bearded Warbonnet are likely of minor ecological 

significance in the U.S. Arctic. 

Physical Description/Attributes: Elongate, compressed, slightly 

eel-like body colored pinkish orange with lilac-red bands and 

vague spots on dorsal fins. For specific diagnostic characteristics, 

see Fishes of Alaska (Mecklenburg and others, 2002, p. 752) [1]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood 

serum: Unknown.

Range: Point Barrow, Alaska [6]. Elsewhere in Alaska, Bering Sea, Aleutian Islands west to Adak Island, and to northwestern 

Gulf of Alaska [4]. Worldwide, Sea of Okhotsk, west coast of Sakhalin Island, Russia (northern Sea of Japan), and Pacific coast 

of Hokkaido, Japan [1, 2, 4]. 

Relative Abundance: Uncommon in U.S. Chukchi Sea, at least in offshore waters typically sampled, documented by single 

voucher specimens from five locations [6]. Possibly common in shallower, nearshore waters, although uncommon north and rare 

south of the Alaska Peninsula [1].

Bearded Warbonnet (Chirolophis snyderi), 122 mm TL, 

Chukchi Sea, 2010. Photograph by C.W. Mecklenburg, Point 

Stephens Research. The fish shown had been frozen and 

thawed before it was photographed, with consequent loss 

of color.

Geographic distribution of Bearded Warbonnet (Chirolophis snyderi) within Arctic OCS Planning Areas [5] 

based on review of literature and specimens from historical and recent collections [1, 3, 6].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Bearded Warbonnet

Chirolophis snyderi

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Bearded Warbonnet 503

Depth Range: 3–490 m, typically nearshore and less than 70 m [2]. The five known records from U.S. Chukchi Sea are 

from 17 to 46 m [6].

Benthic and reproductive distribution of Bearded Warbonnet (Chirolophis snyderi).

31010gBeardedWarbonnet

Some fish

to 490 m

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

500

400

300

200

100

0

Juveniles and adults

Chirolophis snyderi

Bearded Warbonnet

Benthic distribution

Open ocean No data for this species. Shore

Depth, in meters

Spawning and eggs

Depths documented in U.S. Chukchi Sea

Reproductive distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

 Larvae depths unknown

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

 Overall depth data from

outside U.S. Beaufort-Chukchi Seas

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic and adhesive [7]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Unknown. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: To 41.7 cm TL [4]. 

Habitat: Benthic, coastal species [1]. 

Substrate—Soft and rocky bottoms [1]. 

Physical/chemical—Temperature: 2.4–10.6 °C [6]. Salinity: Unknown. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. In general, warbonnet adults brood their eggs [7]. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous[7]. 

Spawning season—Unknown. 

Fecundity—Unknown.
504 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Unknown. Likely, small benthic invertebrates [2]. 

Trophic level—3.48 standard error 0.43 [8].

Biological Interactions

Predators—Unknown. 

Competitors—Unknown, but likely to be other pricklebacks, as well as such diminutive benthic species as 

sculpins, snailfishes, and some eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [8]. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Bearded Warbonnet are not commercially harvested.

Potential Effects of Climate Change

Bearded Warbonnet, as a predominantly a Boreal Pacific species already having some presence in the Arctic 

marine environment [3], would be expected to increase in abundance and expand its distribution in the U.S. 

Chukchi and Beaufort Seas, wherever suitable shallow nearshore habitat occurs.

Areas for Future Research [B]

Limited information is available regarding the biology and ecology of this species in the U.S. Arctic. Research 

needs include: (1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year 

benthic recruitment, (3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and 

ontogenetic movements, (6) population studies, (7) prey, and (8) predators.
Bearded Warbonnet 505

References Cited

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

8. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
506 Alaska Arctic Marine Fish Ecology Catalog

Fourline Snakeblenny (Eumesogrammus praecisus) 

(Krøyer, 1836)

Family Stichaeidae 

Note: Except for physical description, relative abundance, and 

geographic range data, all information is from areas outside of the 

study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown, although its abundance in the 

U.S. Chukchi and western U.S. Beaufort Seas implies potential 

ecological significance in benthic ecosystems.

Physical Description/Attributes: Elongate, compressed, slightly eel-like chocolate brown to gray body with vague, darker 

bands; 1–3 black spots, often ringed with white, near front of dorsal fin. For specific diagnostic characteristics, see Fishes 

of Alaska (Mecklenburg and others, 2002, p. 746) [1]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood serum: 

Unknown.

Range: U.S. Chukchi and Beaufort Seas [3]. Elsewhere in Alaska, in northeastern Bering Sea. Worldwide, East Siberian Sea 

through North American Arctic to west Greenland and south to Sea of Okhotsk and Gulf of St. Lawrence [1, 3].

Relative Abundance: Uncommon in U.S. Chukchi and Beaufort Seas [5]. Common off northwest and southwest Greenland [3].

Fourline Snakeblenny (Eumesogrammus praecisus), 150 mm 

TL, Bering Strait, 2007. Photograph by C.W. Mecklenburg, 

Point Stephens Research.

Geographic distribution of Fourline Snakeblenny (Eumesogrammus praecisus) within 2008–09 lease areas [4] 

based on review of published literature and specimens from historical and recent collections [1, 3, 5].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Fourline Snakeblenny

Eumesogrammus praecisus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Fourline Snakeblenny 507

Depth Range: 5–6 m [5] to 400 m, typically less than 70 m [1–3]. Taken in U.S. Chukchi Sea at 14–60 m [5, 6] and in U.S. 

Beaufort Sea at 183 m [5].

Benthic and reproductive distribution of Fourline Snakeblenny (Eumesogrammus praecisus). 

3-10-11gFourlineSnakeblenny

Some fish to 400 m

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

500

400

300

200

100

0

Juveniles and adults

Eumesogrammus praecisus

Fourline Snakeblenny

Benthic distribution

Open ocean No data for this species Shore

Depth, in meters

Spawning and eggs

Depths documented in U.S. Chukchi Sea

Reproductive distribution

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

 Larvae depths unknown

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Overall depth data from outside

U.S. Chukchi and Beaufort Seas

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic and adhesive [6]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Unknown. 

Juveniles—Age and size: Unknown. Habitat: Benthic [3]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 23 cm TL [1]. 

Habitat: Benthic [3]. Coastal algae-rock areas off Greenland [7]. 

Substrate—Sand or slightly silty bottom mixed with stones, pebbles, and gravel [1, 7]. 

Physical/chemical—Temperature: From -1.3–4 °C [5], to 2 °C or more in the Bering Sea [7]. Salinity: 

Marine [7]. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—In general, Fourline Snakeblenny adults brood their eggs [9]. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [9]. 

Spawning season—Unknown. 

Fecundity—Unknown.
508 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Amphipods in Greenland [7]. Likely, small benthic invertebrates [2]. 

Trophic level—3.5 standard error 0.50 [10].

Biological Interactions

Predators—Black Guillemots in Hudson Bay [11]. 

Competitors—Unknown, but likely other small and benthic fishes, such as sculpins, snailfishes, and flatfishes.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [10]. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Fourline Snakeblenny are not commercially harvested currently.

Potential Effects of Climate Change

Fourline Snakeblenny reproduce in Arctic and Boreal Pacific waters and in the western North Atlantic. The 

species appears to be reestablishing a circumpolar distribution in response to long-term climate change (believed 

to be circumpolar in pre-Bering Land Bridge times) [3].

Areas for Future Research [B]

Little is known about the ecology of this species. Research needs include: (1) depth and location of pelagic 

larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth ranges for 

juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population studies, 

(7) prey, and (8) predators.
Fourline Snakeblenny 509

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [7]

Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p. [6]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p. [2]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

7. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

8. Backus, B.H., 1957, The fishes of Labrador: Natural History, v. 113, no. 4, p. 273–338.

9. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

10. Froese, R., and Pauly, D., eds., 2012, FishBase—World Wide Web electronic publication—Global information system on 

fishes: FishBase, accessed July 8, 2012, at http://www.fishbase.org.

11. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.
510 Alaska Arctic Marine Fish Ecology Catalog

Daubed Shanny (Leptoclinus maculatus) 

(Fries, 1838)

Family Stichaeidae 

Note: Except for physical description, relative abundance, and 

geographic range data, all information is from areas outside of the 

study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown, but its scarcity in the U.S. Arctic implies 

it is of minimal ecological importance.

Physical Description/Attributes: Elongate, compressed, slightly eel-like body colored creamy white to yellowish brown with 

dark blotches, including four or five blackish brown saddles. Dorsal fin has dark spots or oblique bars, the caudal fin has three to 

five narrow dark bands, and other fins are unmarked and yellowish [1]. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 756) [1]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Present [3].

Range: U.S. Chukchi and Beaufort Seas [4]. Elsewhere in Alaska, from Bering Sea to Gulf of Alaska. Worldwide, from East 

Siberian and western (Russian) Chukchi Seas to Arctic Canada and southward to Sea of Okhotsk and Tatar Strait, Sea of Japan and 

Puget Sound, Washington. In the Atlantic Ocean, they also are found from Barents Sea, Svalbard Island and White Sea, Iceland 

and southward to southern Greenland, and Cape Cod, Massachusetts [4].

Relative Abundance: Common in U.S. Chukchi and Beaufort Seas [6]. Common in the eastern Bering Sea [7]. 

Daubed Shanny (Leptoclinus maculatus), 155 mm TL, 

Chukchi Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Daubed Shanny (Leptoclinus maculatus) within Arctic OCS Planning Areas [5] 

based on review of literature and specimens from historical and recent collections [1, 4, 6]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Daubed Shanny

Leptoclinus maculatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Daubed Shanny 511

Depth Range: 2–773 m, usually less than 170 m [1, 4, 6], and possibly to 607 m [8]. Larvae were taken in near-surface waters 

off Kodiak Island [9]. May spawn in shallow waters [10].

Benthic and reproductive distribution of Daubed Shanny (Leptoclinus maculatus). 

31012gDaubedShanny

Some fish

to 607 m

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

500

400

300

200

100

0

Juveniles and adults

Leptoclinus maculatus

Daubed Shanny

Benthic distribution

Open ocean No data for this species Shore

Depth in Meters

Spawning and eggs

Reproductive distribution

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Data from outside U.S. Beaufort-Chukchi Seas

Characteristic location of larvae

Habitats and Life History

Eggs—Size: 1.5 mm [3]. Time to hatching: Unknown. Habitat: Benthic and adhesive [3, 11]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: 7–8 cm [3, 12] Days to juvenile 

transformation: 2–3 years [3]. Habitat: Pelagic [3]. Off Kodiak Island, larvae were taken from April to August, 

with highest densities in April [9]. 

Juveniles—Age and size: At least 2 years and 7 cm [3]. Habitat: Young juveniles are pelagic [3] and older 

juvenile are benthic [3, 4]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 22 cm TL [13]. 

Habitat: Benthic, soft and low-relief hard sea floors [1, 3, 4, 14]. 

Substrate—Mud, sand, or stone and pebble bottoms [1]. 

Physical/chemical—Temperature: -1.6–11.5 °C [6]. Salinity: Marine and slightly brackish (as low as 26 ppt) [14, 

15].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—May come into shallow waters to spawn [10]. In general, Daubed Shanny adults brood their 

eggs [11]. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [11]. 

Spawning season—Perhaps winter in Russia [14]. December–February in North Atlantic [16]. 

Fecundity—One female contained 970 eggs [14].
512 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Small crustaceans and polychaetes [14]. Post-larval fish feed primarily on copepods [3]. 

Trophic level—3 standard error 0.00 [17].

Biological Interactions

Predators—In Canadian Arctic waters, Daubed Shanny are eaten by Black Guillemots and Thick-billed 

Murres [18]. In the North Pacific and Bering Sea, predators include Arrowtooth Flounder, Kamchatka Flounder, 

Greenland Halibut, Pacific Cod, Arctic Cod, skates, Walleye Pollock, Steller sea lions, and seals [3, 19–24]. 

Competitors—Unknown, but likely other small, benthic fishes (for example, sculpins, snailfishes, eelpouts, and 

pricklebacks).

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [17]. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Daubed Shanny are not commercially harvested.

Potential Effects of Climate Change

Daubed Shanny have an Arctic-Boreal pattern of distribution. Populations are located in the North Pacific and 

North Atlantic Oceans. A possible effect of climate warming could be an increase in abundance of the species in 

Arctic seas as it reestablishes its former circumpolar Arctic distribution of pre-Bering Land Bridge times [3].

Areas for Future Research [B]

Little is known about the species from the region. Research needs include: (1) depth and location of pelagic 

larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth ranges for 

juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population studies, 

(7) prey, and (8) predators.
Daubed Shanny 513

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [14]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]

Meyer Ottesen, C.A., Hop, H., Christiansen, J.S., and Falk-Petersen, S., 2011, Early life history of the daubed shanny (Teleostei: 

Leptoclinus maculatus) in Svalbard waters: Marine Biodiversity, v. 41, no. 3, p. 383–394. [3]

Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p. [15]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p.

3. Meyer Ottesen, C.A., Hop, H., Christiansen, J.S., and Falk-Petersen, S., 2011, Early life history of the daubed shanny 

(Teleostei: Leptoclinus maculatus) in Svalbard waters: Marine Biodiversity, v. 41, no. 3, p. 383–394.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

8. Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic marine fishes of Canada: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, no. 2674, 112 p.

9. Rogers, D.E., Rabin, D.J., Rogers, B.J., Garrison, K.J., and Wangerin, M.E., 1979, Seasonal composition and food web 

relationships of marine organisms in the nearshore zone of Kodiak Island—including ichthyoplankton, meroplankton 

(shellfish), zooplankton, and fish: Seattle, University of Washington College of Fisheries, Fisheries Research Institute, 

Technical Report FRI-UW-7906, 123 p.

10. Collette, B.B., and Klein-MacPhee, G., 2002, Bigelow and Schroeder’s fishes of the Gulf of Maine (3rd ed.): Washington, 

D.C., Smithsonian Institution Press, 882 p.

11. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

12. Dragoo, D.E., 2006, Seabird, fish, marine mammals and oceanography coordinated investigations (SMMOCI) near Bluff, 

Norton Sound, Alaska, July 2002: U.S. Fish and Wildlife Service Report AMNWR 06/03, 35 p.

13. Elliott, K.H., and Gaston, A.J., 2008, Mass-length relationships and energy content of fishes and invertebrates delivered to 

nestling thick-billed murres Uria lomvia in the Canadian Arctic, 1981–2007: Marine Ornithology, v. 36, no. 1, p. 25–34.
514 Alaska Arctic Marine Fish Ecology Catalog

14. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

15. Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p.

16. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

17. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

18. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.

19. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-158, 249 p.

20. Lang, G.M., Livingston, P.A., Pacunski, R.E., Parkhurst, J., and Yang, M.-S., 1991, Groundfish food habits and predation on 

commercially important prey species in the eastern Bering Sea from 1984 to 1986: Seattle, Washington, U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFS F/NWC-207, 240 p.

21. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.

22. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990.

23. Orlov, A.M., and Maukhametov, I.N., 2003, Feeding characteristics of Greenland halibut Reinhardtius hippoglossoides

matsuurae and Kamchatka flounder Atheresthes evermanni in the northwestern part of the Pacific Ocean: Journal of 

Ichthyology, v. 43, no. 9, p. 789–801.

24. Yang, M.-S., Dodd, K., Hibpshman, R., and Whitehouse, A., 2006, Food habits of groundfishes in the Gulf of Alaska 

in 1999 and 2001: U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical 

Memorandum, NMFS-AFSC-164, 189 p.
Slender Eelblenny 515

Slender Eelblenny (Lumpenus fabricii) 

Reinhardt, 1836

Family Stichaeidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown in U.S. Chukchi and Beaufort Seas. 

However, this species is common in these waters and is important 

prey for seabirds, marine mammals, and fishes in other parts of 

its range.

Physical Description/Attributes: Elongate, compressed, eel-shaped 

tan or cream colored body with irregular brown blotches or broken 

diagonal bars extending from the back to the lower sides. For specific diagnostic characteristics, see Fishes of Alaska 

(Mecklenburg and others, 2002, p. 759) [1]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [1, 3]. Elsewhere in Alaska, from eastern Bering Sea to Auke Bay, Alaska, in the 

eastern Gulf of Alaska [1] and Unalaska Island in the Aleutian Islands [4]. Worldwide, from the Barents Sea, eastward across 

Siberia and Arctic North America to western Greenland and south to Nova Scotia and southward in Pacific Ocean to western 

Bering Sea (off Pavla and Nataliya Bays) and northern Sea of Okhotsk; not in the eastern North Atlantic and Canadian High 

Arctic archipelago [3, 4].

Relative Abundance: Common throughout central and eastern U.S. Chukchi Sea, and U.S. Beaufort Sea [6–9]. Common in 

Canadian Beaufort Sea at least as far eastward as Tuktoyaktuk Harbor, Yukon Territory, Canada, [10] and in eastern Bering Sea 

[11]. Uncommon in Gulf of Alaska [12].

Slender Eelblenny (Lumpenus fabricii), 136 mm TL, Chukchi 

Sea, 2004. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Slender Eelblenny (Lumpenus fabricii) within Arctic Outer Continental Shelf Planning 

Areas [5] based on review of published literature and specimens from historical and recent collections [1, 3, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Slender Eelblenny

Lumpenus fabricii

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
516 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: From subtidal to 183 m, typically less than 50 m; rarely intertidal [12–14]. Primarily inhabits the inner 

continental shelf [12]. Taken in U.S. Chukchi Sea at less than 14–72 m [6, 9, 13]. Pelagic larvae were found between surface and 

48 m in western Chukchi Sea [13].

Benthic and reproductive distribution of Slender Eelblenny (Lumpenus fabricii).

Some fish to 

183 m

Open ocean Shore

Depth, in meters

250

200

150

100

50

0

250

200

150

100

50

0

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Lumpenus fabricii

Slender Eelblenny

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Larvae in U.S.

Chukchi Sea

Overall depth data from outside U.S. Beaufort-Chukchi Seas

Juveniles and adults

Depths documented in U.S. Chukchi Sea

Depth range of spawning is uncertain.

Potential location of spawning and eggs is

based on depth range of benthic individuals 

Data from outside U.S. Beaufort-Chukchi Seas

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. In James Bay, eastern Arctic Canada, eggs are reported to 

hatch in May and June [15]. Habitat: Benthic [14]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile transformation: 

Unknown. Habitat: Pelagic [13]. 

Juveniles—Age and size: Unknown. Juveniles as small as 50 mm TL have been taken in bottom trawls [6]. 

Habitat: Pelagic to benthic [1, 3, 13]. 

Adults—Age and size at first maturity: Unknown. One Kara Sea female with ripe eggs was 16.4 cm [14]. 

Maximum age: 17 years [10]. Maximum size: 36.5 cm TL [1]. Males may be larger at age (particularly in older 

fish) and may live longer [10]. Habitat: Benthic [1, 3], fish as large as 20.9 cm TL have been taken in water 

column [16]. Substrate-oriented, living among eelgrass, in algal beds, over rocky reefs [17, 18], and on relatively 

featureless seafloors of rock, sand, mud, and even anoxic mud [9]. 

Substrate—Rock, sand, mud, and mixed bottoms [9, 17, 18]. 

Physical/chemical—Temperature: -1.8–15.6 °C [9, 17]. Salinity: Marine and estuarine (to as low as 12 ppt 

[19, 20].

Behavior

Diel—Unknown. 

Seasonal—Unknown.

Reproductive—Lays its eggs among algae [14]. In general, adults of the Stichaeidae family brood their eggs 

[21]. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.
Slender Eelblenny 517

Reproduction

Mode—Oviparous. 

Spawning season—Autumn in Russian Arctic and in southeastern Beaufort Sea [19, 22] and July off west 

Greenland [14]. 

Fecundity—One female contained 490 eggs [19].

Food and Feeding

Food items—In southeastern Beaufort Sea, a diverse array of benthic and epibenthic prey including polychaetes, 

amphipods, snails, fish eggs, clam siphons, insects, bryozoans, and priapulids [23, 24]. 

Trophic level—3.28 standard error 0.37 [25].

Biological Interactions

Predators—In the eastern Canadian, Arctic Cod, ringed seals, and Black Guillemots [26–28]. Elsewhere, Great 

and Plain Sculpins, Pacific Cod, Pacific Halibut, Starry Flounder, and Walleye Pollock [29–32]. 

Competitors—Unknown, but likely many small, benthic fishes (for example, sculpins, snailfishes, flatfishes, and 

pricklebacks).

1980 1990 2000 2010

Resilience

Very low, minimum population doubling time: more than 14 years (Preliminary K or Fecundity) [25]. 

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Slender Eelblenny are not commercially harvested.

Potential Effects of Climate Change

Slender Eelblenny reproduce in Arctic and Boreal waters. Warming Arctic waters appear to be reestablishing the 

circumpolar distribution they perhaps enjoyed in pre-Bering Land Bridge times [3].

Areas for Future Research [B]

Little is known about the ecology of this species from the region. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators.
518 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.] [19]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., and Mecklenburg, T.A., 2011, Slender Eelblenny—Lumpenus fabricii Reinhardt, 1836: Arctic Ocean 

Diversity Web site, accessed July 2011, at http://www.arcodiv.org/Fish/Lumpenus_fabricii.html. [14]

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187. [9]

Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi Sea: 

Deep-Sea Research II, v. 57, no. 1–2, p. 57–70. [13] 

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

7. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

8. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

9. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

10. Hopky, G.E., and Ratynski, R.A., 1983, Relative abundance, spatial and temporal distribution, age and growth of fishes 

in Tuktoyaktuk Harbour, N.W.T., 28 June to 5 September, 1981: Canadian Manuscript Report of Fisheries and Aquatic 

Sciences 1983, 77 p.

11. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

12. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

13. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.
Slender Eelblenny 519

14. Mecklenburg, C.W., and Mecklenburg, T.A., 2011, Slender Eelblenny—Lumpenus fabricii Reinhardt, 1836: Arctic Ocean 

Diversity Web site, accessed July 2011, at http://www.arcodiv.org/Fish/Lumpenus_fabricii.html.

15. Ochman, S., and Dodson, J.J., 1982, Composition and structure of the larval and juvenile fish community of the Eastman 

River and estuary, James Bay: Naturaliste Canada, v. 109, p. 803–813.

16. Dragoo, D.E., 2006, Seabird, fish, marine mammals and oceanography coordinated investigations (SMMOCI) near Bluff, 

Norton Sound, Alaska, July 2002: U.S. Fish and Wildlife Service Report AMNWR 06/03, 35 p.

17. Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: Toronto, University of Toronto Press, 730 p.

18. Laur, D.R., and Haldorson, L.J., 1996, Coastal habitat studies—The effect of the Exxon Valdez oil spill on shallow subtidal 

fishes in Prince William Sound: American Fisheries Society Symposium 18, p. 659–670.

19. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. Department of Commerce, Springfield, Virginia.]

20. Schneider-Vieira, F., Baker, R., and Lawrence, M., 1993, The estuaries of Hudson Bay—A case study of the physical and 

biological characteristics of selected sites: Report prepared for the Hudson Bay Program by North-South Consultants Inc., 

26 p.

21. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

22. Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbor, N.W.T.: Canadian Technical 

Report of Fisheries and Aquatic Sciences, no.1218, 21 p.

23. Lacho, G., 1991, Stomach content analyses of fishes from Tuktoyaktuk Harbour, N.W.T., 1981: Winnipeg, Manitoba, 

Canadian Data Report of Fisheries and Aquatic Sciences, Central and Arctic Region, Department of Fisheries and Oceans, 

no. 853, 15 p.

24. Atkinson, E.G., and Percy, J.A., 1992, Diet comparison among demersal marine fish from the Canadian Arctic: Polar 

Biology, v. 11, no. 8, p. 567–573.

25. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

26. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.

27. McLaren, I.A., 1958, The biology of the ringed seal (Phoca hispida Schreber) in the eastern Canadian Arctic: Fisheries 

Research Board of Canada, Bulletin No. 118, 97 p.

28. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.

29. Jewett, S.C., and Feder, H.M., 1980, Autumn food of adult starry flounders, Platichthys stellatus, from the northeastern 

Bering Sea and the southeastern Chukchi Sea: Journal de Conseil International pour l’Exploration de la Mer., v. 39, no. 1, 

p. 7–14.

30. Tokranov, A.M., 1987, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok

(Cuvier) (Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114.

31. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.

32. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-158, 249 p.
520 Alaska Arctic Marine Fish Ecology Catalog

Snake Prickleback (Lumpenus sagitta) 

Wilimovsky, 1956

Family Stichaeidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown. This species is known in the U.S. 

Chukchi Sea from only one record. It has not been reported from the 

Beaufort Sea. In these Arctic waters, it is replaced by the Slender 

Eelblenny (L. fabricii). 

Physical Description/Attributes: Elongate, compressed, slightly 

eel-like body colored light green to tannish or gray dorsally and cream ventrally. Midbody has row of dark, dash-like or 

oval marks and upper body has small dark blotches or streaks. For specific diagnostic characteristics, see Fishes of Alaska 

(Mecklenburg and others, 2002, p. 760) [1]. Swim bladder: Absent [2]. Antifreeze glycoproteins in blood serum: Unknown.

Range: One (uncertain) record from U.S. Chukchi Sea north of Cape Lisburne at 69°04’N, 166°12’W [3, 4]. Elsewhere in 

Alaska, from Bering Sea to eastern Aleutian Islands and southeastern Gulf of Alaska. Worldwide, in Sea of Japan and Sea of 

Okhotsk to Commander Islands, Russia, and south to Humboldt Bay, northern California [1, 3].

Relative Abundance: If present, rare in U.S. Chukchi Sea [3]. Common in southeastern Alaska [6]. Common in Sea of 

Japan [7].

Snake Prickleback (Lumpenus sagitta), 201 mm, northern 

Bering Sea, 2011. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Snake Prickleback (Lumpenus sagittal) within Arctic Outer Continental Shelf 

Planning Areas [5] based on review of published literature and specimens from historical and recent 

collections [3, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Snake Prickleback

Lumpenus sagitta

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

Occurrence records

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Snake Prickleback 521

Depth Range: Nearshore at intertidal depths to 425 m [1, 3], typically shallower than 200 m [3]. Larvae are pelagic, in surface 

waters [8, 9].

Benthic and reproductive distribution of Snake Prickleback (Lumpenus sagittal).

31014SkPiklbk

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Some fish to 425 m

Lumpenus sagitta

Snake Prickleback

Benthic distribution

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas 500

400

300

200

100

0

Open ocean Shore

Depth, in meters

Reproductive distribution

Depth range of spawning is uncertain. 

Potential location of eggs is based on 

depth range of benthic individuals

Adults and juveniles

Larvae

Spawning and eggs

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Larvae have been taken as early as February in Gulf of 

Alaska [8]. Habitat: Likely benthic [8]. 

Larvae—Size at hatching: 5 mm [10]. Size at juvenile transformation: 48–52 mm SL [10, 11]. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [8, 9]. 

Juveniles—Age and size: Habitat: Benthic, among eelgrass, kelp, and bare bottoms [1, 3, 6]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 51 cm TL [1]. 

Habitat: Benthic, shelf species [1], among eelgrass, kelp, and over bare bottoms [3, 6, 8]. 

Substrate—Sand and mud bottoms, sometimes with small pebbles or broken shells, and cobble [11, 12]. 

Physical/chemical—Temperature: Unknown. Salinity: Nearly fresh water to marine [13]. 

Behavior

Diel—Unknown. 

Seasonal—Migrates into shallow waters in summer and early autumn off British Columbia, Canada [14]. 

Reproductive—Unknown. In general, adult prickleback brood their eggs [15]. 

Schooling—Unknown. 

Feeding—Mostly benthic feeder [16].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [15]. 

Spawning season—Unknown. 

Fecundity—Unknown. 
522 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Mainly benthic organisms. In Puget Sound and the Strait of Juan de Fuca, bivalves dominated, 

followed by tanaids and polychaetes, and then gammarids and harpacticoids [16]. Juveniles in the nearshore zone 

of Kodiak Island, Alaska, fed on polychaetes, gammarids, clam siphons, ostracods, and fish eggs [17]. Larval 

diets consist almost entirely of copepods [9] 

Trophic level—3.1 standard error 0.32 [18].

Biological Interactions

Predators—A wide variety of fishes including Pacific Halibut, Pacific Cod, Flathead Sole, Pacific Staghorn 

Sculpin, Okhotsk Snailfish, Chinook Salmon, Sand Sole, and Spotted Spiny Dogfish [19–24]; harbor and ribbon 

seals [25, 26]; and cormorants, pigeon guillemots, and common murres [27–29]. 

Competitors—Unknown, but likely to be various small benthic-feeding taxa, including other pricklebacks, 

sculpins, and flatfishes.

1980 1990 2000 2010

Resilience

Very low, minimum population doubling time: more than 14 years (Preliminary K or Fecundity) [18].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Snake Prickleback are not commercially harvested.

Potential Effects of Climate Change

The Snake Prickleback are predominantly a Boreal Pacific species. A northward shift if the species distribution 

could be expected. 

Areas for Future Research [B]

Little is known about this species from the region. Research needs include: (1) depth and location of pelagic 

larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth ranges for 

juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population studies, 

(7) prey, and (8) predators. 

References Cited

Hart, J.L., 1973, Pacific fishes of Canada: Ottawa, Fisheries Research Board of Canada, 740 p. [9]

Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 

98 p. [6]
Snake Prickleback 523

Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 281 p. 

[8]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

7. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

8. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

9. Hart, J.L., 1973, Pacific fishes of Canada: Ottawa, Fisheries Research Board of Canada, 740 p.

10. Garrison, K.J., and Miller, B.S., 1982, Review of the early life history of Puget Sound fishes: Seattle, University of 

Washington, Fisheries Research Institute, 729 p.

11. Peden, A.E., and Wilson, D.E., 1976, Distribution of intertidal and subtidal fishes of northern British Columbia and 

southeastern Alaska: Syesis, v. 9, p. 221–248.

12. Thedinga, J.F., Johnson, S.W., Neff, A.D., and Lindeberg, M.R., 2008, Fish assemblages in shallow, nearshore habitats of 

the Bering Sea: Transactions of the American Fisheries Society, v. 137, no. 4, p. 1,157–1,164.

13. Haertel, L., and Osterberg, C., 1967, Ecology of zooplankton, benthos and fishes in the Columbia River Estuary: Ecology, 

v. 48, no. 3, p. 459–472.

14. Lamb, A., and Edgell, P., 2010, Coastal fishes of the Pacific Northwest: Madeira Park, British Columbia, Harbour 

Publishing, 335 p.

15. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.
524 Alaska Arctic Marine Fish Ecology Catalog

16. Simenstad, C.A., Miller, B.S., Nyblade, C.F., Thornburgh, K., and Bledsoe, L.J., 1979, Food web relationships of northern 

Puget Sound and the Strait of Juan de Fuca—A synthesis of the available knowledge: National Oceanic and Atmospheric 

Administration/Marine Ecosystems Analysis Puget Sound Project, Prepared for Office of Environmental Engineering and 

Technology, United States Environmental Protection Agency, 334 p.

17. Rogers, D.E., Rabin, D.J., Rogers, B.J., Garrison, K.J., and Wangerin, M.E., 1979, Seasonal composition and food web 

relationships of marine organisms in the nearshore zone of Kodiak Island—including ichthyoplankton, meroplankton 

(shellfish), zooplankton, and fish: Seattle, University of Washington College of Fisheries, Fisheries Research Institute, 

Technical Report FRI-UW-7906, 123 p.

18. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

19. Miller, B.S., 1967, Stomach contents of adult starry flounder and sand sole in East Sound, Orcas Island, Washington: 

Journal of the Fisheries Research Board of Canada, v. 24, no. 12, p. 2,515–2,521.

20. Tokranov, A.M., and Vinnikov, A.B., 1991, Diet of the Pacific cod, Gadus morhua macrocephalus, and its position in the 

food chain in Kamchatkan coastal waters: Journal of Ichthyology, v. 31, no. 5, p. 84–98.

21. Armstrong, D.A., Dinnel, P.A., Orensanz, J.M., Armstrong, J.L., McDonald, T.L., Cusimano, R.F., Nemeth, R.S., Landolt, 

M.L., Skalski, J.R., Lee, R.F., and Huggett, R.J., 1995, Status of selected bottomfish and crustacean species in Prince 

William Sound following the Exxon Valdez oil spill, in Wells, P.G., Butler, J.N., and Hughes, J.S., eds., Exxon Valdez oil 

spill, fate and effects in Alaskan waters: Philadelphia, Pennsylvania, ASTM International, p. 485–547.

22. Tokranov, A.M., 2000, Feeding of Liparidae in the Pacific waters off southeastern Kamchatka and the northern Kuril 

Islands: Journal of Ichthyology, v. 40, no. 7, p. 536–542.

23. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

24. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-158, 249 p.

25. Frost, K.J., and Lowry, L.F., 1980, Feeding of ribbon seals (Phoca fasciata) in the Bering Sea in spring: Canadian Journal 

of Zoology, v. 58, no. 9, p. 1601–1607.

26. Harvey, J.T., 1987, Population dynamics, annual food consumption, movements, and dive behaviors of harbor seals, Phoca 

vitulina richardsi, in Oregon: Corvalis, Oregon State University, Ph.D. dissertation, 177 p.

27. Drent, R.H., 1965, Breeding biology of the pigeon guillemot, Cepphus columba: Ardea, v. 53, nos. 3–4, p. 99–159.

28. Robertson, I., 1974, The food of nesting double-crested and pelagic cormorants at Mandarte Island, British Columbia, with 

notes on feeding ecology: The Condor, v. 76, no. 3, p. 346–348.

29. Krasnow, L.D., and Sanger, G.A., 1982, Feeding ecology of marine birds in the nearshore waters of Kodiak Island: 

Anchorage, Alaska, Final Report to the Outer Continental Shelf Environmental Assessment Program, U.S. Fish and Wildlife 

Service, National Fisheries Research Center, Migratory Bird Section, Contract 01-5-022-2538.
Arctic Shanny 525

Arctic Shanny (Stichaeus punctatus) 

(Fabricius, 1780)

Family Stichaeidae 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown in the U.S. Chukchi and Beaufort Seas.

In the eastern Bering Sea and Hudson Bay, they are preyed upon by 

Black Guillemots and Thick-billed Murres [1, 2]. 

Physical Description/Attributes: Elongate, compressed, slightly 

eel-like body colored yellowish brown to bright scarlet with brown 

streaks and blotches. For specific diagnostic characteristics, see 

Fishes of Alaska (Mecklenburg and others, 2002, p. 747) [3]. Swim 

bladder: Absent [4]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [5]. Found in all Alaskan marine waters. Worldwide, from East Siberian Sea through 

Canadian Arctic to west Greenland, southward to Okhotsk and Japan Seas, northern British Columbia, Hudson Bay, Gulf of St. 

Lawrence, and banks off Newfoundland and Nova Scotia to Gulf of Maine [5].

Relative Abundance: Common in U.S. Chukchi and Beaufort Seas [5, 8]. In Pacific region, common at least in Cook Inlet, 

eastern and northern Bering Sea, and in northern Sea of Okhotsk, [9–11].

Arctic Shanny (Stichaeus punctatus), 105 mm, Chukchi Sea, 

2009. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Arctic Shanny (Stichaeus punctatus) within Arctic Outer Continental Shelf Planning 

Areas [6] based on review of published literature and specimens from historical and recent collections [3, 5, 7]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Shanny

Stichaeus punctatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
526 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: Shallow subtidal to 100 m, typically less than 55 m [3, 4, 11, 12]. Larvae have been taken from the surface to 110 

m [13]. In U.S. Chukchi Sea, 3 pelagic larvae were collected between the surface and 48 m [14].

Benthic and reproductive distribution of Arctic Shanny (Stichaeus punctatus). 

3-10-15g_ArcticShanny

Open ocean Shore

Depth, in meters 125

100

75

50

25

0

125

100

75

50

25

0

Adults and juveniles

Some fish 

to 100 m

Stichaeus punctatus

Arctic Shanny

Benthic distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Spawning and eggs

Larvae

Depth, in meters

Reproductive distribution

Depth range of spawning is uncertain. 

Potential location of spawning and eggs 

is based on depth range of benthic individuals

Depths documented in 

U.S. Beaufort-Chukchi Seas

Habitats and Life History

Eggs—Size: 1.7 mm [13]. Time to hatching: Unknown. Habitat: Benthic [13]. 

Larvae—Size at hatching: Unknown. Size at juvenile transformation: As small as 2.5–3.0 cm TL [15, 16]. Days 

to juvenile transformation: Unknown. Habitat: Pelagic [16]. 

Juveniles—Age and size: 2.5 to at least 11 cm [16]. Habitat: Benthic [3, 5], structure-covered, nearshore sea 

floors among eelgrass beds and understory kelps [15, 17–19]. 

Adults—Age and size at first maturity: Age unknown. A few mature at 11.0 cm SL [15]. Maximum age: 5 years 

[15]. Maximum size: 22 cm TL [3]. Habitat: Benthic [3, 5], structure-covered, nearshore sea floors and eelgrass 

beds [15, 17–19], as well as offshore [3, 11]. 

Substrate—In the northwest Atlantic Ocean, juveniles are found most often in pebble and fine cobble and adults 

in coarse cobble and boulders [15]. In U.S. Chukchi Sea, juveniles and adults were collected on shell hash, 

gravel, rock, sand, and mud [11].

Physical/chemical—Temperature: 1.4–10.5 °C [11, 20]. Salinity: Marine and, at least occasionally brackish 

waters [9, 15]. Collected from Bering Strait at salinities of 30.62–32.56 ppt [11].

Behavior

Diel—Off Newfoundland, Canada, juveniles are territorial, a behavior which appears to decrease with age [13]. 

Seasonal—Larvae have been collected in August in U.S. Chukchi Sea [14] and small larvae are present in 

Gulf of Alaska in spring [16]. Off Newfoundland, larger fish move into shallow waters in June and July and by 

November, most fish of all sizes appear to migrate out of those waters [13]. 

Reproductive—Unknown. In general, members of the Stichaeidae family brood their eggs [21]. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous. 

Spawning season—Off Newfoundland, probably in mid-winter, perhaps in February and March [15]. 

Fecundity—At least 1,624–2,475 eggs, based on two females taken off Newfoundland and off Greenland [15].
Arctic Shanny 527

Food and Feeding

Food items—Off Newfoundland, fish up to 2 years feed on copepods, amphipods, and smaller quantities of 

polychaetes, isopods, mysids, and ostracods [15]. 

Trophic level—3.08 standard error 0.23 [22].

Biological Interactions

Predators—In eastern Bering Sea and Hudson Bay, Black Guillemots and Thick-billed Murres 

[1, 2]. 

Competitors—Unknown, but likely to be a range of small, benthic fishes, including various sculpins, flatfishes, 

and eelpouts.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (K=0.24) [22].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Arctic Shanny are not commercially harvested.

Potential Effects of Climate Change

Arctic Shanny reproduce in Arctic and Boreal Pacific waters. Changes in marine habitat conditions with climate 

warming may allow the species to reestablish the circumpolar distribution it is believed to have held in the past 

[5].

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.

Remarks

Farwell and others (1976) [15] give a maximum age of 6 years. However, they labeled individuals that are less than 1 year old 

(young-of-the-year) as “1 year olds.” Thus, the three fish designated as 6 years old are actually age-5 fish. 
528 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Brown, J., and Green, J.M., 1976, Territoriality, habitat selection, and prior residency in underyearling Stichaeus punctatus

(Pisces: Stichaeidae): Canadian Journal of Zoology, v. 54, no. 11, p. 1,904–1,907. [13]

Farwell, M.K., Green, J.M., and Pepper, V.A., 1976, Distribution and known life history of Stichaeus punctatus in the northwest 

Atlantic: Copeia, v. 1976, no. 3, p. 598–602. [15]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [5]

Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi Sea: 

Deep-Sea Research II, v. 57, no. 1–2, p. 57–70. [14]

Bibliography

1. Hunt, G.L., Burgeson, B., and Sanger, G.A., 1981, Feeding ecology of seabirds of the Eastern Bering Sea, in Hood, D.W., 

and Calder, J.A., eds., The Eastern Bering Sea Shelf—Oceanography and Resources: National Oceanic and Atmospheric 

Administration, p. 629–641.

2. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Mecklenburg, C.W., and Sheiko, B.A., 2004, Family Stichaeidae Gill 1864—Pricklebacks: California Academy of Sciences 

Annotated Checklists of Fishes no. 35, 36 p.

5. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

8. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

9. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

10. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

11. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.
Arctic Shanny 529

12. Collette, B.B., and Klein-MacPhee, G., 2002, Bigelow and Schroeder’s fishes of the Gulf of Maine (3rd ed.): Washington, 

D.C., Smithsonian Institution Press, 882 p.

13. Brown, J., and Green, J.M., 1976, Territoriality, habitat selection, and prior residency in underyearling Stichaeus punctatus

(Pisces: Stichaeidae): Canadian Journal of Zoology, v. 54, no. 11, p. 1,904–1,907.

14. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

15. Farwell, M.K., Green, J.M., and Pepper, V.A., 1976, Distribution and known life history of Stichaeus punctatus in the 

northwest Atlantic: Copeia, v. 1976, no. 3, p. 598–602.

16. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

17. Laur, D.R., and Haldorson, L.J., 1996, Coastal habitat studies—The effect of the Exxon Valdez oil spill on shallow subtidal 

fishes in Prince William Sound: American Fisheries Society Symposium 18, p. 659–670.

18. Dean, T.A., Haldorson, L.J., Laur, D.R., Jewett, S.C., and Blanchard, A., 2000, The distribution of nearshore fishes in 

kelp and eelgrass communities in Prince William Sound, Alaska—Associations with vegetation and physical habitat 

characteristics: Environmental Biology of Fishes, v. 57, no. 3, p. 271–287.

19. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

20. Blackburn, J.E., and Jackson, P.B., 1982, Seasonal composition and abundance of juvenile and adult marine finfish and crab 

species in the nearshore zone of Kodiak Island’s eastside during April 1978 through March 1979: Outer Continental Shelf 

Environmental Assessment Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 377–570.

21. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

22. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
530 Alaska Arctic Marine Fish Ecology Catalog

Banded Gunnel (Pholis fasciata) 

(Bloch & Schneider, 1801)

Family Pholidae 

Note: Except for geographic range data, all information is from 

areas outside of the study area. 

Colloquial Name: None within U.S. Chukchi and Beaufort Seas.

Ecological Role: Its apparent scarceness in the U.S. Chukchi and 

Beaufort Seas implies this species is of little ecological importance.

Physical Description/Attributes: Body elongate and strongly 

compressed. Bright reddish orange to greenish yellow with sinuous reddish black bands reaching the ventral surface, white 

blotches along the back and dorsal fin containing black spots [1]. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 777) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea (and presumably U.S. Beaufort Sea [4]. Elsewhere in Alaska, Bering Sea to western Gulf of Alaska at 

Kodiak Island. Worldwide, Canadian Beaufort Sea eastward to west Greenland, southward to Bay of Fundy and banks off Nova 

Scotia, and southward to Seas of Okhotsk and Japan [2].

Relative Abundance: Occasional in U.S. Chukchi Sea. As yet, no records from the U.S. Beaufort Sea [2]. Common off 

Greenland [5].

Banded Gunnel (Pholis fasciata), 159 mm, eastern Bering 

Strait, 2004. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

Geographic distribution of Banded Gunnel (Pholis fasciata) within Arctic Outer Continental Shelf Planning 

Areas [3] based on review of published literature and specimens from historical and recent collections [1, 2, 4]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Banded Gunnel

Pholis fasciata

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Banded Gunnel 531

Depth Range: Shallow subtidal to 110 m [1, 5]. Less than 50 m in the Arctic Ocean, typically shallower than 20 m [1]. 

Documented in Bering Strait at 50 m [6, 7] Pelagic larvae were taken from 0 to 7 m in James Bay, Canada [8].

Benthic and reproductive distribution of Banded Gunnel (Pholis fasciata).

3-10-16g_BandedGunnel

Open ocean Shore

Depth, in meters 150

120

90

60

30

0

150

120

90

60

30

0

Adults and juveniles

Some fish 

to 110 m

Some fish in 

Arctic to 50 m

Pholis fasciata

Banded Gunnel

Benthic distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Depth, in meters

Reproductive distribution

Depth range of spawning is uncertain. 

Potential location of eggs is based on 

depth range of benthic individuals

Larvae

Spawning and eggs

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Benthic, adhesive [9]. 

Larvae—Size at hatching: Unknown. Yolk-sac larvae taken in James Bay were 11–14 mm TL [8]. Size at 

juvenile transformation: Unknown. Days to juvenile transformation: Unknown. Habitat: Pelagic [9]. 

Juveniles—Age and size: Unknown. Habitat: Benthic, over rocky substrate and among clumps of algae [1]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 30 cm TL [1]. 

Habitat: Benthic, over rocky substrate and among clumps of algae [1]. 

Substrate—Sand, broken shell, gravel, and rock [1, 7]. 

Physical/chemical—Temperature: -1.0–10.5 °C [7, 8]. Salinity: Marine. Pelagic larvae were taken between 4.0 

and 17.0 ppt [8]. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. Gunnels in general tend to guard their eggs [9]. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [9]. 

Spawning season—May and early June in James Bay, Canada [8]. 

Fecundity—Unknown. 
532 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Unknown. Small crustaceans and mollusks for gunnels in general [10]. 

Trophic level—3.27 standard error 0.39 [11].

Biological Interactions

Predators—Black Guillemots in Hudson Bay in summer [12]. Sculpins, cods, other bottom fishes, and seabirds 

[13]. 

Competitors—Likely other microcarnivores, including sculpins, gunnels, and flatfishes.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time: 1.4–4.4 years (Preliminary K or Fecundity) [11].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Banded Gunnel are not commercially harvested.

Potential Effects of Climate Change

Unknown.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs include: (1) depth and location 

of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, (3) preferred depth 

ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic movements, (6) population 

studies, (7) prey, and (8) predators. 

References Cited

Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p. [9]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]
Banded Gunnel 533

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [2]

Ochman, S., and Dodson, J.J., 1982, Composition and structure of the larval and juvenile fish community of the Eastman River 

and estuary, James Bay: Naturaliste Canada, v. 109, p. 803–813. [8]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Møller, P.R., Nielsen, J.G., Knudsen, S.W., Poulsen, J.Y., Sünksen, K., and Jørgensen, O.A., 2010, A checklist of the fish 

fauna of Greenland waters: Zootaxa, v. 2,378, p. 1–84.

6. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

7. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

8. Ochman, S., and Dodson, J.J., 1982, Composition and structure of the larval and juvenile fish community of the Eastman 

River and estuary, James Bay: Naturaliste Canada, v. 109, p. 803–813.

9. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

10. Mecklenburg, C.W., 2003, Family Pholidae Gill 1893—Gunnels: California Academy of Sciences Annotated Checklists of 

Fishes no. 9, 11 p.

11. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

12. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Canadian Government 

Publishing Centre, Canada Communication Group, Canadian Wildlife Service report series, no. 46, 63 p.

13. Mecklenburg, C.W., and Mecklenburg, T.A., 2012, Banded Gunnel—Pholis fasciata (Bloch and Schneider, 1801): Arctic 

Ocean Diversity Web site, accessed February 2012, at http://www.arcodiv.org/Fish/Pholis_fasciata.html.
534 Alaska Arctic Marine Fish Ecology Catalog

Northern Wolffish to Greenland Halibut

Northern Wolffish (Anarhichas denticulatus) 

Krøyer, 1845

Family Anarhichadidae 

Note: Except for geographic range data, all information is 

from areas outside of the study area. 

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Rare in both seas, which suggests a limited function 

role in marine ecosystem dynamics. 

Physical Description/Attributes: Moderately thick, elongate gray to 

dark brown body, with bright violet shades or with brown tones. Body and dorsal fins covered with dark spots. For specific 

diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 784) [1]. Swim bladder: Absent [2]. 

Antifreeze glycoproteins in blood serum: Unknown.

Range: Known in the U.S. Chukchi Sea from one carcass found on the ice near Barrow and in the U.S. Beaufort Sea from a few 

carcasses on the beach at Kaktovik. Worldwide, Barents and Norwegian Seas to Spitsbergen, Iceland, Faroe Islands, Greenland, 

south along east coast of North America to southern New England, and Canadian Arctic [3].

Relative Abundance: Rare in U.S. Chukchi and Beaufort Seas [3]. Elsewhere in world, uncommon in Gulf of St. Lawrence [6]. 

Previously common but declining in northwestern Atlantic Ocean [7].

Northern Wolffish (Anarhichas denticulatus), Barents 

Sea, 2010. Photograph by Arve Lynghammar, University 

of Tromsø, Norway.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Northern Wolffish

Anarhichas denticulatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Northern Wolffish (Anarhichas denticulatus), within Arctic Outer Continental Shelf 

Planning Areas [4] based on review of published literature and specimens from historical and recent collections 

[3, 5]. 
Northern Wolffish 535

Depth Range: Surface to 1,504 m and possibly to 1,700 m [5]; mainly 150–900 m [7]. Spawns below 400 m [8].

tac14-5222_3-11-1g_NorthernWolffish

Open ocean

Depth, in meters

Adults make daily 

vertical excursions

Anarhichas denticulatus

Northern Wolffish

Benthic distribution

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Some fish to 1,504 m

Shore Open ocean Shore Depth, in meters

Spawning and eggs

Reproductive distribution

Larvae pelagic but specific 

depth unknown

1,500

1,200

900

600

300

0

1,500

1,200

900

600

300

0

Depth range of eggs is uncertain. 

Potential location is based on depth 

range of spawning individuals

Data from outside 

U.S. Beaufort-Chukchi Seas

 Data from outside 

U.S. Beaufort-Chukchi Seas

Juveniles and adults

Benthic and reproductive distribution of Northern Wolffish (Anarhichas denticulatus).

Habitats and Life History

Eggs—Size: 6–8 mm [6, 8]. Time to hatching: Unknown. Habitat: Benthic, on rocky bottoms [6, 7]. 

Larvae—Size at hatching: 25–26 mm [6]. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [6]. 

Juveniles—Age and size: Unknown. Habitat: Benthopelagic to pelagic [2, 6, 8]. 

Adults—Age and size at first maturity: A few males mature starting at 5 years [7]. Females mature at 6–8 years 

(about 80 cm) [8]. 

Maximum age: 16 years [8]. Maximum size: To at least 138 cm TL [5] and 32 kg [8]. Habitat: Benthopelagic to 

pelagic [2, 6, 8] over soft bottoms near boulders [7]. 

Substrate—Mud, sand, pebbles, small rock, and hard bottoms [6, 8]. Rocky bottoms for spawning [6]. 

Physical/chemical—Temperature: -1.4–7.0 °C [8], most common from 1 to 5 °C [6]. Spawns at 1.6–4 °C [9]. 

Salinity: Prefers high salinity [8].

Behavior

Diel—Has extensive daily vertical migrations [8], but limited horizontal migrations [6]. Somewhat territorial 

[6, 10]. 

Seasonal—Migrates between spawning, feeding, and wintering grounds [8]. 

Reproductive—Builds nests [6]. 

Schooling—Solitary [6, 8]. Feeding: In Canadian Atlantic, feeds mid-water on both bathypelagic and 

mesopelagic prey [6]. Stops or reduces foraging in February–March when teeth are shed [6, 8].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes; oviparous [11]. 

Spawning season—Between April and October on continental slope of Barents and Norwegian Seas. Peak is 

during summer months [8]. 

Fecundity—23,380–42,500 eggs [6, 8].
536 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Pelagic larvae consume planktonic invertebrates, fish eggs and fish larvae [7]. Juveniles consume 

planktonic crustaceans (copepods, hyperiids, and euphausiids) [12]. Young adults consume fish, hyperiids, 

pteropods, sea urchins, ctenophores, and jellyfish [2, 8, 12, 13]. Older adults consume predominantly fish [13]. 

Trophic level—3.75 (standard error 0.46) [14].

Biological Interactions

Predators—In Canada, ringed seals, Golden Redfish, cod, and Greenland Shark [7]. 

Competitors—For juveniles and young adults, likely various gadids, poachers, eelpouts, and flatfishes. For older 

adults, likely larger gadids and flatfishes.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (K=0.08–0.10) [14].

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Northern Wolffish are not commercially harvested.

Potential Effects of Climate Change

Unknown. 

Areas for Future Research [B]

Little is known about the ecology and life history of this species in the study area. In particular, research needs in 

the study area include: (1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-theyear benthic recruitment; (3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal 

and ontogenetic movements; (6) population studies; (7) prey; and (8) predators.

References Cited

Kulka, D., Hood, C., and Huntington, J., 2007, Recovery strategy for northern wolffish (Anarhichas denticulatus) and spotted 

wolffish (Anarhichas minor), and management plan for Atlantic Wolffish (Anarhichas lupus) in Canada: St. John’s, 

Fisheries and Oceans Canada, Newfoundland and Labrador Region, 103 p. [6]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]
Northern Wolffish 537

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

O’Dea, N.R., and Haedrich, R.L., 2001, COSEWIC status report on the northern wolffish Anarhichas denticulatus in Canada, in

Committee on the Status of Endangered Wildlife in Canada, Status report on the northern wolffish Anarhichas denticulatus

in Canada: Ottawa, Committee on the Status of Endangered Wildlife in Canada, p. 1–21. [7]

Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, A., 

Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502‐8828, 274 p. [8]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., 2003, Family Anarhichadidae Bonaparte 1846—Wolffishes: California Academy of Sciences 

Annotated Checklists of Fishes, no. 10, 6 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Kulka, D., Hood, C., and Huntington, J., 2007, Recovery strategy for northern wolffish (Anarhichas denticulatus) and 

spotted wolffish (Anarhichas minor), and management plan for Atlantic Wolffish (Anarhichas lupus) in Canada: St. John’s, 

Fisheries and Oceans Canada, Newfoundland and Labrador Region, 103 p.

7. O’Dea, N.R., and Haedrich, R.L., 2001, COSEWIC status report on the northern wolffish Anarhichas denticulatus in 

Canada, in Committee on the Status of Endangered Wildlife in Canada, Status report on the northern wolffish Anarhichas 

denticulatus in Canada: Ottawa, Committee on the Status of Endangered Wildlife in Canada, p. 1–21.

8. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502‐8828, 274 p.

9. Rose, G.A., 2005, On distributional responses of North Atlantic fish to climate change: ICES Journal of Marine Science, 

v. 62, no. 7, p. 1,360–1,374.

10. Godø, O.R., Huse, I., and Michalsen, K., 1997, Bait defence behaviour of wolffish and its impact on long-line catch rates: 

ICES Journal of Marine Science, v. 54, no. 2, p. 273–275.

11. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

12. Grigorev, S.S., 1983, New catches of young northern wolfish, Anarhichas denticulatus (Anarhichadidae): Journal of 

Ichthyology, v. 23, no. 2, p. 157–160.

13. González, C., Paz, X., Román, E., and Alvárez, M., 2006, Feeding habits of wolffishes (Anarhichas denticulatus, A. lupus, 

A. minor) in the North Atlantic: Northwest Atlantic Fisheries Organization, NAFO SCR Doc. 06/52, 16 p.

14. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
538 Alaska Arctic Marine Fish Ecology Catalog

Bering Wolffish (Anarhichas orientalis) 

Pallas, 1814

Family Anarhichadidae

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: This is an uncommon species in the U.S. Chukchi 

and Beaufort Seas. Its role in benthic ecosystem functioning 

probably is of modest significance regarding competition with 

marine invertebrate competitors.

Physical Description/Attributes: Elongate laterally compressed body, colored brown, reddish-brown, or black with mottling 

and blotches. Heads are blunt with large forward-projecting canine teeth. Juveniles have dark stripes. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 783) [1]. Swim bladder: Absent [2]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [3]. Elsewhere in Alaska, this fish has been found southward to Prince William Sound, 

Gulf of Alaska. Worldwide, Bering Wolffish are found in the Sea of Okhotsk, Peter the Great Bay, Sea of Japan, and eastward in 

Canadian Beaufort Sea to Bathurst Inlet, Nunavut [3]. 

Relative Abundance: Occasional in U.S. Chukchi and Beaufort Seas [1, 6, 7]. Widespread and common in northern Sea of 

Okhotsk [8] and in eastern Bering Sea [9]. 

Bering Wolffish (Anarhichas orientalis), 193 mm, Chukchi 

Sea, 2007. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Bering Wolffish

Anarhichas orientalis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Bering Wolffish (Anarhichas orientalis), within Arctic Outer Continental Shelf 

Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [1, 3, 5].
Bering Wolffish 539

tac14-5222_3-11-2g_BeringWolffish

Open ocean

Depth, in meters

125

100

75

50

25

0

125

100

75

50

25

0

Anarhichas orientalis

Bering Wolffish

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning and eggs

Reproductive distribution

Depth range of spawning is uncertain. 

Potential location of spawning and eggs 

is based on depth range of benthic individuals

Larvae and younger juveniles are pelagic 

but specific depths unknown

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Shore

Older juveniles and adults

Overall benthic depth range.

Specific depth range of either 

juveniles or adults is unknown Some fish

 to 100 m

Benthic and reproductive distribution of Bering Wolffish (Anarhichas orientalis)

Habitats and Life History

Eggs—Size: 4–8 mm [10, 11]. Time to hatching: Unknown. Habitat: Benthic [10]. 

Larvae—Size at hatching: About 17–18 mm SL [10]. Size at juvenile transformation: About 40 mm SL [10]. 

Days to juvenile transformation: Unknown. Habitat: Pelagic [12]. 

Juveniles—Age and size: About 40–150 mm SL [10, 13]. Habitat: Benthopelagic [3]; juveniles to lengths of at 

least 50 mm TL, are sometimes found in surface waters [7, 14]. 

Adults—Age and size at first maturity: Age, unknown. Size, about 15–17 cm TL [11]. Maximum age: At least 

17 years [11]. Maximum size: 124 cm TL and 19.5 kg and 15 kg [1]. Habitat: Benthopelagic [3]. Nearshore, 

intertidal area, among rocks and kelp [8, 12]. 

Substrate—Gravel and sand, around rocks [1, 13]. 

Physical/chemical—Temperature: -0.2–11.9 °C [5, 13]. Salinity: Marine [12, 13]. 

Behavior

Diel—Observed being nocturnally active near Saint Michael Island, northeastern Bering Sea [15]. 

Seasonal—Benthic; individuals are believed to migrate nearshore after ice melts [12]. 

Reproductive—One or both parents guard eggs [10]. 

Schooling—Unlikely. Generally, wolffish are solitary or occur in small groups [1]. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [10]. Fertilization is external [16]. 

Spawning season—October and November in the Sea of Japan [17]. One female with well-developed eggs was 

taken in late May off Kamchatka [12]. 

Fecundity—Unknown.

Depth Range: From shallow waters near shore to 100 m, typically in near-shore waters [3]. 
540 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Little is known. Hermit crabs, crabs, eggs, and snails based on a few fish from Russia [11, 

12]. Juveniles (to 21 cm TL) in Russia fed on hyperiid amphipods, fishes (young Irish lords), pteropods, and 

euphausiids [18]. 

Trophic level—3.798 (standard error 0.60) [16].

Biological Interactions

Predators—Pacific Cod and northern fur seals in the eastern Bering Sea [19, 20]. 

Competitors—Unknown, but likely to include pricklebacks, eelpouts, sculpins, and flatfishes.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (assuming t

m>5) [16]. 

Traditional and Cultural Importance

None reported. Historically, on Saint Michael Island in Norton Sound, wolffish were a popular food with the 

Iñuits who caught them with hooks baited with grass. The tanned skin of this species was inserted between the 

seams of boots and other waterproof clothing as the skin was believed to swell when moistened [15].

Commercial Fisheries

Currently, Bering Wolffish are not commercially harvested.

Potential Effects of Climate Change

Because Bering Wolffish are a predominantly boreal Pacific species that is typically found in nearshore, ice-free 

habitats, abundance would be expected to increase in both the U.S. Chukchi and Beaufort Seas wherever suitable 

habitat and diet occur.

Areas for Future Research [B]

Little is known about the ecology and life history of this species. Research needs in the study area include: 

(1) depth and location of pelagic larvae, (2) depth, location, and timing of young-of-the-year benthic recruitment, 

(3) preferred depth ranges for juveniles and adults, (4) spawning season, (5) seasonal and ontogenetic 

movements, (6) population studies, (7) prey, and (8) predators.
Bering Wolffish 541

References Cited

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R—Keys to the fauna in the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from the U.S. Department of Commerce, Springfield, Virginia.] [12]

Fruge, D.J., and Wiswar, D.W., 1991, First records of the Bering wolffish, Anarhichas orientalis, for the Alaskan Beaufort Sea: 

Canadian Field-Naturalist, v. 105, no. 1, p. 107–109. [13]

Houston, J., and McAllister, D.E., 1990, Status of the Bering wolfish, Anarhichas orientalis, in Canada: The Canadian FieldNaturalist, v. 104, no. 1, p. 20–23. [11]

Matarese, A.C., Kendall, J., Arthur W., Blood, D.M., and Vinter, B.M., 1989, Laboratory guide to early life history stages of 

northeast Pacific fishes: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National Marine 

Fisheries Service Technical Report NMFS 80, 652 p. [10]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., 2003, Family Anarhichadidae Bonaparte 1846—Wolffishes: California Academy of Sciences 

Annotated Checklists of Fishes, no. 10, 6 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2 p. 195–209.

7. Barton, L.H., 1978, Finfish resource surveys in Norton Sound and Kotzebue Sound: Alaska Department of Fish and Game, 

Commercial Fisheries Division, p. 75–313.

8. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

9. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

10. Matarese, A.C., Kendall, J., Arthur W., Blood, D.M., and Vinter, B.M., 1989, Laboratory guide to early life history stages 

of northeast Pacific fishes: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National 

Marine Fisheries Service Technical Report NMFS 80, 652 p.
542 Alaska Arctic Marine Fish Ecology Catalog

11. Houston, J., and McAllister, D.E., 1990, Status of the Bering wolfish, Anarhichas orientalis, in Canada: The Canadian 

Field-Naturalist, v. 104, no. 1, p. 20–23.

12. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R—Keys to the fauna in the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from the U.S. Department of Commerce, Springfield, Virginia.]

13. Fruge, D.J., and Wiswar, D.W., 1991, First records of the Bering wolffish, Anarhichas orientalis, for the Alaskan Beaufort 

Sea: Canadian Field-Naturalist, v. 105, no. 1, p. 107–109.

14. Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, Fisheries Research Institute, FRI-UW-7719.

15. Turner, L.M., 1886, Contributions to the natural history of Alaska—Arctic series of publications, no. 2: Washington, D.C., 

U.S. Government Printing Office, 226 p.

16. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

17. Kolpakov, N.V., 2005, Diversity and seasonal dynamics of ichthyocenosis of the Circumlittoral of Russkaya Bight 

(Northern Primor’s): Journal of Ichthyology, v. 45, no. 9, p. 744–753.

18. Chuchukalo, V.I., Efimkin, A.Y., and Lapko, V.V., 1995, Feeding of some planktivorous fishes in the Sea of Okhotsk during 

the summer: Russian Journal of Marine Biology, v. 21, p. 112–115.

19. Harry, G.Y., and Hartley, J.R., 1981, Northern fur seals in the Bering Sea, in Hood, D.W., and Calder, J.A., eds., The Eastern 

Bering Sea shelf—Oceanography and resources: Seattle, University of Washington Press, p. 847–867.

20. Yang, M.-S., Dodd, K., Hibpshman, R., and Whitehouse, A., 2006, Food habits of groundfishes in the Gulf of Alaska 

in 1999 and 2001: U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical 

Memorandum, NMFS-AFSC-164, 189 p.
Prowfish 543

Prowfish (Zaprora silenus) 

Jordan, 1896

Family Zaproridae 

Note: Except for geographic range data, all information is from 

areas outside of the U.S. Chukchi and Beaufort Seas. 

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Unknown. Only one specimen is known from the 

U.S. Chukchi Sea and the species is absent from the U.S. Beaufort 

Sea. The ecological role is probably insignificant.

Physical Description/Attributes: Blunt snout and elongate, 

compressed, somewhat flaccid body. Head pores large, outlined 

in white, yellow, or pale blue. Adults are grayish blue to 

green and may have darker spots and yellow blotches. Juveniles are orange-brown and have spots and blotches. For specific 

diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 786) [1]. Swim bladder: Absent [2]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: Southern U.S. Chukchi Sea [3]. Elsewhere in Alaska, south from Bering Sea through Aleutian Islands to southern Gulf of 

Alaska. Worldwide, Pacific Ocean from Hokkaido, Japan, and Sea of Okhotsk south to San Miguel Island, southern California [1].

Relative Abundance: Rare in U.S. Chukchi Sea, where presence is documented by one juvenile found west of Kivalina at 

67°32’N, 165°54’W in 2007 [3]. Common at least from Kuril Islands and southeastern Kamchatka to eastern Bering Sea and 

southward to about Vancouver Island [6–8].

Prowfish (Zaprora silenus), juvenile, 139 mm, northeastern 

Bering Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Prowfish

Zaprora silenus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Occurrence records

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit) UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Prowfish (Zaprora silenus), within Arctic Outer Continental Shelf Planning Areas [4] 

based on review of published literature and specimens from historical and recent collections [3, 5].
544 Alaska Arctic Marine Fish Ecology Catalog

Depth Range: From 10 to 801 m, typically between 100 and 250 m, along deeper continental shelf and shallower continental 

slope [6, 7, 9]. Larvae and juveniles are pelagic in surface waters [6]. One juvenile taken pelagically in U.S. Chukchi Sea over 

bottom depth of 43 m [3].

tac14-52223-11-3gProwfish

Open ocean Shore

Depth, in meters

1,000

800

600

400

200

0

1,000

800

600

400

200

0

 Adults and

older juveniles

Some fish to 801 m

Zaprora silenus

Prowfish

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Spawning Larvae and young juveniles pelagic.

Specific depth range unknown

Depths of spawning is unknown. 

Location is based on depths 

of adult fish. Location of eggs is

unknown

Data from outside U.S. Beaufort-Chukchi Seas

One larvae documented in U.S. 

Chukchi Sea from 0-48 m

Benthic and reproductive distribution of Prowfish (Zaprora silenus). 

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Unknown. 

Larvae—Size at hatching: 4 mm SL [10]. Size at juvenile transformation: 3.0 cm TL [11, 12]. Days to juvenile 

transformation: Unknown. Habitat: Pelagic, common on outer shelf edges [6]. 

Juveniles—Age and size: Age unknown, 3.0–57 cm TL [6, 10]. Pelagic to at least 18 cm [6, 12]. Habitat: Young 

juveniles are pelagic. Older juveniles are benthopelagic [3, 6]. 

Adult—Age and size at first maturity: At about 5 years, 50 percent of females are mature at about 57 cm TL 

[6]. Females are slightly heavier at length than males. Males and females have similar growth rates and, in the 

northeast Pacific, females and males reach the same length [6]. Fish in northeast Pacific are larger at age than 

those in western Pacific and females are slightly heavier at length than males [6]. Off southeastern Kamchatka 

and northern Kuril Islands, females appear to grow larger than males [12]. Maximum age: At least 20 years old 

[6]. Maximum size: 1 m TL or more [1] and 9.3 kg [13]. Habitat: Benthopelagic [3, 6], in caves and other rocky 

habitats [14, 15]. 

Substrate—Cobble, mud, sand, and gravel [14–16]. 

Physical/chemical—Temperature: 0.2–8.5 °C off Russia [12, 13]. Salinity: Marine to slightly brackish water [17].

Behavior

Diel—Pelagic larvae and juveniles may be obligate commensals of medusae, because they often are found in 

association with large jellyfish. Juveniles usually swim near tops and sides of bells, but will dive within or behind 

the tentacles when frightened. As many as nine juveniles have been seen associating with one medusa [18]. 

Seasonal—Off northern Kuril Islands and southeastern Kamchatka, prowfish may migrate from deep shelf to 

shallow slope waters in the late autumn and early winter [12]. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Mostly water column feeders [6, 19, 20].

Populations or Stocks

There have been no studies.
Prowfish 545

Reproduction

Mode— Unknown. 

Spawning season—Spawning may occur primarily in the winter and spring, as most newly hatched larvae are 

captured during this period [10]. 

Fecundity—Unknown. 

Food and Feeding

Food items—In eastern Bering Sea and northern Pacific, dominant prey are gelatinous organisms such as 

medusae, pelagic tunicates, and comb jellies [2, 12]. Other important prey includes copepods, amphipods, 

euphausiids, larvaceans, fish larvae, and polychaetes [6, 19, 20]. 

Trophic level—3.6 (standard error 0.5) [21].

Biological Interactions

Predators—Black Rockfish, Chinook and Coho salmon, Pacific Cod, other prowfish, Whiteblotched Skates, and 

Tufted Puffins [22–27].

Competitors—In U.S. Chukchi Sea, likely competitors include midwater planktivores such as Arctic and Saffron 

Cods.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (K= 0.18, t

m= 5.1;) [21].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Prowfish are not commercially harvested.

Potential Effects of Climate Change

The Prowfish is a boreal Pacific species [3], which could be expected to increase in abundance in the Chukchi 

Sea and perhaps expand its range into the Beaufort Sea.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment; 

(3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic 

movements; (6) population studies; (7) prey; and (8) predators. 
546 Alaska Arctic Marine Fish Ecology Catalog

References Cited

Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p. [10]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Smith, K.R., Somerton, D.A., Yang, M.-S., and Nichol, D.G., 2004, Distribution and biology of prowfish (Zaprora silenus) in 

the northeast Pacific: Fishery Bulletin, v. 102, no. 1, p. 168–178. [6]

Tokranov, A.M., 1999, Some features of biology of the prowfish Zaprora silenus (Zaproridae) in the Pacific waters of the 

northern Kuril Islands and southeastern Kamchatka: Journal of Ichthyology, v. 39, no. 6, p. 475–478. [12]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., 2003, Family Zaproridae Jordan & Evermann 1898—Prowfishes: California Academy of Sciences 

Annotated Checklists of Fishes No. 13.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Smith, K.R., Somerton, D.A., Yang, M.-S., and Nichol, D.G., 2004, Distribution and biology of prowfish (Zaprora silenus) 

in the northeast Pacific: Fishery Bulletin, v. 102, no. 1, p. 168–178.

7. Orlov, A.M., 2005, Bottom trawl-caught fishes and some features of their vertical distribution in the Pacific waters off the 

north Kuril Islands and south-east Kamchatka, 1993–1999: Aqua, Journal of Ichthyology and Aquatic Biology of Fishes, 

v. 9, no. 4, p. 139–160.

8. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

9. Zenger, H.H., Jr., 2004, Data report—2002 Aleutian Islands bottom trawl survey: U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS-AFSC-143, 247 p.

10. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

11. Matarese, A.C., Kendall, J., Arthur W., Blood, D.M., and Vinter, B.M., 1989, Laboratory guide to early life history stages 

of northeast Pacific fishes: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National 

Marine Fisheries Service Technical Report NMFS 80, 652 p.

12. Tokranov, A.M., 1999, Some features of biology of the prowfish Zaprora silenus (Zaproridae) in the Pacific waters of the 

northern Kuril Islands and southeastern Kamchatka: Journal of Ichthyology, v. 39, no. 6, p. 475–478.
Prowfish 547

13. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

14. O’Connell, M., Dillon, M.C., Wright, J.M., Bentzen, P., Merkouris, S.E., and Seeb, J.E., 1998, Genetic structuring among 

Alaskan Pacific herring populations identified using microsatellite variation: Journal of Fish Biology, v. 53, no. 1, 

p. 150–163.

15. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, no. 1–3, p. 15–28.

16. Johnson, S.W., Thedinga, J.F., and Lindeberg, M.R., 2012, Nearshore fish atlas of Alaska: National Oceanic and 

Atmospheric Administration Fisheries, accessed February 2012 at http://www.fakr.noaa.gov/habitat/fishatlas/. 

17. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

18. Brodeur, R.D., 1998, In situ observations of the association between juvenile fishes and scyphomedusae in the Bering Sea: 

Marine Ecology Progress Series, v. 163, p. 11–20.

19. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

20. Yang, M.-S., 2003, Food habits of the important groundfishes in the Aleutian Islands in 1994 and 1997: National Marine 

Fisheries Service, AFSC Processed Report 2003-07, 233 p.

21. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

22. Rosenthal, R.J., Haldorson, L.J., Field, L.J., Moran-O’Connell, V., LaRiviere, M.G., Underwood, J., and others, 1982, 

Inshore and shallow offshore bottomfish resources in the southeastern Gulf of Alaska (1981–82): Alaska Coastal Research 

and University of Alaska, Juneau, 166 p.

23. Baird, P.A., and Gould, P., 1983, The breeding biology and feeding ecology of marine birds in the Gulf of Alaska: U.S. Fish 

and Wildlife Service, Denver Wildlife Research Center, Final Report, Outer Continental Shelf Environmental Assessment 

Program, Research Unit 341, Final Report 45 (1986), p. 1–120.

24. Wing, B.L., 1985, Salmon stomach contents from the Alaska troll logbook program 1977-84: U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration National Marine Fisheries Service, Technical Memorandum 

NMFS F/NWC-91, 43 p.

25. Albers, W.D., and Anderson, P.J., 1985, Diet of Pacific cod, Gadus macrocephalus, and predation on the northern pink 

shrimp, Pandalus borealis, in Pavlof Bay, Alaska: Fishery Bulletin, v. 83, no. 4, p. 601–610.

26. Hatch, S.A., and Sanger, G.A., 1992, Puffins as samplers of juvenile pollock and other forage fish in the Gulf of Alaska: 

Marine Ecology Progress Series, v. 80, p. 1–14.

27. Orlov, A.M., 1999, Some aspects of trophic relations among Pacific predatory fishes off the northern Kuril Islands and 

southeastern Kamchatka, in MacKinlay, D., and Houlihan, D., eds., Gutshop ‘98: American Fisheries Society, p. 41–52.
548 Alaska Arctic Marine Fish Ecology Catalog

Arctic Sand Lance (Ammodytes hexapterus) 

Pallas, 1814

Family Ammodytidae

Note: Until recently, it was assumed that the sand lance species 

found throughout the U.S. Arctic and southwards along both the 

eastern and western Pacific was Ammodytes hexapterus. Research 

has determined that beginning in the southeastern Bering Sea and 

particularly southward a second species, Ammodytes personatus,

replaces A. hexapturus. Because of this historical confusion data 

presented below may refer to one or both species.

Iñupiaq Name: Panmaksraq [1].

Ecological Role: Important high-lipid prey for many fishes, birds, and mammals throughout their range. Arctic Sand Lances 

are a critical summer food of nesting seabirds at Capes Lisburne and Thompson along the northwest coast of the Chukchi Sea. 

Their widespread geographic distribution and abundance and their mid-level trophic and food web importance makes them a 

significant component of marine ecosystems and a key species in the Alaskan high Arctic. The Arctic Sand Lance is a key forage 

species in the marine biological community due to its intermediate food web position and significant role in energy transfer 

between primary and higher-level consumers.

Physical Description and Attributes: Arctic Sand Lance have a metallic blue, elongate narrow body with a series of diagonal 

skin folds on sides, a fleshy ridge along body either side of the belly, a long dorsal fin that folds into a groove, and a projecting 

lower jaw (Fishes of Alaska [Mecklenburg and others, 2002, p. 795]) [2]. Swim bladder: Absent [2]. Antifreeze glycoproteins in 

blood serum: Unknown.

Range: Throughout the U.S. Chukchi and Beaufort Seas. Elsewhere in East Siberian, Chukchi, and Beaufort Seas eastward 

to Hudson and Ungava Bays; southward through eastern Bering Sea to Unimak Pass, and western Bering Sea to southeastern 

Kamchatka and Sea of Okhotsk [8].

Arctic Sand Lance (Ammodytes hexapterus), 149 mm TL, U.S. 

Chukchi Sea, 2007. Photograph by C.W. Mecklenburg, Point 

Stephens Research.
Arctic Sand Lance 549

Relative Abundance: Common, although patchily distributed, throughout the U.S. Chukchi and Beaufort Seas [6, 9–12]. 

Elsewhere regionally, common but not abundant in Canadian Beaufort Sea [11]. 

Geographic distribution of Arctic Sand Lance (Ammodytes hexapterus), within Arctic Outer Continental 

Shelf Planning Areas [7], based on review of published literature and specimens from historical and recent 

collections [6, 8]. 

tac14-5222_fig3-11-4m_Ammodytes_hexapterus

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Sand Lance

Ammodytes hexapterus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
550 Alaska Arctic Marine Fish Ecology Catalog

Benthic and reproductive distribution of Arctic Sand Lance (Ammodytes hexapterus). 

Pelagic larvae and young juveniles– 

as far offshore as central 

Chukchi Sea 

Open ocean Shore

Depth, in meters

Adults

Some fish to 172 m

Ammodytes hexapterus 

Arctic Sand Lance

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Juveniles Spawning

and eggs 

Adult data from outside

U.S.Beaufort-Chukchi Seas

Data from U.S. Chukchi Sea

Data from outside U.S.

250 Beaufort-Chukchi Seas

200

150

100

50

0

250

200

150

100

50

0

Habitats and Life History

Eggs—Size: perhaps 0.67–0.91 mm [20]. Time to hatching: 13–67 days (depending on water temperatures) 

[21–23]. Habitat: Eggs are demersal and slightly adhesive on coarse sand or fine gravel [17–19]. 

Larvae—Days to juvenile transformation: Unknown. Size: 4–7 mm at hatching to 40–80 mm FL at 

transformation [20]. Habitat: Pelagic in the Chukchi Sea, in open water at depths of at least 45 m below surface

[13, 14]. In coastal, shelf, and slope waters, depth to at least 149 m in the Gulf of Alaska [likely A. personatus] 

[24, 25]. 

Juveniles—Age and size: Mature during second full year [26], 40–80 mm to 88–113 mm FL [17, 20, 27]. 

Habitat: Has not been identified. Elsewhere throughout range, mainly pelagic. Juveniles frequently recruit to 

shallow neritic waters and remain for some time [16, 28]. Frequently occupied habitats include eelgrass and algae 

beds, sand, and bedrock [29, 30]. 

Adults—Age and size at first maturity: Unknown. In the Gulf of Alaska [likely A. personatus], most fish mature 

at slightly less than 2 years old (age-1) (a few as old as 4 years). Smallest ripe males can be 88 mm FL and 

smallest ripe females 113 mm FL [17]. Maximum age: Unknown. In the Gulf of Alaska [likely A. personatus], at 

least 6 years [17] and to about 11 years in Asian waters [16]. Maximum size: 28 cm TL [2, 5]. Habitat: Pelagic. 

Habitats frequently occupied include eelgrass and algae beds, sand, and bedrock [29, 30] in very shallow neritic 

waters; also found along the shallower parts of the continental shelf [28].

Substrate—Arctic Sand Lance burrow under soft substrate at night and during colder-water months. Other, they 

dwell in water column [23, 31].

Physical/chemical— Temperature: In the U.S. Beaufort Sea, -1.0 °C or less [10]. Salinity: in marine, brackish, 

and nearly fresh waters in Bering Sea and Gulf of Alaska [33–35].

Behavior

Diel—Unknown. In Gulf of Alaska [likely A. personatus], larvae migrate into deep waters at night [36]. In other 

regions, juveniles and adults school in water column by day (a few bury) and bury to depths of 10 cm in soft 

sediments of nearshore and shelf sea floors at night [18, 31, 34]. 

Seasonal—Unknown. In other areas, almost all fish spend coldest months under the sea floor in intertidal or 

subtidal waters, although occasionally individuals are taken in the water column pelagically [23]. Similarly, in 

warm-water seasons most or all fish remain buried at night and are most active in the water column during the day 

[31]. Burrowing fish apparently remain alert; even during the coldest months fish disturbed while in the sediment 

quickly move off [23]. 

Depth Range: Unknown. Larvae, juveniles, and adults documented near surface to 49 m [8, 13, 14] and at least in other areas 

to 100–120 m [8]. Spawning: Poorly known, although it is likely that A. personatus spawns in the intertidal and perhaps shallow 

subtidal waters [17–19].
Arctic Sand Lance 551

Reproductive—Unknown. In other areas, spawn both day and night in intertidal waters (and perhaps in the 

shallow subtidal) on fine gravel or coarse sand beaches in marine and brackish waters [17–19]. May spawn on 

the same beaches year after year [17]. Prior to spawning, large mixed schools of males and females (sometimes 

males predominate) form in shallow water. Large schools may remain close to spawning beaches for days at a 

time; large numbers occasionally are stranded on shore as tides fall [18]. Spawning can occur over any part of 

the tidal cycle; may peak at high tide. Just prior to spawning, females excavate shallow pits in which the eggs are 

laid. 

Schooling—Schooling primarily during daylight hours. Schools can be dense, monospecific, or composed of 

other species such as Pacific Herring [18, 31, 34]. 

Feeding—Unknown.

Populations or Stocks

DNA studies of fishes from the Hudson Bay-Beaufort Sea-Chukchi Sea-Bering Sea imply no distinct stocks [8].

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [17, 21, 22]. 

Spawning season—Highly variable among geographic locations and may occur from August into late spring 

[17, 19, 37]. 

Fecundity—Off Kamchatka Peninsula, 6,150–59,900 [16].

Food and Feeding

Food items—Food habits and nutritional requirements have not been described. Elsewhere, feed primarily on 

planktonic and epibenthic prey such as calanoid copepods, gammarid and hyperiid amphipods, crustacean larvae, 

and polychaete larvae [38–40]. 

Trophic level—3.5 [41].

Biological Interactions

Predators—In U.S. Chukchi Sea, Arctic Sand Lance are an important prey of both ringed (November to June) 

and bearded (February–March) seals, and seabirds (June–September) [42]. Other predators include Dolly 

Varden and Pacific Herring [9, 43, 44]. Elsewhere, sand lance are heavily preyed upon by a wide range of fishes, 

birds, and marine mammals [45]. Some seabird populations including Pigeon Guillemot are partially or mainly 

dependent on them [17, 46, 47].

Competitors—Presumably, competitors are Capelin and Arctic Cod and other consumers of midwater 

zooplankton 

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time is 1.4–4.4 years (t

m=2–3) [48].

Traditional and Cultural Importance

Unknown in the Alaskan High Arctic. Elsewhere in Alaska were used as food and bait by indigenous peoples, but 

not of great importance [49]. Occasionally harvested for bait [21].
552 Alaska Arctic Marine Fish Ecology Catalog

Commercial Fisheries

Currently, Arctic Sand Lance are not commercially harvested.

Potential Effects of Climate Change

As this species finds its southern boundary near the Aleutian Islands, it might be expected to retreat northwards 

as waters warm.

Areas for Future Research [A]

The ecology of Arctic Sand Lance is not well known for the U.S. Chukchi and Beaufort Seas. Of particular 

interest are standing stock, geographic and depth seasonal distribution, movements and migrations, spawning and 

overwintering grounds, and predator-prey relationships.

References Cited

Dick, M.H., and Warner, I.M., 1982, Pacific sand lance, Ammodytes hexapterus Pallas, in the Kodiak island group, Alaska: 

Syesis, v. 15, p. 43–50. [18]

Hobson, E.S.,1986, Predation on the Pacific sand lance, Ammodytes hexapterus (Pisces: Ammodytidae), during the transition 

between day and night in southeastern Alaska: Copeia, v. 1986, no. 1, p. 223–226. [31]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Robards, M.D., and Piatt, J.F., 1999, Biology of the genus Ammodytes, the sand lances, in Robards, M.D., and others, eds., 

Sand land—A review of biology and predator relations and annotated bibliography: U.S. Forest Service, Pacific Northwest 

Research Station, Research Paper PNW-RP-521. [23]

Robards, M.D., Piatt, J.F., and Rose, G.A., 1999, Maturation, fecundity, and intertidal spawning of Pacific sand lance in the 

northern Gulf of Alaska: Journal of Fish Biology v. 54, no. 5, p. 1,050–1,068. [17]

Bibliography

1. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

4. Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic marine fishes of Canada: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, no. 2674, 112 p.

5. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: U.S. Geological Survey, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

6. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.
Arctic Sand Lance 553

7. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

8. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

9. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

10. Moulton, L.L., and Tarbox, K.E., 1987, Analysis of Arctic cod movements in the Beaufort Sea nearshore region, 1978–79: 

Arctic, v. 40, no. 1, p. 43–49.

11. Bond, W.A., and Erickson, R.N., 1993, Fisheries investigations in coastal waters of Liverpool Bay, Northwest Territories: 

Winnipeg, Manitoba, Canada Department of Fisheries and Oceans, Central and Arctic Region, Canadian Manuscript Report 

of Fisheries and Aquatic Sciences No. 2204, 59 p.

12. Fechhelm, R.G., Baker, J.S., Griffiths, W.B., and Schmidt, D.R., 1989, Localized movement patterns of least cisco 

(Coregonus sardinella) and Arctic cisco (C. autumnalis) in the vicinity of a solid-fill causeway: Biological Papers of the 

University of Alaska, v. 24, p. 75–106.

13. Quast, J.C., 1974, Density distribution of juvenile Arctic cod, Boreogadus saida, in the eastern Chukchi Sea in the fall of 

1970: Fishery Bulletin, v. 72, no. 4, p. 1,094–1,105.

14. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

15. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

16. Tokranov, A.M., 2007, Distribution and some biological features of the Pacific sand lance Ammodytes hexapterus

(Ammodytidae) in waters off Kamchatka in the Sea of Okhotsk: Journal of Ichthyology, v. 47, no. 4, p. 288–295.

17. Robards, M.D., Piatt, J.F., and Rose, G.A., 1999, Maturation, fecundity, and intertidal spawning of Pacific sand lance in the 

northern Gulf of Alaska: Journal of Fish Biology v. 54, no. 5, p. 1,050–1,068.

18. Dick, M.H., and Warner, I.M., 1982, Pacific sand lance, Ammodytes hexapterus Pallas, in the Kodiak island group, Alaska: 

Syesis, v. 15, p. 43–50.

19. Moulton, L.L., and Penttila, D.E., 2000, Forage fish spawning distribution in San Juan County and protocols for sampling 

intertidal and nearshore regions: Mount Vernon, Washington, San Juan County Forage Fish Assessment Project, Northwest 

Straits Commission, 53 p.

20. Alaska Fisheries Science Center, 2013, Ichthyoplankton Information System: Alaska Fisheries Science Center, Accessed 

November 29, 2013, at http://access.afsc.noaa.gov/ichthyo/index.cfm.

21. Field, L.J., 1988, Pacific sand lance, Ammodytes hexapterus, with notes on related Ammodytes species, in Wilimovsky, N.J., 

Incze, L.S., and Westrheim, S.J., eds., Species synopses, life histories of selected fish and shellfish of the northeast Pacific 

and Bering Sea: Seattle, University of Washington, Washington Sea Grant Program and Fisheries Research Institute.

22. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

23. Robards, M.D., and Piatt, J.F., 1999, Biology of the genus Ammodytes, the sand lances, in Robards, M.D., and others, eds., 

Sand land—A review of biology and predator relations and annotated bibliography: U.S. Forest Service, Pacific Northwest 

Research Station, Research Paper PNW-RP-521.

24. Siefert, D.L., Incze, L.S., and Ortner, P.B., 1988, Vertical distribution of zooplankton, including ichthyoplankton, in 

Shelikof Strait, Alaska—Data from fisheries-oceanography coordinated investigations (FOCI) cruise in May 1986: National 

Marine Fisheries Service, NWAFC Processed Report, v. 88, no. 28, 232 p.
554 Alaska Arctic Marine Fish Ecology Catalog

25. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.

26. Hamada, T., 1966, Studies on fluctuations in the abundance of larval sand-lance in the Harima-nada and Osaka Bay I—

Relation between the progeny abundance and the age composition of parent fish: Bulletin of the Japanese Society of 

Fisheries Oceanography, v. 32, no. 5, p. 393–398.

27. Blackburn, J.E., and Jackson, P.B., 1982, Seasonal composition and abundance of juvenile and adult marine finfish and crab 

species in the nearshore zone of Kodiak Island’s eastside during April 1978 through March 1979: Outer Continental Shelf 

Environmental Assessment Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 377–570.

28. Orsi, J.A., and Landingham, J.H., 1985, Numbers, species, and maturity stages of fish captured with beach seines during 

spring 1981 and 1982 in some nearshore marine waters of southeastern Alaska: National Oceanic and Atmospheric 

Administration, Technical Memorandum NMFS F/NWC-86, 68 p.

29. Fresh, K.L., 1979, Distribution and abundance of fishes occurring in the nearshore surface waters of northern Puget Sound, 

Washington: Seattle, University of Washington, Master’s thesis.

30. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 98 p.

31. Hobson, E.S.,1986, Predation on the Pacific sand lance, Ammodytes hexapterus (Pisces: Ammodytidae), during the 

transition between day and night in southeastern Alaska: Copeia, v. 1986, no. 1, p. 223–226.

32. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

33. Rosenthal, R.J., 1980, Shallow water fish assemblages in the northeastern Gulf of Alaska—Habitat evaluation, species 

composition, abundance, spatial distribution and trophic interaction, in Bureau of Land Management, Environmental 

assessment of the Alaskan Continental Shelf, final reports of principal investigators: U.S. Department of Commerce, 

National Oceanic and Atmospheric Administration, p. 451–540.

34. Blackburn, J.E., Jackson, P.B., Warner, I.M., and Dick, M.H., 1981, A survey for spawning forage fish on the east side of the 

Kodiak Archipelago by air and boat during spring and summer 1979: Outer Continental Shelf Environmental Assessment 

Program, Final Reports of Principal Investigators, RU 0552, p. 309–376.

35. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

36. Brodeur, R.D., and Rugen, W.C., 1994, Diel vertical distribution of ichthyoplankton in the northern Gulf of Alaska: Fishery 

Bulletin, v. 92, no. 2, p. 223–235.

37. Busby, M.S., Matarese, A.C., and Mier, K.L., 2000, Annual, seasonal, and diel composition of larval and juvenile fishes 

collected by dip-net in Clam Bay, Puget Sound, Washington, from 1985 to 1995: Seattle, Washington, U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-111, 48 p.

38. Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, Fisheries Research Institute, FRI-UW-7719.

39. Simenstad, C.A., Isakson, J.S., and Nakatani, R.E., 1977, Marine fish communities, in Merritt, M.L., and Fuller, R.G., eds., 

The environment of Amchitka Island, Alaska: National Technical Information Center, Energy Research and Development 

Administration TID-26712, p. 451–492.

40. Cross, J.N., Fresh, K., Miller, B.S., Simenstad, C.A., Steinfort, S.N., and Fegley, J.C., 1978, Nearshore fish and 

invertebrates assemblages along the Strait of Juan de Fuca including food habits of the common inshore fishes: U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, Environmental Research Laboratories, 

NOAA Technical Memorandum ERL MESA-32.

41. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.
Arctic Sand Lance 555

42. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.

43. Furniss, R.A., 1975, Inventory and cataloging of Arctic area waters: Alaska Department of Fish and Game, Federal Aid in 

Fish Restoration, Annual Report of Performance, Project F-9-7, Study G-1-1, v. 16, 47 p.

44. Bond, W.A., and Erickson, R.N., 1987, Fishery data from Phillips Bay, Yukon, 1985: Winnipeg, Manitoba, Canadian Data 

Report of Fisheries and Aquatic Sciences, Central and Artic Region, Department of Fisheries and Oceans, no. 635, 47 p.

45. Willson, M.F., Armstrong, R.H., Robards, M.D., and Piatt, J.F., 1999, Sand lance as cornerstone prey for predator 

populations in Robards, M.D., and others, eds., Sand land—A review of biology and predator relations and annotated 

bibliography: U.S. Forest Service, Pacific Northwest Research Station, Research Paper PNW-RP-521.

46. Hayes, D.L., and Kuletz, K.J., 1997, Decline of pigeon guillemot populations in Prince William Sound, Alaska, and 

apparent changes in distribution and abundance of their prey, in Forage fishes in marine ecosystems—Proceedings of the 

International Symposium on the Role of Forage Fishes in Marine Ecosystems: University of Alaska, Fairbanks, Alaska Sea 

Grant College Program, p. 699–702.

47. Litzow, M.A., Piatt, J.F., Prichard, A.K., and Roby, D.D., 2002, Response of pigeon gullemots to variable abundance of 

high-lipid and low-lipid prey: Oecologia, v. 132, no. 2, p. 286–295.

48. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

49. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.
556 Alaska Arctic Marine Fish Ecology Catalog

Bering Flounder (Hippoglossoides robustus) 

Gill & Townsend, 1897

Family Pleuronectidae 

Note: This species may be a synonym of Hippoglossoides elassodon 

Jordan & Gilbert, 1880 [3].

Colloquial Name: None in U.S. Chukchi and Beaufort Seas. 

Ecological Role: Bering Flounder are the most common flatfish in the 

U.S. Chukchi Sea [1, 2] and are one of the most abundant fishes found 

there [2, 3]. Shifts in this species range in the northern Bering Sea make 

it an important population to monitor with respect to climate change.

Physical Description/Attributes: Member of the right eyed flounders.

Eyed side is reddish brown to grayish brown and blind side is white. 

For specific diagnostic characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, p. 829) [4]. Swim bladder: Absent [4]. 

Antifreeze glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea as far north as 72°19’N, 175°55’W [2], and U.S. Beaufort Sea [5, 6]. Elsewhere in Alaska, Bering 

Flounder is found from Bering Sea off Alaska Peninsula, west to Akutan Island, Aleutian Islands [4]. Worldwide, from East 

Siberian Sea eastward to Bathurst Inlet, south of Dease Strait, Nunavut in Canadian Arctic, and in northern Sea of Japan off 

Hokkaido, Okhotsk Sea, Pacific Ocean off Kamchatka Peninsula, and Commander Islands [4, 6, 7]. 

Relative Abundance: Common in U.S. Chukchi and western Beaufort Seas [2, 5, 6]. Elsewhere, common in Bering Sea and to 

Kuril and Sakhalin Islands, Russia [9–11].

Bering Flounder (Hippoglossoides robustus), 185 mm, 

western Chukchi Sea, 2004. Photograph by C.W. 

Mecklenburg, Point Stephens Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Bering Flounder

Hippoglossoides robustus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Bering Flounder (Hippoglossoides robustus), within Arctic Outer Continental Shelf 

Planning Areas [8] based on review of published literature and specimens from historical and recent collections 

[3, 6]. 
Bering Flounder 557

Benthic and reproductive distribution of Bering Flounder (Hippoglossoides robustus).

Juveniles and adults

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

 Overall depth range data from 

outside U.S. Beaufort-Chukchi Seas 

Spawning data from outside

U.S. Beaufort-Chukchi Seas

Eggs and larvae

Spawning

Depth range of eggs and larvae

are from U.S. Chukchi Sea

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Hippoglossoides robustus

Bering Flounder

Benthic distribution

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Reproductive distribution

Some fish to 532 m

Depths documented in 

U.S. Chukchi Sea 

Habitats and Life History

Eggs—Size: 2.4–2.7 mm [16]. Time to hatching: Unknown. Habitat: Pelagic [17]. 

Larvae—Size at hatching: 4 mm SL [17]. Size at juvenile transformation: Greater than 28.6 mm SL [18]. Days 

to juvenile transformation: Unknown. Habitat: Pelagic [17]. 

Juveniles—Age and size: Unknown, greater than 28.6 mm SL to 25 cm TL [18, 19]. Habitat: Demersal [6]. 

Adults—Age and size at first maturity: Females mature as small as 25 cm TL. Females grow larger and faster 

than males [19]. Maximum age: 30 years [20]. Maximum size: 52 cm TL [4]. Habitat: Demersal [6]. 

Substrate—Mud sea floors [2]. 

Physical/chemical—Temperature: -1.8–7.9 °C in U.S. Chukchi Sea [6]. Elsewhere, between -1.7 and 9.2 °C [5, 

13]. Salinity: Marine. Documented between 29.4 and 33.5 parts per thousand in northeastern Chukchi Sea [1].

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Spawns in shallow bays and gulfs [21]. 

Schooling—Unknown. 

Feeding—Unknown.

Populations or Stocks

There have been no studies. Population information is available from the Bering Sea.

Reproduction

Mode—Unknown. 

Mating/Spawning Season—Between April and June in western Bering Sea and off Kamchatka Island and may 

extend into August in Sea of Okhotsk [13]. In U.S. Chukchi Sea, pelagic eggs were taken in August [12].

Fecundity—Unknown. 

Depth Range: Documented at 40–72 m in southern U.S. Chukchi Sea [12]. Elsewhere, intertidal to 425 m, typically 150 m or 

less [3, 6, 13–15]. Spawning occurs primarily at 25–130 m [13]. In northern Chukchi Sea, pelagic eggs documented from near 

surface to 81 m. Larvae in midwaters from near surface to 81 m [12].
558 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—In U.S. Chukchi Sea, fishes (for example, eel blennies, poachers, sculpins, and cods) are a major 

part of the diet, along with such benthic and epibenthic crustaceans as amphipods, shrimps, and hermit crabs [22]. 

Trophic level—3.7 [23].

Biological Interactions

Predators—Arctic Cod, in U.S. Chukchi Sea [22]. Pacific Halibut, bearded seals, and beluga whales in Bering Sea, 

[1, 24]. 

Competitors—Presumably a wide range of other zoobenthos feeders such as Arctic Cod, Walleye Pollock, other 

flounders, eelpouts, and sculpins. 

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (t

max=11; K=0.21) [25].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Currently, Bering Flounder are not commercially harvested.

Potential Effects of Climate Change

As a predominantly northern species with a low water temperature preference [26], it would be expected that 

climate warming would result in a northward shift in range. 

Areas for Future Research [A]

Little is known about the ecology of this species in the study area. Research needs for this species include: 

(1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment; 

(3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic movements; 

(6) population studies; (7) prey; (and 8) predators. The Bering Flounder is an important shelf species and 

monitoring programs should be designed such that shifts in abundance and changes in vital statistics can 

be detected.
Bering Flounder 559

References Cited 

Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R—Keys to the fauna in the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from the U.S. Department of Commerce, Springfield, Virginia.] [19]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [4]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [6]

Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi Sea: 

Deep-Sea Research II, v. 57, no. 1–2, p. 57–70. [12]

Smith, R.L., Vallarino, M., Barbour, E., Fitzpatrick, E., and Barber, W.E., 1997, Population biology of the Bering flounder in the 

Northeastern Chukchi Sea: American Fisheries Society Symposium, v. 19, p. 127–132. [1]

Bibliography

1. Smith, R.L., Vallarino, M., Barbour, E., Fitzpatrick, E., and Barber, W.E., 1997b, Population biology of the Bering flounder 

in the Northeastern Chukchi Sea: American Fisheries Society Symposium, v. 19, p. 127–132.

2. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

3. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

4. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

5. Pruter, A.T., and Alverson, D.L., 1962, Abundance, distribution, and growth of flounders in the south-eastern Chukchi Sea: 

Journal de Conseil International pour l’Exploration de la Mer., v. 27, p. 79–99.

6. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

7. Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic marine fishes of Canada: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences, Fisheries and Oceans Canada, no. 2674, 112 p.

8. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

9. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

10. Dudnik, Y.I., and Dolganov, V.N., 1992, Distribution and abundance of fish on the continental slopes of the Sea of Okhotsk 

and of the Kuril Islands during the summer of 1989: Journal of Ichthyology, v. 32, no. 9, p. 58–76.

11. Kim, S.T., and Shepeleva, O.N., 2001, The structure of shelf ichthyocenoses of northeastern Sakhalin and Terpeniya Bay: 

Journal of Ichthyology, v. 41, no. 9, p. 711–722.

12. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

13. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224.
560 Alaska Arctic Marine Fish Ecology Catalog

14. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

15. Fedorov, V.V., 2000, Species composition, distribution and habitation depths of the northern Kuril Islands fish and fish like 

species, in Koteneva, B.N., ed., Commercial and biological studies of fishes in the Pacific waters of the Kuril Islands and 

adjacent areas of the Okhotsk and Bering Seas in 1992–1998: Moscow, VNIRO Publishing, collected papers, p. 7–41. [In 

Russian.]

16. Grigorev, S.S., and Fadeev, N.S., 1995, Distribution of Alaska plaice, flathead sole, and Bering flounder eggs in the eastern 

Bering Sea during April–July, 1988–1990, in Alaska Sea Grant Program, eds., Proceeding of the International Symposium 

on North Pacific Flatfish: Fairbanks, University of Alaska, Alaska Sea Grant Program, AK-SG-95-04.

17. Matarese, A.C., Kendall, J., Arthur W., Blood, D.M., and Vinter, B.M., 1989, Laboratory guide to early life history stages 

of northeast Pacific fishes: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National 

Marine Fisheries Service Technical Report NMFS 80, 652 p.

18. Ahlstrom, E.H., Amaoka, K., Hensley, D.A., Moser, H.G., and Sumida, B.Y., 1984, Pleuronectiformes—Development, 

Moser, H.G., and others, eds., in Ontogeny and systematics of fishes: American Society of Ichthyologists and 

Herpetologists.

19. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R—Keys to the fauna in the U.S.S.R.: Academy of Sciences 

of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific Translation, 

Jerusalem, 1964, 617 p., available from the U.S. Department of Commerce, Springfield, Virginia.]

20. National Oceanic and Atmospheric Administration, 2010, Age and growth program: National Oceanic and Atmospheric 

Administration, accessed April 2011 at http://www.afsc.noaa.gov/refm/age/Stats/max_age.htm.

21. Pertseva-Ostroumova, T.A., 1961, The reproduction and development of far-eastern flounders: Moscow, Academy Nauk 

U.S.S.R. Institute Okeanol, 486 p. [In Russian.]

22. Coyle, K.O., Gillispie, J.A., Smith, R.L., and Barber, W.E. 1997, Food habits of four demersal Chukchi Sea fishes, in

Reynolds, J.B., ed., Symposium 19, Proceedings of the Fish Ecology in Arctic North America Symposium: American 

Fisheries Society, Fairbanks, Alaska, May 19–21, 1992, p. 310–318.

23. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.

24. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-158, 249 p.

25. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

26. Moiseev, P.A., 1953, Cod and flounders of far-eastern seas: Izvestiya TINRO, Proceedings Pacific Science Research 

Institute of Marine Fisheries and Oceanography, v. 40, p. 1–287.
Pacific Halibut 561

Pacific Halibut (Hippoglossus stenolepis) 

Schmidt, 1904

Family Pleuronectidae

Note: Except for geographic range data, all information is from 

areas outside the study area.

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Pacific Halibut are absent from the U.S. Beaufort 

Sea and only rarely observed in the southeastern Chukchi Sea. At 

present, the species ecological role in marine ecosystem function is 

minor.

Physical Description/Attributes: Member of the right eyed 

flounders. Eyed side of body is grayish to greenish brown or 

black with various light and dark mottling. The blind side is 

white. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, p. 823) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Unknown.

Range: Northeastern U.S. Chukchi Sea to 71°12’N, 163°05’W [2]. Elsewhere in Alaska, throughout Bering Sea, Aleutian 

Islands, Gulf of Alaska and all southeastern Alaska waters [1]. Worldwide, to Pacific off Hokkaido, Japan and Sea of Okhotsk, 

and to Punta Camalu, Baja California [1, 3–5]. 

Pacific Halibut (Hippoglossus stenolepis), 633 mm, 

southeastern Bering Sea. Photograph by C.W. Mecklenburg, 

Point Stephens Research.
562 Alaska Arctic Marine Fish Ecology Catalog

Relative Abundance: Absent from U.S. Beaufort Sea and rare in U.S. Chukchi Sea [2]. Uncommon in northern Bering Sea, 

but common in eastern Bering Sea as far north as Norton Sound [2]. Resource assessment data suggest a northward shift in this 

species’ distribution in the Bering Sea [8]. Common west to Sea of Okhotsk and northern Kuril Islands, Russia, and southward 

to the Pacific Ocean off Oregon [1, 3–5]. 

tac14-5222_fig3-11-6m_Hippoglossus_stenolepis

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Pacific Halibut

Hippoglossus stenolepis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Pacific Halibut (Hippoglossus stenolepis), within Arctic Outer Continental Shelf Planning 

Areas [6] based on review of published literature and specimens from historical and recent collections [2, 7]. 
Pacific Halibut 563

Some fish to 1,100 m

Open ocean Shore

Depth, in meters

1,250

1,000

750

500

250

0

1,250

1,000

750

500

250

0

Larvae

Spawning

Eggs

Depths documented in U.S. Chukchi Sea

Overall depth data from outside

U.S. Chukchi and Beaufort Seas 

Data from outside 

U.S. Chukchi and Beaufort Seas

Hippoglossus stenolepsis

Pacific Halibut

Benthic distribution

Open ocean Shore

Depth, in meters

Reproductive distribution

Adults

Juveniles

Benthic distribution and reproductive distribution of Pacific Halibut (Hippoglossus stenolepis). 

Habitats and Life History

Eggs—Size: 2.9–3.8 mm [12, 13]. Time to hatching: 15–20 days at 5–6 °C and 12–14 days at 7–8 °C [12, 13]. 

Habitat: Epipelagic to mesopelagic [12–15]. 

Larvae—Size at hatching: 8.0 mm SL [13]. Size at juvenile transformation: 2.2 cm TL [17]. Days to juvenile 

transformation: About 180 days [12, 17, 19]. Habitat: Pelagic, over continental shelf and slope [13–16]. 

Juveniles—Age and size: Age unknown. 2.2 cm TL to 72–120 cm FL [12, 17, 23, 24]. Habitat: Benthic [25]. 

Younger juveniles settle on fine sand sea floors in or near bays [12, 17–19]. Older juveniles are on coarsergrained sediments and migrate into deeper waters [12, 19]. 

Adults—Age and size at first maturity: 8 years (72 cm FL) for males and 12 years (120 cm FL) for females 

[12, 23, 24]. Females grow faster than males and males mature when younger. Growth rates vary by area [12]. 

Maximum age: To at least 55 years old [12]. Maximum size: 267 cm TL [1]. The heaviest documented fish 

weighed 227 kg, but fish to 318 kg have been reported. Males rarely reach 36 kg [12]. Habitat: Benthic [25]. 

Substrate—Mostly associated with soft sea floors, but also rest on rocky bottoms [26]. 

Physical/chemical—Temperature: Between -1.7 and 12.2 °C [27–29], mainly at 3–8 °C [12]. Salinity: Marine, 

occasionally found in somewhat estuarine waters [25].

Behavior

Diel—Juveniles and adults bury themselves in bottom sediments to escape predators, reduce their metabolic rate 

to resting phase, and to avoid fast currents [19, 30]. 

Seasonal—Overall movements vary considerably between individuals and between areas. Adults tend to remain 

on the same grounds from year to year, making only seasonal migrations from shallow feeding grounds in 

summer to deeper spawning grounds in winter [12]. Individuals appear to home to these grounds year after year 

[22]. Some individuals exhibit only limited movements [22, 31, 32]. However, fish tagged in both the Bering Sea 

and in the north and northeast Pacific Ocean traveled many hundreds of kilometers [12, 31]. Generally, relatively 

few fish appear to move between the Gulf of Alaska and Bering Sea [12, 32]. Egg and larvae spawned off 

southeastern Alaska and British Columbia tend to drift northwest into the Gulf of Alaska, and some juveniles that 

have settled there move back to the southeast as they mature [12, 33]. 

Depth Range: Adults and juveniles documented in U.S. Chukchi Sea from 2–51 m [1, 2, 9]. Overall, 6–1,200 m [7], typically 

less than 600 m [10, 11]. Eggs are found at 92–700 m [12–14] and larvae from surface waters to 300 m [15]. Larvae are found 

over the continental shelf and slope [16]. Juveniles are mostly in depths of 40 m or less [12, 17–19]. Spawning is often at 

183–457 m [20–22].
564 Alaska Arctic Marine Fish Ecology Catalog

Reproductive—As waters cool, adults migrate offshore to spawning grounds on deeper parts of continental shelf 

and slope [15, 21]. Spawning grounds are relatively circumscribed and discrete and occur from at least British 

Columbia to the Pribilof Canyon area in the eastern Bering Sea and likely along the Aleutian Islands [12, 20–22]. 

Males ripen earlier than females and extrude milt after females have finished spawning. Females shed ripe eggs 

gradually and spawn over an extended period, as all eggs are not ripe at any given time [34]. Some adults may 

not spawn every year [21, 22]. Potential courtship behavior may include a spawning pair rapidly ascending 

100–175 m off the seafloor, releasing gametes, and immediately returning to the bottom [21, 28]. 

Schooling—Adults have been found schooling at the edge of the continental shelf in October and November 

prior to spawning [34]. 

Feeding—They are strong swimmers who are found feeding off-bottom [22].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external. 

Spawning season—Spawning occurs from November to March in eastern Bering Sea [12, 24]. 

Fecundity—Between 102,000 and more than 4 million eggs [12, 24].

Food and Feeding

Food items—Young-of-the-year halibut feed on small crustaceans (for example, mysids, cumaceans, and 

gammarid amphipods), whereas older juveniles add crabs, shrimps, and small fishes to their diets [35, 36]. As 

halibut grow, crabs remain important, but fishes (for example, Walleye Pollock, Capelin, and Pacific Sand Lance) 

may make up more than half of the diet. Other important preys include hermit crabs, squids, octopi, and snails 

[36–38]. 

Trophic level—4.6 [8].

Biological Interactions

Predators—Alaska Skate, Pacific Cod, Pacific Halibut, Walleye Pollock, bald eagles, Steller sea lions, and orcas 

[36, 38–42]. 

Competitors—Larger fishes, such as Pacific Cod and Walleye Pollock, and marine mammals such as orcas and 

seals.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (rm=0.2; K=0.05; t

m=5–20; tmax=30) [43].

Traditional and Cultural Importance

Pacific Halibut were a major food fish for indigenous peoples living from the Bering Sea to Washington [44, 45], 

but their rarity in the Chukchi Sea precludes widespread use by coastal residents. 
Pacific Halibut 565

Commercial Fisheries

Currently, Pacific Halibut are not commercially harvested.

Potential Effects of Climate Change

A northward shift in distribution is expected into the Chukchi Sea. Pacific Halibut are likely to increase in 

abundance, especially in the southeastern Chukchi Sea and perhaps gradually expand their range into the 

Beaufort Sea.

Areas for Future Research [B]

This is a well-studied marine fish due to its commercial importance elsewhere. As the species range expands, 

additional research will be needed on the location of important habitats, species reproductive ecology (depth and 

location of eggs and larvae), life history (age and size at maturity), and population dynamics. 

Remarks

A condition called “Chalky” halibut occurs in a substantial number of landed fish. This condition, which renders the fish 

unsalable due to dry and powdery musculature, is caused by high lactic acid buildup in the muscles, and is apparently linked to 

stress at capture, along with precapture physiology and water conditions [46]. 

References Cited 

International Pacific Halibut Commission, 1998, The Pacific halibut—Biology, fishery, and management: Seattle, Washington, 

International Pacific Halibut Commission Technical Report, no. 40. [12]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Seitz, A.C., Loher, T., and Nielsen, J.L., 2008, Seasonal movements and environmental conditions experienced by Pacific halibut 

along the Aleutian Islands, examined by pop-up satellite tags: International Pacific Halibut Commission Scientific Report, 

no. 85, 24 p. [22]

Seitz, A.C., Norcross, B.L., Wilson, D., and Nielsen, J.L., 2005, Identifying spawning behavior in Pacific halibut, Hippoglossus 

stenolepis, using electronic tags: Environmental Biology of Fishes, v. 73, p. 445–451. [21]

Stoner, A.W., and Abookire, A.A., 2002, Sediment preferences and size-specific distribution of young-of-the-year Pacific halibut 

in an Alaska nursery: Journal of Fish Biology, v. 61, no. 3, p. 540–559. [19]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.
566 Alaska Arctic Marine Fish Ecology Catalog

4. Hamazaki, T., Fair, L., Watson, L., and Brennan, E., 2005, Analyses of Bering Sea bottom-trawl surveys in Norton Sound—

Absence of regime shift effects on epifauna and demersal fish: ICES Journal of Marine Science, v. 62, no. 8, p. 1,597–1,602.

5. Orlov, A.M., 2003, Impact of eddies on spatial distributions of groundfishes along waters off the northern Kuril Islands, and 

southeastern Kamchatka (north Pacific Ocean): Indian Journal of Marine Sciences, v. 32, no. 2, p. 95–111.

6. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

7. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

8. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320. 

9. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

10. Hoff, G.R., and Britt, L.L., 2003, The 2002 Eastern Bering Sea upper continental slope survey of groundfish and 

invertebrate resources: U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical 

Memorandum, NMFS-AFSC-141, Department of Commerce, 286 p.

11. Hoff, G.R., and Britt, L.L., 2005, Results of the 2004 Eastern Bering Sea upper continental slope survey of groundfish and 

invertebrate resources: National Oceanic and Atmospheric Administration Technical Memorandum, 300 p. 

12. International Pacific Halibut Commission, 1998, The Pacific halibut—Biology, fishery, and management: Seattle, 

Washington, International Pacific Halibut Commission Technical Report, no. 40.

13. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

14. Bailey, K.M., Abookire, A.A., and Duffy-Anderson, J.T., 2008, Ocean transport paths for the early life history stages of 

offshore-spawning flatfishes—A case study in the Gulf of Alaska: Fish and Fisheries, v. 9, no. 1, p. 44–66.

15. Bailey, K.M., and Picquelle, S.J., 2002, Larval distribution of offshore spawning flatfish in the Gulf of Alaska—Potential 

transport pathways and enhanced onshore transport during ENSO events: Marine Ecology Progress Series, v. 236, 

p. 205–217.

16. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.

17. Norcross, B.L., Müter, F.-J., and Holladay, B.A., 1997, Habitat models for juvenile pleuronectids around Kodiak Island, 

Alaska: Fishery Bulletin, v. 95, no. 3, p. 504–520.

18. Norcross, B.L., Blanchard, A., and Holladay, B.A., 1999, Comparison of models for defining nearshore flatfish nursery areas 

in Alaska waters: Fisheries Oceanography, v. 8, p. 50–67.

19. Stoner, A.W., and Abookire, A.A., 2002, Sediment preferences and size-specific distribution of young-of-the-year Pacific 

halibut in an Alaska nursery: Journal of Fish Biology, v. 61, no. 3, p. 540–559.

20. Beamish, R.J., McFarlane, G.A., and King, J.R., 2005, Migratory patterns of pelagic fishes and possible linkages between 

open ocean and coastal ecosystems off the Pacific coast of North America: Deep-Sea Research II, v. 52, p. 739–755.

21. Seitz, A.C., Norcross, B.L., Wilson, D., and Nielsen, J.L., 2005, Identifying spawning behavior in Pacific halibut, 

Hippoglossus stenolepis, using electronic tags: Environmental Biology of Fishes, v. 73, p. 445–451.
Pacific Halibut 567

22. Seitz, A.C., Loher, T., and Nielsen, J.L., 2008, Seasonal movements and environmental conditions experienced by Pacific 

halibut along the Aleutian Islands, examined by pop-up satellite tags: International Pacific Halibut Commission Scientific 

Report, no. 85, 24 p.

23. Best, E.A., 1981, Halibut ecology, in Hood, D.W., and Calder, J.A., eds., The eastern Bering Sea shelf—Oceanography and 

resources: Seattle, University of Washington Press, p. 495–508.

24. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

25. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

26. Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat associations of demersal fishes and crabs in the Pribilof Islands 

region of the Bering Sea: Fisheries Research, v. 75, no. 1–3, p. 15–28.

27. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

28. Seitz, A.C., Norcross, B.L., and Nielsen, J.L., 2003, Pip-up archival transmitting (PAT) tags—A method to investigate the 

migration and behavior of Pacific halibut Hippoglossus stenolepsis in the Gulf of Alaska: Alaska Fishery Research Bulletin, 

v. 10, no. 2, p. 124–135.

29. Novikov, N.P., 1964, Basic elements of the biology of the Pacific halibut (Hippoglossus hippoglossus stenolepis Schmidt) 

in the Bering Sea, in Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part II: Jerusalem, Israel 

Program for Scientific Translations, p. 175–219.

30. Moles, A., and Norcross, B.L., 1995, Sediment preference in juvenile Pacific flatfishes: Netherlands Journal of Sea 

Research, v. 34, no. 1–3, p. 177–182.

31. Loher, T., and Seitz, A.C., 2006, Seasonal migration and environmental conditions of Pacific halibut Hippoglossus 

stenolepis, elucidated from pop-up archival transmitting (PAT) tags: Marine Ecology Progress Series, v. 317, p. 259–271.

32. Seitz, A.C., Loher, T., and Nielsen, J.L., 2008, Seasonal movements and environmental conditions experienced by Pacific 

halibut along the Aleutian Islands, examined by pop-up satellite tags: International Pacific Halibut Commission Scientific 

Report, no. 85, 24 p.

33. Hauser, L., Spies, I., and Loher, T., 2006, Microsatellite screening in Pacific halibut (Hippoglossus stenolepis) and a 

preliminary examination of population structure based on observed DNA variation: Seattle, Washington, International 

Pacific Halibut Commission Scientific Report, no. 81.

34. St.-Pierre, G., 1984, Spawning locations and season for Pacific halibut: Seattle, Washington, International Pacific Halibut 

Commission Scientific Report no. 70, 46 p.

35. Holladay, B.A., and Norcross, B.L., 1995, Diet diversity as a mechanism for partitioning nursery grounds of pleuronectids: 

Proceedings of the International Symposium on North Pacific Flatfish, Alaska Sea Grant AK-SG-95-04, p. 177–203.

36. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration Technical Memorandum NMFS-AFSC-158, 249 p.

37. Yang, M.-S., 1996, Diets of the important groundfishes in the Aleutian Islands in summer 1991: Seattle, Washington, U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFSAFSC-60, 105 p.

38. Yang, M.-S., and Nelson, M.W., 2000, Food habits of the commercially important groundfishes in the Gulf of Alaska 

in 1990, 1993, and 1996: Seattle, Washington, U.S. Department of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFS-AFSC-112, 174 p.

39. Ofelt, C.H., 1975, Food habits of nesting bald eagles in southeast Alaska: The Condor, v. 77, p. 337–338.
568 Alaska Arctic Marine Fish Ecology Catalog

40. Kajimura, H., and Loughlin, T.R., 1988, Marine mammals in the oceanic food web of the eastern subarctic Pacific: 

University of Tokyo, Bulletin of the Ocean Research Institute, v. 26, no. 2, p. 187–233.

41. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990. 

42. Yang, M.-S., 2003, Food habits of the important groundfishes in the Aleutian Islands in 1994 and 1997: National Marine 

Fisheries Service, AFSC Processed Report 2003-07, 233 p.

43. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

44. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

45. Stewart, H., 1977, Indian fishing—Early methods on the northwest coast: Seattle, University of Washington Press, 182 p.

46. Foy, R.J., Crapo, C.A., and Kramer, D.E., 2006, Investigating the roles of temperature and exercise in the development of 

chalkiness in Pacific halibut: International Pacific Halibut Commission, Technical Report No. 50.
Yellowfin Sole 569

Yellowfin Sole (Limanda aspera) 

(Pallas, 1814)

Family Pleuronectidae 

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Yellowfin Sole are likely of only modest 

ecological importance in the U.S. Chukchi Sea and are unimportant 

in the U.S. Beaufort Sea. Increases in abundance associated with 

warming may attract commercial interests.

Physical Description/Attributes: Member of the right eyed 

flounders. Eyed side is brown and smaller individuals have dark 

spots on head and body. Median fins are yellowish, there is a narrow 

black line at base of dorsal and anal fins, and blind side is white. For 

specific diagnostic characteristic, see Fishes of Alaska (Mecklenburg 

and others, 2002, p. 844) [1]. Swim bladder: Absent [1]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea to U.S. Beaufort Sea at about 71°14’N, 155°35’W, east of Point Barrow [2]. Elsewhere in Alaska, 

throughout Bering Sea, eastern Aleutian Islands, Gulf of Alaska and all southeastern waters. Worldwide, in western Pacific 

Ocean southward to Sea of Japan off Korea and Okhotsk Sea and southward in eastern Pacific Ocean to Barkley Sound, British 

Columbia [1].

Yellowfin Sole (Limanda aspera), 185 mm, U.S. Chukchi Sea, 

2007. Photograph by C.W. Mecklenburg, Point Stephens 

Research.
570 Alaska Arctic Marine Fish Ecology Catalog

Geographic distribution of Yellowfin Sole (Limanda aspera), within Alaska Outer Continental Shelf Planning 

Areas [3] based on review of published literature and specimens from historical and recent collections 

[2, 4]. 

tac14-5222_fig3-11-7m_Limanda_aspera

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Yellowfin Sole

Limanda aspera

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Relative Abundance: Common, although not abundant, in southern U.S. Chukchi Sea, rare farther north and east and in 

U.S. Beaufort Sea [2]. Common throughout much of Bering Sea (particularly to as far north as 61°N) and eastward to Kenai 

Peninsula and Cook Inlet [5–8]. Common from Sea of Japan [9] and Sea of Okhotsk [10].
Yellowfin Sole 571

Depth Range: At 10–600 m, typically 150 or less [1]. Winters at 100–270 m in eastern Bering Sea [5]. Spawning occurs 

nearshore to a depth of 50 m [5], but mainly at 30 m or less [11]. In U.S. Chukchi Sea, eggs of Limanda sp. taken as deep as 29 

m, or to surface [12]. Eggs at surface in Sea of Japan [13]. In U.S. Chukchi Sea, larvae and young juveniles taken in midwater as 

deep as 37 m, as shallow as 25, m and perhaps to surface [12]. Older juveniles are nearshore, from intertidal to 50 m [14, 15].

Benthic and reproductive distribution of Yellowfin Sole (Limanda aspera). 

Some fish to 600 m

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Limanda aspera

Yellowfin Sole

Benthic distribution

Depth, in meters

Reproductive distribution

Open ocean Shore

125

100

75

50

25

0

Egg and larvae data from U.S. Chukchi Sea

Spawning data from outside U.S. Beaufort-Chukchi Seas

Larvae Eggs and larvae

Adults

Juveniles

Data from outside U.S. Beaufort-Chukchi Seas

Spawning

Habitats and Life History 

Eggs—Size: 0.8–0.9 mm [18]. Time to hatching: 100 hours at 13 °C [16]. Lower threshold temperature for 

successful egg development is about 4.0 °C [17]. Eggs are colorless or pale gray with a very slight greenish tinge 

[19]. Habitat: Epipelagic [12, 13, 20]. 

Larvae—Size at hatching: 2.2–3.1 mm SL [17, 20]. Size at juvenile transformation: As small as 1.2 cm SL [14, 

15], but usually 1.5–1.7 cm SL [20]. Days to juvenile transformation: One month, but may continue as much as 

4 months [18]. Habitat: Epipelagic [12, 17, 18, 20]. 

Juveniles—Age and size: From one month to at least 7 years [18], 2.2 mm to 1.7 cm SL [17, 20]. Habitat: 

Benthic [1], shallow, sometimes intertidal, waters over soft seafloors (often among eelgrass) [14, 15, 21]. Remain 

on mixed sand-mud bottoms for several years [5, 22]. At 3–5 years, fish begin to migrate into deeper waters and 

within a few years occupy same depth and habitat as adults [17]. 

Adults—Age and size at first maturity: Age varies with location, bottom depth, and year [23]; size varies with 

bottom depth [23]. Studies in the eastern Bering Sea showed female length at 50 percent maturity varied between 

27 and 34 cm TL and about 10–13 years [23]. Other studies have yielded a similar range of length (as small 

as 25 cm) and ages of 7–11 years [5, 11]. Males mature when smaller and younger, between 10.5 and 20.3 cm 

and perhaps 4 years younger than females [5, 17]. From about 10 years and older, females are larger at age than 

males. Growth rates vary with area. For instance, fish living in northwestern (and colder) parts of eastern Bering 

Sea are larger at age than those living more to the southeast [23]. Length-weight relations are similar between 

sexes. Maximum age: 39 years [24]. Males and females may have similar life spans [25]. Maximum size: 49 cm 

TL [1]. Habitat: Benthic [1, 2]. 

Substrate—Mud, sand, and mixed soft sediments [8, 14, 19, 26]. 

Physical/chemical—Temperature: -1.5–13 °C, mostly 0–11 °C [7, 16, 17, 27]. In southeastern Bering Sea, from 

-1.7 to 11.7 °C (mainly 1.8–4.9 °C) [28]. Salinity: Marine and estuarine, to at least 16 parts per thousand [8, 

14, 29]. 

Behavior 

Diel—Unknown. Flatfish generally tend to bury themselves in bottom sediments to escape predators, reduce 

their metabolic rate to resting phase, and to avoid fast currents [30].

Seasonal—Extensive seasonal migrations of 402–482 km or more in eastern Bering Sea. In winter, they occupy 

three major grounds on the lower continental shelf-upper slope region. In spring (as early as March through 

May), they migrate nearshore to spawn, then into slightly deeper waters to feed. The spring migration is thought 

to follow the ice edge as it recedes [17]. In autumn, adults move offshore to wintering grounds [5, 25, 29, 31]. 
572 Alaska Arctic Marine Fish Ecology Catalog

Reproductive—Spawn in nearshore waters [5]. May rise to surface waters to spawn [19]. Females are batch 

spawners and an individual female will spawn 8–11 batches per egg series with some females spawning more 

than 1 series per season [32]. 

Schooling—Occurs in very dense concentrations during winter [31]. 

Feeding—An opportunistic, benthopelagic feeder [5]. Generally feeds on or in bottom, but may ascend to 

surface waters to feed on midwater animals [19, 33]. Few fish feed during winter. Feeding intensity increases 

before and during spring migration and particularly before spawning [5, 19, 34, 35].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [5]. 

Spawning season—May through September in eastern Bering Sea and May through November in Gulf of 

Alaska. During warmer-water years, spawning may occur earlier in the season [25, 36–38]. 

Fecundity—Batch fecundity ranges from 2,400 to 408,000 eggs and total fecundity for a series ranges from 

295,615 to 3,635,108 [32]. 

Food and Feeding

Food items—Young fish prey on small crustaceans (gammarid amphipods, harpacticoid copepods, and 

ostracods) and polychaetes [14, 39]. Older fish consume mostly benthic invertebrates (for example, polychaetes, 

clams, brittle stars, echiurioids, shrimps, crabs, sand dollars, sea cucumbers) and some fishes; diets vary 

depending on habitat type and geographic location [19, 34, 40, 41]. 

Trophic level—3.6 [42].

Biological Interactions

Predators—In U.S Chukchi Sea, off Point Hope, ringed seals ate small numbers in April [43]. In eastern Bering 

Sea and Gulf of Alaska, Bigmouth Sculpin, Great and Plain sculpins, Pacific Cod, Pacific Halibut, Skates, and 

Steller sea lions [41, 44–46].

Competitors—Likely a wide range of benthic feeding fishes, including eelpouts, Walleye Pollock, sculpins, and 

other flatfishes.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (K=0.1–0.15; t

m=4–10; tmax=26; Fecundity=1 million) 

[47].

Traditional and Cultural Importance

Yellowfin Sole are of little importance in subsistence fisheries in Arctic waters.

Commercial Fisheries

Currently, Yellowfin Sole are not commercially harvested.
Yellowfin Sole 573

Potential Effects of Climate Change

As a boreal Pacific fish common throughout much of the Bering Sea and only fairly common to rare in the U.S. 

Chukchi and Beaufort Seas, Yellowfin Sole would be expected to increase in abundance in the study area as 

climate warms. 

Areas for Future Research [A]

Yellowfin Sole are of commercial interest in the southeastern Bering Sea. As a potential indicator species, 

long-term monitoring programs should be designed to understand trends in population growth, survival, and 

recruitment. The location of spawning grounds and overwintering areas should be described along with seasonal 

information about habitat usage including trophic interactions.

Remarks

Allozyme analyses have shown some population differences between fish living in the Bering Sea and the Gulf of Alaska [48]. 

References Cited 

Bakkala, R.G., 1981, Population characteristics and ecology of yellowfin sole, in Hood, D.W., and Calder, J.A., eds.: The eastern 

Bering Sea shelf—Oceanography and resources: National Oceanic and Atmospheric Administration, p. 553–574. [17]

Fadeev, N.S., 1970, The fishery and biological characteristics of yellowfin soles in the eastern part of the Bering Sea in

Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part V: Jerusalem, Israel Program for Scientific 

Translations, p. 332–396. [19]

McGregor, S.B., 1990, Seasonal and ontogenetic changes in meiofauna in the diets of postmetamorphic flatfishes: University of 

Alaska, Fairbanks, Master’s thesis. [14]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Wilderbuer, T.K., Walters, G.E., and Bakkala, R.G., 1992, Yellowfin sole, Pleuronectes asper, of the eastern Bering Sea—

Biological characteristics, history of exploitation, and management: Marine Fisheries Review, v. 54, no. 4, p. 1–18. [5]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

5. Wilderbuer, T.K., Walters, G.E., and Bakkala, R.G., 1992, Yellowfin sole, Pleuronectes asper, of the eastern Bering Sea—

Biological characteristics, history of exploitation, and management: Marine Fisheries Review, v. 54, no. 4, p. 1–18.
574 Alaska Arctic Marine Fish Ecology Catalog

6. Martin, M.H., and Clausen, D.M., 1995, Data report—1993 Gulf of Alaska Bottom Trawl Survey: U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration Technical Memorandum, NMFS-AFSC-59, 217 p.

7. Kupriyanov, S.V., 1996, Distribution and biological indices of yellowfin sole (Pleuronectes asper) in the southwestern 

Bering Sea, in Mathisen, O.A., and Coyle, K.O., eds., Ecology of the Bering Sea—A review of Russian literature: 

Fairbanks, University of Alaska, Alaska Sea Grant Report no. 96-01, 306 p.

8. Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, Nearshore fish distributions in an Alaska estuary in relation to 

stratification, temperature and salinity: Estuarine, Coastal and Shelf Science, v. 51, no. 1, p. 45–59.

9. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

10. Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., Shestakov, A.V., Yabe, M., and others, 2001, Annotated list of 

fish-like vertebrates and fish in Tauisk Bay (northern part of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, H.H., 

eds., Biodiversity and ecological status along the northern coast of the Sea of Okhotsk—A collection of study reports: 

Dalnauka Vladivostok, Russia, Institute of Biological Problems of the North, p. 64–86.

11. Nichol, D.G., 1995, Spawning and maturation of female yellowfin sole in the eastern Bering Sea, in Alaska Sea Grant 

Program, Proceedings of the International Symposium on North Pacific Flatfish: Alaska Sea Grant Program, AK-SG-95-04, 

p. 35–50.

12. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

13. Sokolovskii, A., and Sokolovskaya, T., 2009, Distribution of ichthyoplankton in the eastern Peter the Great Bay, Sea of 

Japan, in June and July of 2007: Russian Journal of Marine Biology, v. 35, no. 3, p. 250–254.

14. McGregor, S.B., 1990, Seasonal and ontogenetic changes in meiofauna in the diets of postmetamorphic flatfishes: 

Fairbanks, University of Alaska, Master’s thesis.

15. Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries Science Center, Technical Memorandum NMFS-AFSC-157, 

98 p.

16. Salveson, S.J., and Alton, M.S., 1976, Yellowfin sole (family Pleuronectidae) in Pereya, W.T., Reeves, J.E., and Bakkala, 

R.G., eds.: Demersal fish and shellfish resources of the Eastern Bering Sea in the baseline year 1975: Seattle, Washington, 

National Marine Fisheries Service, Northwest and Alaska Fisheries Center, Processed Report, p. 439–459.

17. Bakkala, R.G., 1981, Population characteristics and ecology of yellowfin sole, in Hood, D.W., and Calder, J.A., eds.: The 

eastern Bering Sea shelf—Oceanography and resources: National Oceanic and Atmospheric Administration, p. 553–574.

18. Tarasyuk, S.N., 2002, Survival of yellowfin sole (Limanda aspera Pallas) in the northern part of the Tatar Strait (Sea of 

Japan) during the second half of the 20th century, in Batchelder, H.P., and others, eds., PICES-Globe International Program 

on Climate Change and Carrying Capacity—Report of 2110 BASS/MODEL, MONITOR and REX workshops, and The 

2002 MODEL/REX workshop: Sidney, British Columbia, North Pacific Marine Science Organization (PICES).

19. Fadeev, N.S., 1970, The fishery and biological characteristics of yellowfin soles in the eastern part of the Bering Sea in

Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part V: Jerusalem, Israel Program for Scientific 

Translations, p. 332–396.

20. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

21. McConnaughey, T., and McRoy, C.P., 1979, 13C label identifies eelgrass (Zostera marina) carbon in an Alaskan estuarine 

food web: Marine Biology, v. 53, no. 3, p. 263–269.

22. Norcross, B.L., Blanchard, A., and Holladay, B.A., 1999, Comparison of models for defining nearshore flatfish nursery areas 

in Alaska waters: Fisheries Oceanography, v. 8, p. 50–67.
Yellowfin Sole 575

23. Nichol, D.G., 1997, Effects of geography and bathymetry on growth and maturity of yellowfin sole, Pleuronectes asper, in 

the eastern Bering Sea: Fishery Bulletin, v. 95, no. 3, p. 494–503.

24. National Oceanic and Atmospheric Administration, 2010, Age and growth program: National Oceanic and Atmospheric 

Administration, accessed April 2011 at http://www.afsc.noaa.gov/refm/age/Stats/max_age.htm.

25. Nichol, D.G., 1998, Annual and between-sex variability of yellowfin sole, Pleuronectes asper, spring-summer distributions 

in the eastern Bering Sea: Fishery Bulletin, v. 96, no. 3, p. 547–561.

26. Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic marine fish museum specimens, 2nd ed., Metadata report and 

database submitted to ArcOD, Institute of Marine Science: University of Alaska, Fairbanks, by Point Stephens Research, 

metadata report accessed August 8, 2012, at http://www.arcodiv.org/Database/Fish_datasets.html.

27. Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution in the Kronitsky Bay in relation to abiotic factors: Zoologicheskii 

Zhurnal, v. 63, p. 1,363–1,373.

28. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

29. Fadeev, N.S., 1965, Comparative outline of the biology of flatfishes in the southeastern part of the Bering Sea and condition 

of their resources, in Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part IV: Jerusalem, Israel 

Program for Scientific Translations, p. 112–146.

30. Moles, A., and Norcross, B.L., 1995, Sediment preference in juvenile Pacific flatfishes: Netherlands Journal of Sea 

Research, v. 34, no. 1–3, p. 177–182.

31. Moiseev, P.A., 1964, Some results of the work of the Bering Sea expedition, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific, Part III: Jeruselam, Israel Program for Scientific Translations, p. 1–21.

32. Nichol, D.G., and Acuna, E.I., 2001, Annual and batch fecundities of yellowfin sole, Limanda aspera, in the eastern Bering 

Sea: Fishery Bulletin, v. 99, no. 1, p. 108–122.

33. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna in the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from the U.S. Department of Commerce, Springfield, Virginia.]

34. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

35. Kim, S.T., and Shepeleva, O.N., 2001, The structure of shelf ichthyocenoses of northeastern Sakhalin and Terpeniya Bay: 

Journal of Ichthyology, v. 41, no. 9, p. 711–722.

36. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224.

37. Blackburn, J.E., and Jackson, P.B., 1982, Seasonal composition and abundance of juvenile and adult marine finfish and crab 

species in the nearshore zone of Kodiak Island’s eastside during April 1978 through March 1979: Outer Continental Shelf 

Environmental Assessment Program, Alaska Department of Fish and Game, Final Report, Research Unit 552, p. 377–570.

38. North Pacific Fishery Management Council, 2010, Fishery management plan for groundfish of the Bering Sea and Aleutian 

Islands management area—Appendices—Fishery management plan for groundfish of the BSAI Management Area: North 

Pacific Fisheries Management Council, 233 p.

39. Holladay, B.A., and Norcross, B.L., 1995, Diet diversity as a mechanism for partitioning nursery grounds of pleuronectids: 

Proceedings of the International Symposium on North Pacific Flatfish, Alaska Sea Grant AK-SG-95-04, p. 177–203.

40. Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, Fisheries Research Institute, FRI-UW-7719.

41. Lang, G.M., Livingston, P.A., Pacunski, R.E., Parkhurst, J., and Yang, M.-S., 1991, Groundfish food habits and predation on 

commercially important prey species in the eastern Bering Sea from 1984 to 1986: Seattle, Washington, U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum NMFS F/NWC-207, 240 p.
576 Alaska Arctic Marine Fish Ecology Catalog

42. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.

43. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.

44. Brodeur, R.D., and Livingston, P.A., 1988, Food habits and diet overlap of various Eastern Bering Sea fishes: National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS F/NWC-127, 76 p.

45. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990. 

46. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, NOAA Technical Memorandum NMFS-AFSC-158, 249 p.

47. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

48. Grant, W.S., Bakkala, R.G., Utter, F.M., Teel, D.J., and Kobayashi, T., 1983, Biochemical genetic population structure of 

yellowfin sole, Limanda aspera, of the North Pacific Ocean and Bering Sea: Fishery Bulletin, v. 81, no. 4, p. 667–677.
Longhead Dab 577

Longhead Dab (Limanda proboscidea) 

Gilbert, 1896

Family Pleuronectidae 

Colloquial Name: Iñupiaq—Nataagnaq [1].

Ecological Role: Due to its relative scarcity in Arctic waters, the 

Longhead Dab is likely to be of little ecological importance in the 

U.S. Chukchi and Beaufort Seas.

Physical Description/Attributes: Member of the right eyed 

flounders. Grayish brown with small whitish spots on the eyed 

side and lemon yellow on blind side. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 

2002, p. 843) [2]. Swim bladder: Absent [2]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi and Beaufort Seas [3]. Eastward in Canadian 

Arctic to Bathurst Inlet and westward to western Chukchi Sea [3, 4]. Elsewhere in Alaska, eastern Bering Sea north of Unimak 

Island. Worldwide, off Hokkaido, Japan, through Sea of Okhotsk [2, 4]. 

Relative Abundance: Uncommon in U.S. Chukchi Sea and rare in U.S. Beaufort Sea [3, 7, 8]. Rare in Canadian Arctic [3]. 

Common from Sea of Japan and along eastern Kamchatka Peninsula, Russia, to eastern Bering Sea at least as far north as Bristol 

Bay [9–11]. Populations in southeastern Bering Sea appear to be expanding their distribution northward [12].

Longhead Dab (Limanda proboscidea), 170 mm, U.S. Chukchi 

Sea, 2007. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

Geographic distribution of Longhead Dab (Limanda proboscidea), within Arctic Outer Continental Shelf Planning 

Areas [5] based on review of published literature and specimens from historical and recent collections [3, 6].

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Longhead Dab

Limanda proboscidea

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
578 Alaska Arctic Marine Fish Ecology Catalog

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

Overall benthic depth range.

Specific depth range of either

juveniles or adults is unknown 

Limanda proboscidea

Longhead Dab

Benthic distribution

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

One larval fish taken in U.S. Chukchi Sea

between 0-29 m

Reproductive distribution

Juveniles and adults

Overall depth data from outside U.S. Beaufort-Chukchi Seas Spawning data from outside U.S. Beaufort-Chukchi Seas

Eggs are pelagic 

but depths unknown

Depths documented in U.S. Beaufort-Chukchi Seas Depth range in U.S. Chukchi Sea

Spawning

Larvae

Benthic and reproductive distribution of Longhead Dab (Limanda proboscidea). Larval distribution reflects one record from 

the study area.

Habitats and Life History

Eggs—Size: 0.72–0.87 mm [17]. Time to hatching: Unknown. Habitat: Pelagic [18]. 

Larvae—Size at hatching: Less than 4.8 mm [17]. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Pelagic [18]. 

Juveniles—Age and size: Unknown. Habitat: Benthic, on soft sea floors [9]. 

Adults—Age and size at first maturity: Unknown. Maximum age: Unknown. Maximum size: 41 cm TL [2]. 

Habitat: Nearshore, benthic [9]. 

Substrate—Sand and mud [18]. 

Physical/chemical—Temperature: -1.8–11.7 °C [6, 19]. Salinity: Marine [9].

Behavior

Diel—Unknown. 

Seasonal—In summer relatively common in nearshore waters of Bristol Bay and absent from those waters in 

winter [9]. 

Reproductive—Spawn nearshore [16]. 

Schooling—Often with starry flounder [9]. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [20]. 

Spawning season—Between May and September [9, 16, 21]. 

Fecundity—78,700–841,200 eggs per season [9].

Depth Range: Benthic individuals taken in U.S. Beaufort Sea at 5–8 m [13] and U.S. Chukchi Sea at 17 m [7]. Pelagic larvae 

taken in U.S. Chukchi Sea at 29 m or less [14]. Generally, 5–125 m or perhaps to 160 m [6], almost always less than 100 m [2, 

13, 15]. Spawn nearshore [16].
Longhead Dab 579

Food and Feeding

Food items—Polychaetes, crustaceans (for example, copepods, cumaceans, gammarid amphipods, and mysids) 

and clams [22, 23]. 

Trophic level—3.6 [12].

Biological Interactions

Predators—Pacific Cod, Pacific Halibut, Plain Sculpin, and skates [23–25]. 

Competitors—Other relatively small benthic feeders, such as eelpouts and sculpins.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (preliminary K or fecundity) [26].

Traditional and Cultural Importance 

Unknown, but of suspected limited use.

Commercial Fisheries

Currently, Longhead Dab are not commercially harvested [27].

Potential Effects of Climate Change

As Longhead Dab are common in part of the Bering Sea and those populations appear to be expanding their 

distribution northward [12], an increase in abundance in both the U.S. Chukchi and Beaufort Seas would be 

expected as the climate warms. 

Areas for Future Research [B]

Changes in abundance patterns are an expected outcome of global warming. Reliable baseline information and 

regular monitoring are needed to evaluate northerly shifts in this population. The monitoring should include 

estimation of key vital statistics in the population (growth, survival, and recruitment), especially in the 

Chukchi Sea.
580 Alaska Arctic Marine Fish Ecology Catalog

References Cited 

Fadeev, N.S., 1965, Comparative outline of the biology of flatfishes in the southeastern part of the Bering Sea and condition 

of their resources, in Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part IV: Jerusalem, Israel 

Program for Scientific Translations, p. 112–146. [9]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320. [12]

Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries investigations 

in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224. [16]

Bibliography

1. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, Russian–

American long-term census of the Arctic—Benthic fishes trawled in the Chukchi Sea and Bering Strait, August 2004: 

Northwestern Naturalist, v. 88, no. 3, p. 168–187.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

8. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2 p. 195–209.

9. Fadeev, N.S., 1965, Comparative outline of the biology of flatfishes in the southeastern part of the Bering Sea and condition 

of their resources, in Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part IV: Jerusalem, Israel 

Program for Scientific Translations, p. 112–146.

10. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

11. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

12. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320. 
Longhead Dab 581

13. Johnson, S.W., Thedinga, J.F., Neff, A.D., and Hoffman, C.A., 2010, Fish fauna in nearshore waters of a Barrier Island 

in the western Beaufort Sea, Alaska: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, 

Technical Memorandum NMFSAFSC-210, 28 p.

14. Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 2009, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

15. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

16. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224.

17. Matarese, A.C., Kendall, J., Arthur W., Blood, D.M., and Vinter, B.M., 1989, Laboratory guide to early life history stages 

of northeast Pacific fishes: U.S. Department of Commerce, National Oceanic and Atmospheric Administration, National 

Marine Fisheries Service Technical Report NMFS 80, 652 p.

18. Mecklenburg, C., and Mecklenburg, T.A., 2011, Longhead Dab—Limanda proboscidea Gilbert, 1896: Arctic Ocean 

Diversity—Census of Marine Life, accessed April 2011 at http://www.arcodiv.org/Fish/Limanda_proboscidea.html. 

19. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

20. Wilderbuer, T.K., Walters, G.E., and Bakkala, R.G., 1992, Yellowfin sole, Pleuronectes asper, of the eastern Bering Sea—

Biological characteristics, history of exploitation, and management: Marine Fisheries Review, v. 54, no. 4, p. 1–18.

21. Grigorev, S.S., and Fadeev, N.S., 1995, Distribution of Alaska plaice, flathead sole, and Bering flounder eggs in the eastern 

Bering Sea during April–July, 1988–1990, in Alaska Sea Grant Program, eds., Proceeding of the International Symposium 

on North Pacific Flatfish: Fairbanks, University of Alaska, Alaska Sea Grant Program, AK-SG-95-04.

22. Tyler, A.V., 1972, Food resource division among northern, marine, demersal fishes: Journal of the Fisheries Research Board 

of Canada, v. 29, no. 7, p. 997–1,003.

23. Brodeur, R.D., and Livingston, P.A., 1988, Food habits and diet overlap of various Eastern Bering Sea fishes: U.S. 

Department of Commerce, National Oceanic and Atmospheric Administration Technical Memorandum NMFS F/NWC-127, 

76 p.

24. Lang, G.M., Livingston, P.A., Pacunski, R.E., Parkhurst, J., and Yang, M.-S., 1991, Groundfish food habits and predation on 

commercially important prey species in the eastern Bering Sea from 1984 to 1986: U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS F/NWC-207, 240 p.

25. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration Technical Memorandum NMFS-AFSC-158, 249 p.

26. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

27. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna in the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from the U.S. Department of Commerce, Springfield, Virginia.]
582 Alaska Arctic Marine Fish Ecology Catalog

Sakhalin Sole (Limanda sakhalinensis) 

Hubbs, 1915

Family Pleuronectidae 

Colloquial Name: None in U.S. Chukchi and Beaufort Seas.

Ecological Role: Likely of little ecological significance in Chukchi 

Sea marine food webs.

Physical Description/Attributes: Member of the right eyed 

flounders. Slightly elongate oval body. Eyed side uniformly brown 

or with vague dark blotches, blind side off-white. For specific 

diagnostic characteristics, see Fishes of Alaska (Mecklenburg and 

others, 2002, p. 845) [1]. Swim bladder: Absent [1]. Antifreeze 

glycoproteins in blood serum: Unknown.

Range: U.S. Chukchi Sea as far northward as 71°04′N, 158°26′W [2]. Elsewhere in Alaska, to southeastern Bering Sea. 

Worldwide, from Sea of Okhotsk and northern Sea of Japan (Tatar Strait) to Gulf of Anadyr, Russia, and St. Lawrence Island, 

Alaska [1].

Relative Abundance: Uncommon in U.S. Chukchi Sea and northern Bering Sea [4].

Sakhalin Sole (Limanda sakhalinensis), 123 mm, Chukchi 

Sea, 2009. Photograph by C.W. Mecklenburg, Point Stephens 

Research.

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Sakhalin Sole

Limanda sakhalinensis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Sakhalin Sole (Limanda sakhalinensis), within Alaskan Outer Continental Shelf 

Planning Areas [3], based on review of published literature and specimens from historical and recent 

collections [1, 2, 4]. 
Sakhalin Sole 583

tac1452223119gSakhalinSole Depth, in meters

500

400

300

200

100

0

Depths from U.S. Chukchi Sea

Some fish to 360 m

Limanda sakhalinensis

Sakhalin Sole

Benthic distribution

Overall depth data from 

outside U.S. Beaufort-Chukchi Seas 500

400

300

200

100

0

No data for this species

Depth, in meters

Spawning

Reproductive distribution

Potential location

 of eggs and larvae.

 Specific depth unknown

Depth range of spawning is uncertain. 

Potential location is based on 

depth range of benthic individuals

Overall benthic depth range. 

Specific depth range of either 

juveniles or adults is unknown

Juveniles and adults Spawning

Open ocean Shore Open ocean Shore

Benthic and reproductive distribution of Sakhalin Sole (Limanda sakhalinensis).

Habitats and Life History

Eggs—Size: Unknown. Time to hatching: Unknown. Habitat: Likely pelagic, as are most other righteye 

flounders [7].

Larvae—Size at hatching: Unknown. Size at juvenile transformation: Unknown. Days to juvenile 

transformation: Unknown. Habitat: Likely pelagic, as are most other righteye flounders [7]. 

Juveniles—Age and size: Unknown. Habitat: Benthic [1, 4]. 

Adults—Age and size at first maturity: Unknown. Maximum age: 8 years [8]. Maximum size: 35 cm TL [1]. 

Habitat: Benthic [1, 4]. 

Substrate—Soft bottoms [1]. 

Physical/chemical—Temperature: -1.7–9.3 °C [2]. Salinity: Marine. 

Behavior

Diel—Unknown. 

Seasonal—Unknown. 

Reproductive—Unknown. 

Schooling—Unknown. 

Feeding—Unknown. 

Populations or Stocks

There have been no studies.

Reproduction

Mode—Oviparous [7]. 

Spawning season—Unknown. 

Fecundity—Unknown. 

Depth Range: Overall 10–360 m [5], mainly less than 110 m [6]. At depths of 20–51 m in the U.S. Chukchi Sea and northern 

Bering Sea [2], to 95 m in southeastern Bering Sea [1].
584 Alaska Arctic Marine Fish Ecology Catalog

Food and Feeding

Food items—Amphipods, polychaetes, and euphausiids [8]. 

Trophic level—3.68 (standard error 0.55) [8].

Biological Interactions

Predators—Pacific Cod and Great Sculpin [9, 10]. 

Competitors—Presumably a wide range of other zoobenthos feeders such as Arctic Cod, Walleye Pollock, other 

flounders, eelpouts, and sculpins.

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (preliminary K or fecundity) [8].

Traditional and Cultural Importance

None reported. 

Commercial Fisheries

Sakhalin Sole are not commercially harvested currently.

Potential Effects of Climate Change

Warming conditions would likely lead to an increased expansion of this species’ distribution in the U.S. Chukchi 

Sea and possibly into the Beaufort Sea.

Areas for Future Research [B]

Little is known about the ecology and life history of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment; 

(3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic 

movements; (6) population studies; (7) prey; and (8) predators.
Sakhalin Sole 585

References Cited

Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p. [7]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

3. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Sheiko, B.A., and Fedorov, V.V., 2000, Class cephalaspidomorphi-lampreys, Class chondrichthyes-cartilaginous fishes, 

Class holocephali-chimaeras, Class osteichthyes-bony fishes, in Artuykhin, Yu. B, and Sheyko, B.A., eds., Catalog of 

vertebrates of Kamchatka and adjacent waters: Russia, Rossiyskaya Akademiya Nauk, p. 7–69. [In Russian.]

6. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: Seattle, Washington, U.S. Department of the Interior, U.S. Geological Survey, Biological 

Resources Division, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

7. Love, M.S., 2011, Certainly more than you wanted to know about the fishes of the Pacific Coast: Santa Barbara, California, 

Really Big Press, 649 p.

8. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

9. Tokranov, A.M., 1991, Diet of the Pacific cod, Gadus morhua macrocephalus, and its position in the food chain in 

Kamchatkan coastal waters: Journal of Ichthyology, v. 31, no. 5, p. 84–98.

10. Napazakov, V.V., and Chuchukalo, V.I., 2003, Diet and trophic status of mass species of sculpins (Cottidae) in the western 

part of the Bering Sea in autumn: Journal of Ichthyology, v. 43, no. 3, p. 236–244.
586 Alaska Arctic Marine Fish Ecology Catalog

Arctic Flounder (Liopsetta glacialis) 

(Pallas, 1776)

Family Pleuronectidae 

Colloquial Name: Iñupiaq—Nataagnaq, Puyyaqiaq [1].

Ecological Role: Unknown. Despite being common in nearshore 

Arctic waters, little data are available on its role in the food chain. 

Physical Description/Attributes: Member of the right eyed 

flounders. Eyed side is darkish brown or black. Blind side is chalky 

white to lime green. Fins are paler. Dorsal and anal fins usually 

spotted. For specific diagnostic characteristics, see Fishes of Alaska

(Mecklenburg and others, 2002, p. 836) [2]. Swim bladder: Absent 

[2] Antifreeze glycoproteins in blood serum: Unknown. 

Range: U.S. Chukchi and Beaufort Seas [3]. Elsewhere in Alaska, 

to southeastern Bering Sea and Aleutian Islands. Worldwide, from 

Arctic Russia to Labrador, Canada, and in the Atlantic and Okhotsk Sea in western Pacific [2]. 

Relative Abundance: Common near shore in U.S. Chukchi and Beaufort Seas [6]. Common from at least Bristol Bay [7] to 

Canadian Arctic at least to Tuktoyaktuk Harbor [8, 9], but appears to be uncommon east of Cape Bathurst [10].

Arctic Flounder (Liopsetta glacialis), from Love (2011, 

p. 588). Photograph attributed to National Oceanic and 

Atmospheric Administration Fisheries.

Geographic distribution of Arctic Flounder (Liopsetta glacialis), within Arctic Outer Continental Shelf Planning 

Areas [4] based on review of published literature and specimens from historical and recent collections [3, 5]. 

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Arctic Flounder

Liopsetta glacialis

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA
Arctic Flounder 587

Depth Range: Nearshore, barely subtidal to 91 m, rarely deeper than about 10 m in U.S. Chukchi Sea [11]. Spawns between 5 

and 10 m [12].

Benthic and reproductive distribution of Arctic Flounder (Liopsetta glacialis).

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

Liopsetta glacialis

Arctic Flounder

Benthic distribution

Open ocean Shore

Depth, in meters

125

100

75

50

25

0

Reproductive distribution

Adults

Juveniles

Data from U.S. Chukchi Sea Spawning data from outside U.S. Beaufort-Chukchi Seas

Eggs and larvae are pelagic but specific depths unknown

Spawning

Some fish

to 91 m

Habitats and Life History

Eggs—Size: 1.5–1.7 mm [13, 14]. Time to hatching: 22–42 days at a mean temperature of 1.0 °C (range–0.5 to 

3.1 °C) [13]. Habitat: Pelagic [13, 14]. 

Larvae—Size at hatching: 5.6 mm [13]. Size at juvenile transformation: 14–20 mm TL [8, 10, 15]. Days to 

juvenile transformation: Unknown. Habitat: Pelagic [16]. 

Juveniles—Age and size: 1.4–20.0 cm TL [10, 11]. Habitat: Benthic, soft sea floors in nearshore protected 

waters [8, 10, 15]. The smallest fish tend to live in quiet sloughs and other backwaters [17]. 

Adults—Age and size at first maturity: Generally, most fish mature at between 4 and 8 years and 13.0 and 

20.0 cm TL [9, 11, 18]. Males tend to mature at a slightly younger age and smaller size than females [18–20]. 

Females grow larger, live longer, and may grow faster than males, although growth rates vary widely with region. 

Maximum age: At least 28 years [21]). Maximum size: 44 cm (17.6 in) TL [20]. Habitat: Benthic, nearshore [7, 

22, 23]. 

Substrate—Mud and fine sand [7]. 

Physical/chemical—Temperature: Less than –1.0–13.5 °C [7, 23]. Salinity: 0–31 parts per thousand. Tolerant of 

estuarine conditions [7, 16, 23]. Occasionally ascends into fresh water [22]. 

Behavior

Diel—Relatively sedentary, few move more than a few kilometers in a year [21], although one tagged fish in the 

ANWR moved 62 km before being recaptured [24]. Moves closer to shore in evening and mainly on incoming 

tides (Morrow 1980). 

Seasonal—Slight offshore movements in autumn; however, extent of this migration is unknown in U.S. Beaufort 

and Chukchi Seas [14, 23]. Over winters in the deeper parts of Tuktoyatuk Harbor, at depths of 12 m or more 

[25]. In spring, as temperatures warm, large numbers move into shallow, nearshore waters where they remain 

until fall [25]. 

Reproductive—Spawning occurs under ice in winter and spring in nearshore waters, although location of 

spawning aggregations is poorly understood [25, 26]. Mature fish may spawn only once every 2 years [11]. 

Schooling—At certain times, single-sex aggregations are formed [9]. 

Feeding—Unknown. 

Populations or Stocks

Life history parameters have been described from the northeastern Chukchi Sea and the coastal Beaufort Sea. No 

information about population size or stock structure is available.
588 Alaska Arctic Marine Fish Ecology Catalog

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [27]. 

Spawning season—At least March through June in Tuktoyaktuk, Canada, region [9, 28]. From December 

through March in White and Kara Seas [13, 14]. 

Fecundity—50,000–200,000 pelagic eggs [13, 14].

Food and Feeding

Food items—Primarily epibenthic and benthic invertebrates, such as clams, isopods, amphipods, ascidians, and 

mysids [11, 18, 29]. 

Trophic level—3.6 [27].

Biological Interactions

Predators—Unknown. 

Competitors—Unknown, but presumably other benthic feeding predators, such as other flatfishes, as well as 

eelpouts and sculpins.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (t

m=2; Fecundity =31,000–230,000).

Traditional and Cultural Importance

Arctic Flounder form a minor part of the subsistence fisheries of the U.S. Beaufort and Chukchi Seas. It has 

been listed as a food fish from Kotzebue Sound and northward [30]. Another reported “flounder” caught in the 

nearshore waters at Point Hope and Point Lay is likely this species [31]. In the Canadian Beaufort Sea, small 

numbers were reported in subsistence catches along the Yukon coast [32] and some were reportedly fed to dogs 

in the Canadian Arctic [10].

Commercial Fisheries

Currently, Arctic Flounder are not commercially harvested.

Potential Effects of Climate Change

Because Arctic Flounder are predominantly an Arctic species, it would be expected that climate warming would 

push this species northward. 
Arctic Flounder 589

Areas for Future Research [A]

Even though the species is relatively common, especially in coastal waters of the Alaska Beaufort Sea, compared 

to the amphidromous species, little is known about its biology and ecology in the U.S. Chukchi and Beaufort 

Seas. Its distribution and abundance, and the amount of existing information, make it a potential indicator of 

change associated with climate warming. The vulnerability of this species to climate changes in light of possible 

increases in abundance of other flatfish and associated interactions should be assessed. The Arctic Flounder is 

 considered a potential indicator for monitoring effects of climate change.

References Cited 

Aronovich, T.M., Doroshev, S.I., Spectorova, L.V., and Makhotin, V.M., 1975, Egg incubation and larval rearing of navaga 

(Eleginus navaga Pall.), Arctic Cod (Boreogadus saida lepechin) and Arctic flounder (Liopsetta glacialis Pall.) in the 

laboratory: Aquaculture, v. 6, no. 3, p. 233–242. [13]

Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, Simpson 

Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649. [23]

Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research Associates, 

Inc., North Slope Borough, Materials Source Division, 105 p. [18]

Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p. [11]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Bibliography

1. Craig, P.C., 1989b, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers University 

of Alaska, no. 24, p. 27–54.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. George, C., Moulton, L.L., and Johnson, M., 2007, A field guide to the common fishes of the North Slope of Alaska: Alaska 

Department of Wildlife Management, North Slope Borough, 93 p.

7. Isakson, J.S., Houghton, J.P., Rogers, D.E., and Parker, S.S., 1986, Fish use of inshore habitats north of the Alaska 

Peninsula, June–September 1984 and June–July 1985—Final report: U.S. Department of the Interior, Minerals Management 

Service, Alaska OCS Region, OCS EIS/EA, 380 p.

8. Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 1989, Fish population characteristics of Arctic National Wildlife 

Refuge coastal waters, summer 1988: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Fishery Assistance office, Progress 

Report, 73 p.

9. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.
590 Alaska Arctic Marine Fish Ecology Catalog

10. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

11. Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, B.J., 1984, Fish distribution and use of nearshore waters in the 

northeastern Chukchi Sea: LGL Ecological Research Associates, Inc., Outer Continental Shelf Environmental Assessment 

Program, National Oceanic and Atmospheric Administration, OMPA/OCSEAP, Final Report, 190 p.

12. Morrow, J.E., 1980, The freshwater fishes of Alaska: Anchorage, Alaska Northwest Publishing Company, 248 p.

13. Aronovich, T.M., Doroshev, S.I., Spectorova, L.V., and Makhotin, V.M., 1975, Egg incubation and larval rearing of navaga 

(Eleginus navaga Pall.), Arctic Cod (Boreogadus saida lepechin) and Arctic flounder (Liopsetta glacialis Pall.) in the 

laboratory: Aquaculture, v. 6, no. 3, p. 233–242.

14. Andriashev, A.P., 1954, Fishes of the northern seas of the U.S.S.R.—Keys to the fauna in the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute no. 53, 566 p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from the U.S. Department of Commerce, Springfield, Virginia.]

15. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

16. Bond, W.A., and Erickson, R.N., 1989, Summer studies of the nearshore fish community at Phillips Bay, Beaufort Sea coast, 

Yukon: Winnepeg, Manitoba, Canadian Technical Report of Fisheries and Aquatic Sciences, Central and Arctic Region, 

Department of Fisheries and Oceans, no. 1676, 102 p.

17. Martin, D.J., Glass, D.R., Whitmus, C.J., Simenstad, C.A., Milward, D.A., Volk, E.C., Stevenson, M.L., Nunes, P., Savoie, 

M., and Grotefendt, R.A., 1986, Distribution, seasonal abundance, and feeding dependencies of juvenile salmon and 

non-salmonid fishes in the Yukon River Delta: Outer Continental Shelf Environmental Assessment Program, Reports of 

Principal Investigators, U.S. Department of Commerce and U.S. Department of the Interior, 388 p.

18. Craig, P.C., and Schmidt, D.R., 1985, Fish resources at Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source Division, 105 p.

19. Jones, M.L., and Den Beste, J., 1977, Tuft Point and adjacent coastal areas fisheries projects: Calgary, Alberta, Canada, 

Aquatic Environments, Ltd., 152 p.

20. Kobelev, E.A., 1989, Some biological characteristics of the Arctic flounder, Liopsetta glacialis, in the southeastern area of 

the Barents Sea: Journal of Ichthyology, v. 29, no. 6, p. 16–21.

21. Palmer, D.E., and Dugan, L.J., 1990, Fish population characteristics of Arctic National Wildlife Refuge coastal waters, 

summer 1989: Fairbanks, Alaska, U.S. Fish and Wildlife Service, Progress Report, 83 p.

22. West, R.L., and Fruge, D.J., 1989, A review of coastal plain fish surveys and the results of 1986 fish surveys of selected 

coastal lakes and streams Arctic National Wildlife Refuge, Alaska: Alaska Fisheries Technical Report, no. 4, 22 p.

23. Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce, Biological Studies, p. 384–649.

24. Underwood, T.J., Gordon, J.A., Millard, M.J., Thorpe, L.A., and Osborne, B.M., 1995, Characteristics of selected fish 

populations of Arctic National Wildlife Refuge coastal waters, final report, 1988–91: Fairbanks, Alaska, U.S. Fish and 

Wildlife Service, Fairbanks Fishery Resource Office, Alaska Fisheries Technical Report, no. 28, 590 p.

25. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 

1119, 90 p.

26. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224.

27. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Arctic Flounder 591

28. Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbour, N.W.T.: Canadian Technical 

Report of Fisheries and Aquatic Sciences, no. 1218, 21 p.

29. Atkinson, E.G., and Percy, J.A., 1992, Diet comparison among demersal marine fish from the Canadian Arctic: Polar 

Biology, v. 11, no. 8, p. 567–573.

30. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

31. Braund, S.R., and Burnham, D.C., 1984, Subsistence economics and marine resource use patterns, in Truett, J.C., ed., The 

Barrow Arch environment and possible consequences of planned offshore oil and gas development: Outer Continental Shelf 

Environmental Assessment Program, National Oceanic and Atmospheric Administration/Ocean Assessment Division, 

p. 167–215.

32. Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), Beaufort Sea Project, Technical Report 6, 114 p.
592 Alaska Arctic Marine Fish Ecology Catalog

Starry Flounder (Platichthys stellatus) 

(Pallas, 1787)

Family Pleuronectidae 

Colloquial Name: Iñupiaq—Nataagnaq [1].

Ecological Role: Starry flounder are uncommon in 

U.S. Arctic waters. Their abundance is greatest in the southeastern 

Chukchi Sea where they are important predators of the coastal 

marine community.

Physical Description/Attributes: Member of the right eyed 

flounders. Dark brown on eyed side and white on blind side. 

Median fins are white to orange with broad 

black bands. For specific diagnostic characteristics, see Fishes of 

Alaska (Mecklenburg and others, 2002, p. 833) [2]. Swim bladder: 

Absent [2]. Antifreeze glycoproteins in blood serum: Unknown. 

Range: U.S. Chukchi and Beaufort Seas [2]. Elsewhere in Alaska, throughout Bering Sea, Aleutian Islands, Gulf of Alaska and 

all southeastern waters. Worldwide, in Arctic Ocean from the East Siberian Sea eastward to Bathurst Inlet and Viscount Melville 

Sound in the Canadian high Arctic [2, 3]. In Pacific Ocean, southward to Sea of Japan off Korean Peninsula and Sea of Okhotsk, 

and to Los Angeles Harbor, southern California [2].

Starry Flounder (Platichthys stellatus), 290 mm TL, off 

northeastern Sakhalin, Sea of Okhotsk, 2003. Photograph by 

B.A. Sheiko, Russian Academy of Sciences.
Starry Flounder 593

Relative Abundance: Uncommon in U.S. Chukchi and Beaufort Seas [7, 8], becoming more abundant in Canadian Beaufort Sea 

at about the Tuktoyaktuk Peninsula and eastward to at least Coppermine, Amundsen Gulf [9, 10]. Common in Bering Sea [11] to 

Sea of Japan [12], Sea of Okhotsk [13], and southward to central California [14].

tac14-5222_fig3-11-10m_Platichthys_stellatus

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Starry Flounder

Platichthys stellatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Starry Flounder (Platichthys stellatus), within Arctic Outer Continental Shelf 

Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [5, 6].
594 Alaska Arctic Marine Fish Ecology Catalog

Benthic and reproductive distribution of Starry Flounder (Platichthys stellatus).

Some fish to 600 m

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Platichthys stellatus

Starry Flounder

Benthic distribution

Depth, in meters

Reproductive distribution

Open ocean Shore

125

100

75

50

25

0

Depth distribution of eggs 

listed as near surface only

Larvae Eggs

Adults

Juveniles

Data from outstide U.S. Beaufort-Chukchi Seas Data from outstide U.S. Beaufort-Chukchi Seas

Spawning

Habitats and Life History

Eggs—Size: 0.9 to 1.3 mm [20]. Time to hatching: 68 hours at 12.5 °C and 110 hours at 10.5 °C [14]. Pale 

orange in color [21]. Habitat: Pelagic [14, 18]. —Size at hatching: 1.9–2.1 mm SL [20]. Size at juvenile 

transformation: 8.3–10.5 mm SL [22]. Days to juvenile transformation: About 39 days at 12–15 °C in one 

laboratory study [23]. Habitat: Epipelagic, along the coast to the outer shelf [14, 18, 24]. 

Juveniles—Age and size: Age unknown. 8.3–10.5 mm SL to 220 mm TL [22, 25]. Habitat: Benthic [26], in 

backwaters and estuaries, around drift algae, and eelgrass beds [12, 27]. Remain in shallow (often intertidal) 

fresh or brackish waters for several years [12, 28]. 

Adults—Age and size at first maturity: About 8 years and 220 mm TL in Canadian Beaufort Sea based on 

limited data [25]. Elsewhere, varies between 2–4 years and 220–300 mm off central California [14], 2–3 years 

off Washington [29], and about 5 years and 240 mm FL in Sea of Japan [12]. Males mature slightly younger than 

females [14, 29]. Generally, females grow larger than males, live longer, and weigh more at length. Both sexes 

grow faster in warmer temperatures [12, 30]. Maximum age: At least 42 years old in Canadian Beaufort Sea [25]. 

Life spans of fish in warmer waters appear to be much shorter than fish in the Arctic [12, 14]. Maximum size: 

91 cm TL [2]. Habitat: Benthic [26], on soft bottoms and eelgrass beds [12, 27]. 

Substrate—Soft bottoms [12, 27]. 

Physical/chemical—Temperature: -1.8–11.7 °C. Fairly resistant to cold [31]. Spawning occurs at 1.5–6.8 °C 

just after ice break-up in Canadian Beaufort Sea, [30]. Salinity: Mostly marine and estuarine. However, juveniles 

enter fresh water and have been found as much as 121 km upstream in the Columbia River and 32 km upstream 

in the Fraser River [28, 32, 33]. Juveniles (often intertidal) remain in fresh or brackish waters for several years 

[12, 28] and migrate into more saline waters as they mature. Generally, larger adult fish enter estuaries only 

on occasion, for instance at high tide to feed [12, 34, 35]. However, during summer along the Tuktoyaktuk 

Peninsula (Canadian Beaufort Sea), many large fish remained in salinities of 3.7–8.0 parts per thousand [30]. 

Depth Range: Intertidal to 600 m, mainly 100 m or less [15–17]. Larvae are found at least in upper 15 m of water column in 

Gulf of Alaska [18]. In Sea of Japan, juveniles are found less than 55 m [12]. Spawning has been reported at 11–75 m in Bering 

Sea [19].
Starry Flounder 595

Behavior

Diel—Not well understood. At night in summer, adults move into shallow, nearshore waters in Sea of Japan 

[12] and have been found midwaters or near the surface in Puget Sound [27, 36]. Adults exhibit tidally oriented, 

inshore movements [12]. An adult buries itself to avoid predators and can change color to match bottom substrate 

[14]. 

Seasonal—Little is known. In Canadian Beaufort Sea, moves inshore in early summer prior to spawning. Some 

move offshore in autumn [25, 30] and others over winter in nearshore waters [25, 37]. Location of offshore 

wintering is unknown. In Sea of Japan, juveniles winter in coastal estuaries less than 55 m and larger fish winter 

at 45–200 m [12]. 

Reproductive—In Canadian Beaufort Sea, spawning occurs in inshore waters just after ice break-up [30]. In 

California, spawning fish seek shallow water near river mouths and sloughs [14]. 

Schooling—Unknown. 

Feeding—Adults enter estuaries at high tides only to feed [34]. Seems to feed mainly during daylight hours [38].

Populations or Stocks

There have been no studies.

Reproduction

Mode—Separate sexes, oviparous [39]. Fertilization is external [14]. Females are batch spawners [40, 41]. 

Spawning season—June and July in Canadian Beaufort Sea [25, 42]. Between November and February off 

central California and occurs progressively later to the north [14, 25, 42]. 

Fecundity—Between 914,000 and 2,287,000 eggs [40, 41].

Food and Feeding

Food items—In Canadian Beaufort Sea, benthic and epibenthic prey. Clams, crustaceans (for example, mysids 

and isopods), and polychaetes are often most important, and fishes, plants, insects, and priapulids also are 

consumed [25, 30, 43]. Diets are similar in southeastern Chukchi Sea and northeastern Bering Sea, but also 

include brittle stars, cockles, mussels, snails, shrimps, and hermit crabs [44].

Trophic level—3.8 [45].

Biological Interactions

Predators—Ringed seals in U.S. Chukchi Sea [46] and beluga whales in Cook Inlet [47]. Elsewhere fishes such 

as Spotted Spiny Dogfish and White Sturgeon, as well as great blue herons, bald eagles, harbor seals and Stellar 

sea lions [48–52]. 

Competitors—Likely such bottom feeding taxa as sculpins and other flatfishes.

1980 1990 2000 2010

Resilience

Medium, minimum population doubling time 1.4–4.4 years (t

m=2–3; tmax=24) [53].

Traditional and Cultural Importance

Starry Flounder were of minor significance to Iñuits living near the U.S. Chukchi Sea, perhaps due to their 

relative scarcity [54]. Some fish are taken in artisanal fisheries in the Canadian Beaufort Sea and Amundsen 

Gulf, where they are eaten by humans and fed to dogs [3, 37, 55]. Small numbers are commercially harvested 

throughout much of their range and they are frequently taken in the recreational catch. 
596 Alaska Arctic Marine Fish Ecology Catalog

Commercial Fisheries

Currently, Starry Flounder are not commercially harvested.

Potential Effects of Climate Change

Changes in coastal hydrography resulting in increased freshening of coastal waters may expand the habitat and 

range of Starry Flounder in the U.S. Chukchi and Beaufort Seas. Increased abundance will result in increased 

interactions and competition with other marine fishes with unknown ecological effect. Community effects would 

probably be greatest in the nearshore.

Areas for Future Research [B]

Little is known about the biology and ecology of this species from the region. Research needs include: 

(1) depth and location of pelagic larvae; (2) depth, location, and timing of young-of-the-year benthic recruitment; 

(3) preferred depth ranges for juveniles and adults; (4) spawning season; (5) seasonal and ontogenetic 

movements; (6) population studies; (7) prey; and (8) predators.

Remarks

Along the Pacific Coast, Starry Flounder hybridize with English Sole (Parophrys vetulus) and in the western Pacific Ocean, with 

the Stone Flounder (Kareius bicoloratus) [56]. 

References Cited 

Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special reference 

to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, no. 1119, 90 p. [25]

Kolpakov, N.V., 2005, On the biology of Platichthys stellatus (Pleuronectidae) in coastal waters of northern Primorye: Journal 

of Ichthyology, v. 45, no. 8, p. 594–606. [12]

Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p. [30]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [2]

Orcutt, H.G., 1950, The life history of the starry flounder Platichthys stellatus (Pallas): Fish Bulletin, California Fish and Game 

78, 64 p. [14]

Bibliography

1. Craig, P.C., 1989b, Subsistence fisheries at coastal villages in the Alaskan Arctic, 1970–1986: Biological Papers University 

of Alaska, no. 24, p. 131–152.

2. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

3. Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical Report Fisheries and Aquatic Sciences 1910, 135 p.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 
Starry Flounder 597

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

7. Pruter, A.T., and Alverson, D.L., 1962, Abundance, distribution, and growth of flounders in the south-eastern Chukchi Sea: 

Journal de Conseil International pour l’Exploration de la Mer., v. 27, p. 79–99.

8. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

9. Ellis, D.V., 1962, Observations on the distribution and ecology of some Arctic fish: Arctic, v. 15, no. 3, p. 179–189.

10. Bond, W.A., and Erickson, R.N., 1993, Fisheries investigations in coastal waters of Liverpool Bay, Northwest Territories: 

Winnipeg, Manitoba, Canada Department of Fisheries and Oceans, Central and Arctic Region, Canadian Manuscript Report 

of Fisheries and Aquatic Sciences No. 2204, 59 p.

11. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

12. Kolpakov, N.V., 2005, On the biology of Platichthys stellatus (Pleuronectidae) in coastal waters of northern Primorye: 

Journal of Ichthyology, v. 45, no. 8, p. 594–606.

13. Kim, S.T., and Shepeleva, O.N., 2001, The structure of shelf ichthyocenoses of northeastern Sakhalin and Terpeniya Bay: 

Journal of Ichthyology, v. 41, no. 9, p. 711–722.

14. Orcutt, H.G., 1950, The life history of the starry flounder Platichthys stellatus (Pallas): Fish Bulletin, California Fish and 

Game, v. 78, 64 p. 

15. Alverson, D.L., 1960, A study of annual and seasonal bathymetric catch patterns for commercially important groundfishes 

of the Pacific Northwest coast of North America: Pacific Marine Fisheries Commission Bulletin 4, 66 p.

16. Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography of common fishes in the Bering Sea and northeastern Pacific: 

U.S. Department of Commerce, National Oceanic and Atmospheric Administration Technical Report NMFS 66, 151 p.

17. Martin, M.H., and Clausen, D.M., 1995, Data report—1993 Gulf of Alaska Bottom Trawl Survey: U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration, NOAA Technical Memorandum, NMFS-AFSC-59, 217 p.

18. Siefert, D.L., Incze, L.S., and Ortner, P.B., 1988, Vertical distribution of zooplankton, including ichthyoplankton, in 

Shelikof Strait, Alaska—Data from fisheries-oceanography coordinated investigations (FOCI) cruise in May 1986: National 

Marine Fisheries Service, NWAFC Processed Report, v. 88, no. 28, 232 p.

19. Musienko, L.N., 1970, Reproduction and development of Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: Jerusalem, Israel Program for Scientific Translations, p. 161–224.

20. Moser, H.G., 1996, The early stages of fishes in the California current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

21. Hart, J.L., 1973, Pacific fishes of Canada: Ottawa, Fisheries Research Board of Canada, 740 p.

22. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

23. Policansky, D., and Sieswerda, P., 1979, Early life history of the starry flounder, Platichthys stellatus, reared through 

metamorphosis in the laboratory: Transactions of the American Fisheries Society, v. 108, no. 3, p. 326–327.
598 Alaska Arctic Marine Fish Ecology Catalog

24. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.

25. Bond, W.A., 1982, A study of the fishery resources of Tuktoyaktuk Harbour, southern Beaufort Sea coast, with special 

reference to life histories of anadromous coregonids: Canadian Technical Report of Fisheries and Aquatic Sciences, 

no. 1119, 90 p.

26. McGregor, S.B., 1990, Seasonal and ontogenetic changes in meiofauna in the diets of postmetamorphic flatfishes: 

University of Alaska, Fairbanks, Master’s thesis.

27. Miller, B.S., Simenstad, C.A., Moulton, L.L., Fresh, K.L., Funk, F.C., Karp, W.A., and others, 1977, Puget Sound baseline 

program—Nearshore fish survey: University of Washington, Fishery Research Institute, FRI-UW-7710, 219 p.

28. Haertel, L., and Osterberg, C., 1967, Ecology of zooplankton, benthos and fishes in the Columbia River Estuary: Ecology, 

v. 48, no. 3, p. 459–472.

29. Smith, R.T., 1936, Report on the Puget Sound otter trawl investigations: Washington Department of Fisheries, Biological 

Report No. 36B, 120 p.

30. Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey of the coastal fishes of the southeastern Beaufort Sea: Canadian 

Technical Report of Fisheries and Aquatic Sciences, no. 1220, 178 p.

31. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

32. McAllister, D.E., 1959, Records of marine fishes from fresh water in British Columbia: The Canadian Field-Naturalist, 

v. 73, no. 1, p. 13–14.

33. Baxter, J.L., 1966, Inshore fishes of California: California Department of Fish and Game, 80 p.

34. Tokranov, A.M., and Maksimenkov, V.V., 1994, Feeding of the starry flounder, Platichthys stellatus, in the Bol’shaya River 

estuary (Western Kamchatka): Journal of Ichthyology, v. 34, no. 1, p. 76–83.

35. Rooper, C.N., 2002, English sole transport during pelagic stages on the Pacific Northwest coast, and habitat use by juvenile 

flatfish in Oregon and Washington estuaries: Seattle, University of Washington, Ph.D. dissertation.

36. Fresh, K.L., 1979, Distribution and abundance of fishes occurring in the nearshore surface waters of northern Puget Sound, 

Washington: Seattle, University of Washington, Master’s thesis.

37. Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

38. Miller, B.S., 1967, Stomach contents of adult starry flounder and sand sole in East Sound, Orcas Island, Washington: 

Journal of the Fisheries Research Board of Canada, v. 24, no. 12, p. 2,515–2,521.

39. Garrison, K.J., and Miller, B.S., 1982, Review of the early life history of Puget Sound fishes: Seattle, University of 

Washington, Fisheries Research Institute, 729 p.

40. Fadeev, N.S., 1965, Comparative outline of the biology of flatfishes in the southeastern part of the Bering Sea and condition 

of their resources, in Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part IV: Jerusalem, Israel 

Program for Scientific Translations, p. 112–146.

41. Fadeev, N.S., 1970, The fishery and biological characteristics of yellowfin soles in the eastern part of the Bering Sea in

Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part V: Jerusalem, Israel Program for Scientific 

Translations, p. 332–396.

42. Ratynski, R.A., 1983, Mid-summer ichthyoplankton populations of Tuktoyaktuk Harbour, N.W.T.: Canadian Technical 

Report of Fisheries and Aquatic Sciences, no. 1218, 21 p.

43. Lacho, G., 1991, Stomach content analyses of fishes from Tuktoyaktuk Harbour, N.W.T., 1981: Winnipeg, Manitoba, 

Canadian Data Report of Fisheries and Aquatic Sciences, Central and Arctic Region, Department of Fisheries and Oceans, 

no. 853, 15 p.
Starry Flounder 599

44. Jewett, S.C., and Feder, H.M., 1980, Autumn food of adult starry flounders, Platichthys stellatus, from the northeastern 

Bering Sea and the southeastern Chukchi Sea: Journal de Conseil International pour l’Exploration de la Mer., v. 39, no. 1, 

p. 7–14.

45. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320. 

46. Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, M.L., 1966, Marine mammals, in Wilimovsky, N.J., and Wolfe, 

J.N., eds., Environment of the Cape Thompson Region, Alaska: Oak Ridge, Tennesee, United States Atomic Energy 

Commission, Division of Technical Information, p. 877–924.

47. Huntington, H.P., 2000, Traditional knowledge of the ecology of belugas, Delphinapterus leucas, in Cook Inlet, Alaska: 

Marine Fisheries Review, v. 62, p. 134–140.

48. Bonham, K., 1954, Food of the dogfish Squalus acanthias: Washington Department of Fisheries, v. 1, no. 2, p. 25–36.

49. McKechnie, R.J., and Fenner, R.B., 1971, Food habits of white sturgeon, Acipenser transmontanus in San Pablo and Suisun 

Bays, California: California Fish and Game, v. 57, p. 209–212.

50. Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, p. 973–990. 

51. Armstrong, R.H., Carstensen, R.L., and Wilson, M.F., 2004, Hotspots—Bird survey of Mendenhall Wetlands April 2002 to 

May 2003: Juneau Audubon Society and Taku Conservation Society, 9 p.

52. Orr, A.J., Banks, A.S., Mellman, S., Huber, H.R., and DeLong, R.L., 2004, Examination of the foraging habits of Pacific 

harbor sea (Phoca vitulina richardsoni) to describe their use of the Umpqua River, Oregon, and their predation on 

salmonids: Fishery Bulletin, v. 102, no. 1, p. 108–117.

53. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

54. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115.

55. Gillman, D.V., and Kristofferson, A.H., 1984, Biological data on Pacific herring (Clupea harengus pallasi) from 

Tuktoyaktuk Harbour and the Liverpoor Bay area, Northwest Territories, 1981 to 1983: Winnepeg, Manitoba, Canada, 

Canadian Data Report of Fisheries and Aquatic Sciences, Department of Fisheries and Oceans, Western Region, no. 485, 

22 p.

56. Garrett, D.L., Pietsch, T.W., Utter, F.M., and Hauser, L., 2007, The hybrid sole, Inopsetta ischyra (Teleostei: 

Pleuronectiformes: Pleuronectidae)—Hybrid or biological species?: Transactions of the American Fisheries Society, v. 136, 

no. 2, p. 460–468.
600 Alaska Arctic Marine Fish Ecology Catalog

Alaska Plaice (Pleuronectes quadrituberculatus) 

Pallas, 1814

Family Pleuronectidae 

Note: Except for geographic range data, all information is from 

areas outside the study area. 

Colloquial Name: None in U.S. Chukchi and 

Beaufort Seas.

Ecological Role: Largely unknown. This species is absent from 

the U.S. Beaufort Sea and is present only occasionally in the U.S. 

Chukchi Sea. It is unlikely to represent a significant prey resource 

to higher level organisms. 

Physical Description/Attributes: Member of the right eyed 

flounders. Eyed side is greenish gray to almost black and blind 

side is yellow. Young adults are spotted and blotched. For specific 

diagnostic characteristics, see Fishes of Alaska (Mecklenburg and 

others, 2002, p. 835) [1]. Swim bladder: Absent [1]. Antifreeze glycoproteins in blood serum: Present [2].

Range: U.S. Chukchi Sea, as far northward as 70°16’N, 163°58’W [3]. Elsewhere in Alaska, from Bering Sea to eastern 

Aleutian Islands and Gulf of Alaska to southeastern Alaska near Ketchikan [1]. Worldwide, to Sea of Japan; one record from 

Bellingham Bay, Washington [1].

Alaska Plaice (Pleuronectes quadrituberculatus), 60 cm TL, 

off northeastern Sakhalin, Okhotsk Sea, 2003. Photograph by 

B.A. Sheiko, Russian Academy of Sciences.
Alaska Plaice 601

Relative Abundance: Occasional in U.S. Chukchi Sea [3, 6, 7]. Common from eastern Bering Sea to central Gulf of Alaska [8] 

and from Sea of Japan [9] to Kamchatka Peninsula [10].

tac14-5222_fig3-11-12m_Pleuronectes_quadrituberculatus

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Point Lay

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Alaska Plaice

Pleuronectes quadrituberculatus

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Alaska Plaice (Pleuronectes quadrituberculatus) within Alaska Outer Continental 

Shelf Planning Areas [4] based on review of published literature and specimens from historical and recent 

collections [1, 3, 5]. 
602 Alaska Arctic Marine Fish Ecology Catalog

Benthic and reproductive distribution of Alaska Plaice (Pleuronectes quadrituberculatus). 

500

400

300

200

100

0

Adults

Juveniles

Data from outside U.S. Beaufort-Chukchi Seas Data from outside U.S. Beaufort-Chukchi Seas

Open ocean Shore

Depth, in meters

500

400

300

200

100

0

Pleuronectes quadrituberculatus

Alaska Plaice

Benthic distribution

Shore

Depth, in meters

Open ocean

Spawning

Larvae

and eggs

Larvae

Reproductive distribution

Some fish to 900 m 

in winter

Habitats and Life History

Eggs—Size: 1.6–2.2 mm [17–19]. Time to hatching: 16 days to two months [20]. Habitat: Epipelagic [16]. 

Larvae—Size at hatching: 5.6–5.8 mm SL [17–19]. Size at juvenile transformation: 13–17 mm SL [16]. Days to 

juvenile transformation: Unknown. Habitat: Pelagic [16]. 

Juveniles—Age and size: 1.6–17 mm SL. Habitat: Benthic, inshore waters [8, 16]. 

Adults—Age and size at first maturity: In eastern Bering Sea, few females mature at 27 cm TL (about 7 years), 

50 percent at 31 cm (8 years), and 100 percent at 39 cm (11 years) [16]. Off western Kamchatka Peninsula, 

Russia, most males mature at 26–30 cm (6–7 years) and females at 32–36 cm (8–9 years old) [21]. Females grow 

larger and faster than males [16]. Larger females are heavier at length than males [16]. Maximum age: At least 37 

years [22]. Maximum size: 62 cm TL [1]. Habitat: Benthic, mostly on continental shelf [8, 16]. 

Substrate—Sand and mud [8]. 

Physical/chemical—Temperature: Between -1.7 and 12 °C [5, 23]. Salinity: Marine [8].

Behavior

Diel—Unknown. 

Seasonal—During summer in eastern Bering Sea, females tend to be in deeper (60 m) water than males 

(45–55 m) [16]. 

Reproductive—Prior to spawning in spring, mature plaice migrate into somewhat shallower waters. Do not 

aggregate for spawning, but spawn over a wide area of the middle shelf [16]. 

Schooling—Unknown. 

Feeding—Probably do not feed at night or in winter [16, 24].

Populations or Stocks

There have been no studies.

Depth Range: Between 5 and 500 m, mainly 100 m or less in northeast Pacific Ocean and Bering Sea [8, 11]. In winter, occurs 

much deeper (400–900 m) around Sakhalin Island, Sea of Okhotsk [12, 13]. Eggs occur from 0–60 m, mainly 30 m or less [14]. 

Larvae are found between 0–149 m [15], mainly 20 m or less [14]. Juveniles are found inshore, mostly 50 m or less [8]. Most 

spawning occurs at 50–100 m [16].
Alaska Plaice 603

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [16]. 

Spawning season—In eastern Bering Sea spawning occurs from March to July and peak spawning season varies 

with year and area [16, 24]. 

Fecundity—Between 56,000 and 521,000 eggs per season and are probably batch spawners [25, 26].

Food and Feeding

Food items—Benthic and epibenthic organisms are most important in eastern Bering Sea. In particular, 

polychaetes, clams, and such crustaceans as gammarid amphipods dominate the diets. Other preys include 

echiurioids, brittle stars, shrimps, snails, sea urchins, and sand dollars [27, 28]. 

Trophic level—3.5 [29].

Biological Interactions

Predators—Alaska Plaice, Flathead Sole, Great and Plain sculpins, and Walleye Pollock [27, 28, 30, 31]. 

Competitors—Flatfishes such as Yellowfin and Rock soles, as well as various sculpins, poachers, and other 

bottom feeders have been reported south of the Bering Strait [16].

1980 1990 2000 2010

Resilience 

Low, minimum population doubling time 4.5–14 years (K=0.1) [32].

Traditional and Cultural Importance

Historically, Alaska Plaice were sought in various subsistence fisheries [12, 33]. Currently, they are taken 

primarily as bycatch in other fisheries [8].

Commercial Fisheries

Currently, Alaska Plaice are not commercially harvested.

Potential Effects of Climate Change

Because Alaska Plaice are common in the Bering Sea and only occasionally in the U.S. Chukchi Sea, populations 

would be expected to increase as the climate warms. 

Areas for Future Research [B]

Alaska Plaice are common in the Bering Sea. Distributional shifts in the population into the Chukchi Sea are 

possible. Reliable baseline information about distribution and abundance and their use of habitats is needed to 

evaluate possible climate effects on demersal fish assemblages.
604 Alaska Arctic Marine Fish Ecology Catalog

References Cited 

Bailey, K.M., Brown, E.S., and Duffy-Anderson, J.T., 2003, Aspects of distribution, transport and recruitment of Alaska 

plaice (Pleuronectes quadrituberculatus) in the Gulf of Alaska and eastern Bering Sea: comparison of marginal and central 

populations: Journal of Sea Research, v. 50, no. 2–3, p. 87–95. [8]

Duffy-Anderson, J.T., Doyle, M.J., Mier, K.L., Stabeno, P.J., and Wilderbuer, T.K., 2010, Early life ecology of Alaska plaice 

(Pleuronectes quadrituberculatus) in the eastern Bering Sea— Seasonality, distribution, and dispersal: Journal of Sea 

Research, v. 64, no. 1–2, p. 3–14. [14]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [1]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [3]

Zhang, C.I., Wilderbuer, T.K., and Walters, G.E., 1998, Biological characteristics and fishery assessment of Alaska plaice, 

Pleuronectes quadritubercultus, in the eastern Bering Sea: Marine Fisheries Review: v. 60, no. 4, p. 16–27. [16]

Bibliography

1. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

2. Power, G., 1997, A review of fish ecology in Arctic North America: American Fisheries Society Symposium, no. 19, 

p. 13–39.

3. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

4. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

5. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

6. Pruter, A.T., and Alverson, D.L., 1962, Abundance, distribution, and growth of flounders in the south-eastern Chukchi Sea: 

Journal de Conseil International pour l’Exploration de la Mer., v. 27, p. 79–99.

7. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.

8. Bailey, K.M., Brown, E.S., and Duffy-Anderson, J.T., 2003, Aspects of distribution, transport and recruitment of Alaska 

plaice (Pleuronectes quadrituberculatus) in the Gulf of Alaska and eastern Bering Sea: comparison of marginal and central 

populations: Journal of Sea Research, v. 50, no. 2–3, p. 87–95.

9. Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 1998, A list of fishes of Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 38, no. 1, p. 1–11.

10. Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution in the Kronitsky Bay in relation to abiotic factors: Zoologicheskii 

Zhurnal, v. 63, p. 1,363–1,373.

11. Mueter, F.J., and Norcross, B.L., 2002, Spatial and temporal patterns in the demersal fish community on the shelf and upper 

slope regions of the Gulf of Alaska: Fishery Bulletin 100, no. 3, p. 559–581. 

12. Kim, S.T., Biryukov, I.A., and Pometeev, E.V., 1999, Species composition, distribution, and structural changes in the flatfish 

community of the Tatarskii Strait during the fishing period 1994–1997: Journal of Ichthyology, v. 39, no. 6, p. 433–440.

13. Kim, S.T., 2001, Winter migrations of shelf fish to the continental slope: Journal of Ichthyology, v. 41, no. 8, p. 564–574.
Alaska Plaice 605

14. Duffy-Anderson, J.T., Doyle, M.J., Mier, K.L., Stabeno, P.J., and Wilderbuer, T.K., 2010, Early life ecology of Alaska 

plaice (Pleuronectes quadrituberculatus) in the eastern Bering Sea— Seasonality, distribution, and dispersal: Journal of Sea 

Research, v. 64, no. 1–2, p. 3–14.

15. Siefert, D.L., Incze, L.S., and Ortner, P.B., 1988, Vertical distribution of zooplankton, including ichthyoplankton, in 

Shelikof Strait, Alaska—Data from fisheries-oceanography coordinated investigations (FOCI) cruise in May 1986: National 

Marine Fisheries Service, NWAFC Processed Report, v. 88, no. 28, 232 p.

16. Zhang, C.I., Wilderbuer, T.K., and Walters, G.E., 1998, Biological characteristics and fishery assessment of Alaska plaice, 

Pleuronectes quadritubercultus, in the eastern Bering Sea: Marine Fisheries Review: v. 60, no. 4, p. 16–27.

17. Grigorev, S.S., and Fadeev, N.S., 1995, Distribution of Alaska plaice, flathead sole, and Bering flounder eggs in the eastern 

Bering Sea during April–July, 1988–1990, in Alaska Sea Grant Program, eds., Proceeding of the International Symposium 

on North Pacific Flatfish: Fairbanks, University of Alaska, Alaska Sea Grant Program, AK-SG-95-04.

18. Grigorev, S.S., 1995, Characteristics of flatfish eggs and larva samples near eastern Kamchatka and in the western Bering 

Sea during June-July 1991, in Alaska Sea Grant Program, eds., Proceedings of the International Symposium on North 

Pacific Flatfish: University of Alaska, Fairbanks, Alaska Sea Grant Program, AK-SG-95-04, p. 101–116.

19. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

20. Salveson, S.J., 1976, Alaska plaice (family Pleuronectidae), in Pereya, W.T., Reeves, J.E., and Bakkala, R.G., eds., 

Demersal fish and shellfish resources of the Eastern Bering Sea in the baseline year 1975: National Oceanic and 

Atmospheric Administration, p. 489–495.

21. Tokranov, A.M., and Zavarina, S.V., 1992, Size-age structure and sex ratios of Alaska plaice, Pleuronectes 

quadrituberculatus, on the Western Kamchatka Shelf: Journal of Ichthyology, v. 32, no. 7, p. 118–127.

22. National Oceanic and Atmospheric Administration, 2010, Age and growth program: National Oceanic and Atmospheric 

Administration, accessed April 2011 at http://www.afsc.noaa.gov/refm/age/Stats/max_age.htm.

23. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

24. Mineva, T.A., 1964, On the biology of some flatfishes in the eastern Bering Sea, in Moiseev, P.A. ed., Soviet fisheries 

investigations in the northeastern Pacific, Part II: Jerusalem, Israel Program for Scientific Translations, p. 227–235.

25. Fadeev, N.S., 1965, Comparative outline of the biology of flatfishes in the southeastern part of the Bering Sea and condition 

of their resources, in Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part IV: Jerusalem, Israel 

Program for Scientific Translations, p. 112–146.

26. Fadeev, N.S., 1970, The fishery and biological characteristics of yellowfin soles in the eastern part of the Bering Sea, in

Moiseev, P.A., ed., Soviet fisheries investigations in the northeastern Pacific, Part V: Jerusalem, Israel Program for Scientific 

Translations, p. 332–396.

27. Brodeur, R.D., and Livingston, P.A., 1988, Food habits and diet overlap of various Eastern Bering Sea fishes: National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS F/NWC-127, 76 p.

28. Livingston, P.A., Ward, A., Lang, G.M., and Yang, M.-S., 1993, Groundfish food habits and predation on commercially 

important prey species in the eastern Bering Sea from 1987 to 1989: National Oceanic and Atmospheric Administration 

Technical Memorandum NMFS-AFSC-11, 202 p.

29. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.

30. Tokranov, A.M., 1987a, Feeding of giant sculpin Myoxocephalus polyacanthocephalus Pallas and plain sculpin M. jaok

(Cuvier) (Cottidae) in coastal waters of Kamchatka: Journal of Ichthyology, v. 27, p. 104–114.
606 Alaska Arctic Marine Fish Ecology Catalog

31. Pacunski, R.E., Livingston, P.A., and Miller, B.S., 1998, Food of flathead sole Hippoglossoides elassodon in the eastern 

Bering Sea: National Oceanic and Atmospheric Administration, Technical Memorandum NMFS-AFSC-90, 36 p.

32. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.

33. Bean, T.H., 1887, The fishery resources and fishing-grounds of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States Commission of Fish and Fisheries, p. 81–115
Greenland Halibut 607

Greenland Halibut (Reinhardtius hippoglossoides) 

(Walbaum, 1792)

Family Pleuronectidae 

Colloquial Name: None in U.S. Chukchi and 

Beaufort Seas.

Ecological Role: Poorly understood, but likely of limited 

importance. Potentially, a relatively dominant species in offshore 

Beaufort Sea.

Physical Description/Attributes: Member of the right eyed 

flounders. Both eyed and blind sides are pigmented. Eyed side varies 

among black, brown, purplish brown, grey, or blue, and blind side 

is similarly colored, although lighter [1–3]. For specific diagnostic 

characteristics, see Fishes of Alaska (Mecklenburg and others, 2002, 

p. 830) [3]. Swim bladder: Absent [3]. Antifreeze glycoproteins in blood serum: Unknown. 

Range: U.S. Chukchi and Beaufort Seas. Elsewhere in Alaska, throughout Bering Sea, Aleutian Islands, Gulf of Alaska, and all 

southeastern waters [3]. Worldwide, on continental slope north of Alaska, in North Pacific from Seas of Japan and Okhotsk to 

just south of U.S.–Mexico border; in North Atlantic, from Ungava Bay, Canada, to northwest Greenland and south to Gulf of 

Maine; northeast Greenland, Iceland, northern Norway, White Sea, and Kara Sea [4].

Greenland Halibut (Reinhardtius hippoglossoides), 437 mm, 

Chukchi Borderland, 2009. Photograph by C.W. Mecklenburg, 

Point Stephens Research.
608 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_fig3-11-13m_Reinhardtius_hippoglossoides

200

200

50

200

50

50

50

50

YUKON

NORTHWEST

TERRITORIES

NUNAVUT

BRITISH

COLUMBIA

Chukchi Sea

Bering Sea

Bering

Strait

Mackenzie River

Colville River

Yukon River

200

ARCTIC OCEAN

Beaufort Sea

Mackenzie

River Delta

Point

Barrow

Point Hope

Nome

Prudhoe 

Bay

UNITED STATES

CANADA

RUSSIA

Banks

Island

St Lawrence

Island

Wrangel

Island

Victoria

Island

ALASKA

Hanna

Shoal

Canyon

Harrison

Bay

Hulahula R

Norton Sound

B R O O K S

R A N G E

Barrow

Kaktovik

Kotzebue

Sound

Greenland Halibut

Reinhardtius hippoglossoides

170°E 180° 170°W 160°W 150°W 140°W 130°W 120°W 110°W

76°N

74°N

72°N

70°N

68°N

66°N

64°N

62°N

0 100

100

50

50

200 MILES

0 200 KILOMETERS

Base modified from USGS and other digital data. U.S.-Russia Maritime Boundary follows the 

EEZ/200-mile limit line, western edge. Coordinate reference system: projection, Lambert Azimuthal 

Equal Area; latitude of origin, 75.0°; horizontal datum, North American Datum of 1983.

Chukchi-Beaufort lease area

Geographic distribution

EXPLANATION

50 Depth of water, in meters

U.S. Exclusive Economic 

Zone (200-mile limit)

UNITED

STATES

Arctic

Ocean

Pacific

Ocean

RUSSIA CANADA

Geographic distribution of Greenland Halibut (Reinhardtius hippoglossoides), within Arctic Outer Continental 

Shelf Planning Areas [5] based on review of published literature and specimens from historical and recent 

collections [4, 6]. 

Relative Abundance: Uncommon in U.S. Chukchi and Beaufort Seas [4, 7]. Elsewhere in Alaska, common along Aleutian 

Islands to Bering Sea [8, 9]. Worldwide, common in Seas of Japan and Okhotsk [10], and along northern Kuril Islands and 

southeastern Kamchatka Peninsula, Russia [11, 12] but rare south of Alaska [13]. There has been a general northward movement 

of populations in southeastern Bering Sea [14].
Greenland Halibut 609

Benthic and reproductive distribution of Greenland Halibut (Reinhardtius hippoglossoides). 

Open ocean Shore

Depth, in meters

2,000

1,600

1,200

800

400

0

Adults

Some fish to 2,000 m

Juveniles

Reinhardtius hippoglossoides

Greenland Halibut

Benthic distribution

Open ocean Shore

Depth, in meters

Spawning

 Data from outside 

U.S. Beaufort-Chukchi Seas

Reproductive distribution

Larvae

Eggs

1,250

1,000

750

500

250

0

 Data from outside 

U.S. Beaufort-Chukchi Seas

Habitats and Life History

Eggs—Size: 3.7–4.1 mm [15, 19]. Time to hatching: 53 days at 4 °C in Atlantic Ocean [17]. Habitat: 

Bathypelagic, in eastern Bering Sea [15, 19]. 

Larvae—Size at hatching: 10–16 mm SL [15, 20]. Size at juvenile transformation: As small as 34 mm FL in 

western Bering [16] and about 8.0 cm TL in Hudson Bay [21]. Days to juvenile transformation: 8–9 months in 

eastern Bering Sea and Gulf of Alaska [22]. Habitat: Bathypelagic to pelagic [15, 20], from oceanic to coastal 

waters [23]. Larvae 2.7 cm and larger vertically migrate [19]. 

Juveniles—Age and size: As small as 60 mm FL in eastern Bering Sea and Gulf of Alaska [17]. Habitat: 

Benthopelagic, mainly on continental shelf and migrate onto slope at around 4–5 years of age [15, 16]. 

Adults—Age and size at first maturity: Poorly known in eastern Bering Sea, but 50 percent maturity of females 

is likely at 65–70 cm FL [24] and about 10–11 years old [25]. Worldwide, estimates of size of females at 

50 percent maturity range from 48–80 cm FL. Females grow larger than males, are larger at age, and weigh more 

at length [25–28]. Maximum age: Ageing techniques for Greenland Halibut are inadequate and growth rates are 

likely to be much slower than previously estimated [29]. Live to at least 36 years, given limitations of current 

ageing techniques, [28]. Maximum size: 130 cm TL [3] and 44 kg [30]. Habitat: Benthopelagic [4]. 

Substrate—Sand and mud. 

Physical/chemical—Temperature: -1.7 to about 10 °C, primarily 0–5 °C [15, 31–33]. Juveniles seem to tolerate 

colder waters than adults [31]. Salinity: Marine [15, 31–33].

Behavior

Diel—Often found midwaters, unusual behavior for flatfishes. Off Greenland and Norway, nocturnal excursions 

are made hundreds of meters off bottom and fish remain in water column during night [2, 34]. Similar behavior 

may occur in Bering Sea [25]. 

Seasonal—Adults appear to make limited, seasonal bathymetric movements, entering somewhat shallower 

waters in spring and summer [31, 35]. During at least some parts of year, females may tend to be in deeper 

waters than males [11, 36]. In eastern Bering Sea, larvae drift from spawning grounds in southern slope area 

northward to northern shelf area, followed by a gradual shifting of immature and maturing fish to deeper and 

more southern waters [15]. Some individuals make extensive horizontal movements of as much as 687 km [25]. 

Reproductive—Thought to spawn synchronously [37]. 

Schooling—Appears to have no tendency to schooling [38]. 

Feeding—At least some feed well up in water column [39].

Depth Range: Overall, 14–2,000 m, usually at 50–650 m [3, 4]. Young juveniles live mainly on continental shelf (less than 

200 m) [15, 16]. Spawning typically occurs along continental slope between 400 and 1,100 m [17]. In Atlantic Ocean, eggs 

are mesopelagic (600–1,000 m) [17]. In Bering Sea, eggs are at 50–400 m, mostly at 200–300 m [18]. Early larvae are found 

throughout water column to 530 m, mostly down to 450 m [17]. Older larvae in 0–530 m depths, mostly at less than 45 m [18].
610 Alaska Arctic Marine Fish Ecology Catalog

Populations or Stocks

There have been no studies. 

Reproduction

Mode—Separate sexes, oviparous. Fertilization is external [37]. Perhaps a batch spawner [40]. In Atlantic 

Ocean, egg maturation may take longer than 1 year or else some mature females may not reproduce in a given 

year [37]. 

Spawning season—At least August to March (peaking November through February) in Bering Sea, [16, 31].

Fecundity—At least 23,900–149,000 eggs [41]. 

Food and Feeding

Food items—No studies have been conducted in U.S. Chukchi and Beaufort Seas. In Gulf of Alaska and 

along Aleutian Islands, a wide range of fishes (often Walleye Pollock) are dominant, although squids, 

octopi, euphausiids, and polychaetes also are consumed [42–44]. Smaller individuals feed on both fishes and 

invertebrates, whereas larger individuals primarily target fishes 

Trophic level—4.6 [14].

Biological Interactions

Predators—No studies have been conducted. Elsewhere, thick-billed murre, ringed seal, and narwhal in 

Canadian Arctic [45–47]. Flathead Sole, Greenland Halibut, Walleye Pollock, Yellowfin Sole, northern fur seal, 

and ribbon seal in eastern Bering Sea [48–51]. 

Competitors—Unknown in U.S. Chukchi and Beaufort Seas. As it is both a benthic and midwater predator, this 

species likely competes with cods, including Walleye Pollock, and flatfishes. 

1980 1990 2000 2010

Resilience

Low, minimum population doubling time 4.5–14 years (K=0.07–0.10; t

min=7–12; tmax=30; Fecundity=6,800) [52].

Traditional and Cultural Importance

None reported.

Commercial Fisheries

Currently, Greenland Halibut are not commercially harvested. In U.S. waters outside of the Chukchi and 

Beaufort Seas, Greenland Halibut are taken with bottom trawls and longlines in a directed fishery and as bycatch 

in fisheries for sablefish and Pacific cod [22].

Potential Effects of Climate Change

Because Greenland Halibut are common in the Bering Sea but uncommon farther north and their population 

appears to be shifting northward in the southeastern Bering Sea, abundance and limits of distribution would be 

expected to increase in the U.S. Chukchi and Beaufort Seas as climate warms. 
Greenland Halibut 611

Areas for Future Research [A]

The species may be more common in the Beaufort Sea than previously thought. From a zoogeographic 

standpoint, improved information about the geographic distribution and origins of the species is of interest. It is 

hypothesized that the species may be of North Atlantic origins and this could be resolved with genetic research to 

determine biogeographic and phylogenetic relationships. All aspects of the species biology and ecology require 

further attention, but information about important habitats and seasonal movements and migrations is most

 important.

References Cited

Alton, M.S., Bakkala, R.G., Walters, G.E., and Munro, P.T., 1988, Greenland turbot Reinhardtius hippoglossoides of the eastern 

Bering Sea and Aleutian Island region: National Oceanic and Atmospheric Administration, Technical Report NMFS 71, 31 p. 

[15]

Ianelli, J.N., Wilderbuer, T.K., and Nichol, Dan, 2007, Assessment of Greenland turbot in the eastern Bering Sea and Aleutian 

Islands: North Pacific Fisheries Management Council, Bering Sea and Aleutian Islands SAFE Document, p. 513–562, 

accessed April 6, 2014 at http://www.afsc.noaa.gov/refm/docs/2007/BSAIturbot.pdf. [25]

Kodolov, L.S., and Matveychuk, S.P., 1995, Stock condition of Greenland turbot (Reinhardtius hippoglossoides matsuurae

Jordan et Snyder) in the northwestern Bering Sea: Anchorage, Alaska Sea Grant Program AK-SG-95-04. [16]

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p. [3]

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and zoogeography: 

Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1. [4]

Sohn, D., Ciannelli, L., and Duffy-Anderson, J.T., 2010, Distribution and drift pathways of Greenland halibut (Reinhardtius 

hippoglossoides) during early life stages in the eastern Bering Sea and Aleutian Islands: Fisheries Oceanography, v. 19, no. 5, 

p. 339–353. [17]

Bibliography

1. Burton, D.W., 1988, Melanophore comparisons in different forms of ambicoloration in the flatfish Pseudopleuronectes 

americanus and Reinhardtius hippoglossoides: Journal of Zoology, v. 214, no. 2, p. 353–360.

2. Jørgensen, O.A., 1997, Pelagic occurrence of Greenland halibut, Reinhardtius hippoglossoides (Walbaum), in west 

Greenland waters: Journal of Northwest Atlantic Fisheries Science, v. 21, p. 39–50.

3. Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2002, Fishes of Alaska: Bethesda, Maryland, American 

Fisheries Society, 1,116 p.

4. Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–140, Online Resource 1.

5. Minerals Management Service, 2008, Beaufort Sea and Chukchi Sea planning areas—Oil and Gas Lease Sales 209, 212, 

217, and 221: U.S. Department of the Interior, Minerals Management Service Alaska OCS Region, OCS EIS/EA, MMS 

2008-0055, 538 p. 

6. Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., and Steinke, D., 2016, Pacific Arctic marine fishes: Akureyri, 

Iceland, Conservation of Arctic Flora and Fauna, Monitoring Series Report No. 23, 406 p., accessed May 10, 2016, at 

http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes.

7. Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 1997, Demersal fish assemblages of the northeastern Chukchi 

Sea, Alaska: Fishery Bulletin, v. 95, no. 2, p. 195–209.
612 Alaska Arctic Marine Fish Ecology Catalog

8. Ronholt, L.L., Wakabayashi, K., Wilderbuer, T.K., Yamaguchi, H., and Okada, K., 1986, Ground fish resource of the 

Aleutian Island waters based on the U.S.–Japan trawl survey, June–November 1980: Bulletin International North Pacific 

Fisheries Commission 48, p. 251.

9. Hoff, G.R., 2006, Biodiversity as an index of regime shift in the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

10. Kim, S.T., and Shepeleva, O.N., 2001, The structure of shelf ichthyocenoses of northeastern Sakhalin and Terpeniya Bay: 

Journal of Ichthyology, v. 41, no. 9, p. 711–722.

11. Dudnik, Y.I., and Dolganov, V.N., 1992, Distribution and abundance of fish on the continental slopes of the Sea of Okhotsk 

and of the Kuril Islands during the summer of 1989: Journal of Ichthyology, v. 32, no. 9, p. 58–76.

12. Tokranov, A.M., Orlov, A.M., and Biryukov, I.A., 2004, Distribution and size-weight composition of some rare fish species 

in Pacific waters off the northern Kurile Islands and southeast Kamchatka: Journal of Ichthyology, v. 44, no. 3, p. 208–216.

13. Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist of North Pacific and Arctic Ocean species from Baja California 

to the Alaska-Yukon border: U.S. Geological Survey, OCS Study MMS 2005-030 and USGS/NBII 2005-001, 276 p.

14. Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters the biogeography of the Bering Sea continental shelf: Ecological 

Applications, v. 18, no. 2, p. 309–320.

15. Alton, M.S., Bakkala, R.G., Walters, G.E., and Munro, P.T., 1988, Greenland turbot Reinhardtius hippoglossoides of 

the eastern Bering Sea and Aleutian Island region: National Oceanic and Atmospheric Administration, Technical Report 

NMFS 71, 31 p.

16. Kodolov, L.S., and Matveychuk, S.P., 1995, Stock condition of Greenland turbot (Reinhardtius hippoglossoides matsuurae

Jordan et Snyder) in the northwestern Bering Sea: Anchorage, Alaska Sea Grant Program AK-SG-95-04.

17. Sohn, D., Ciannelli, L., and Duffy-Anderson, J.T., 2010, Distribution and drift pathways of Greenland halibut (Reinhardtius 

hippoglossoides) during early life stages in the eastern Bering Sea and Aleutian Islands: Fisheries Oceanography, v. 19, 

no. 5, p. 339–353.

18. Blood, D.M., Duffy-Anderson, J.T., Matarese, A.C., Sohn, D., Stabeno, P.J., and Cheng, W., 2009, Early life history of 

Greenland halibut (Reinhardtius hippoglossoides) in the eastern Bering Sea based on recent field studies (2007–2009)—

Spawning, distribution, and physical processes affecting drift and dispersal: Portland, Oregon, 33rd Annual Larval Fish 

Conference and American Society of Ichthyologists and Herpetologists.

19. Bulatov, O.A., 1983, Distribution of eggs and larvae of Greenland halibut, Reinhardtius hippoglossoides (Pleuronectidae), 

in the eastern Bering Sea: Journal of Ichthyology, v. 23 no. 1, p. 157–159.

20. Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, J.L., 2003, Atlas of abundance and distribution patterns of 

ichthyoplankton from the northeast Pacific Ocean and Bering Sea ecosystems based on research conducted by the Alaska 

Fisheries Science Center (1972–1996): National Oceanic and Atmospheric Administration Professional Paper NMFS 1, 

281 p.

21. Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions to the study of the fishes of Ungava Bay: Journal of the Fisheries 

Research Board of Canada, v. 9, no. 2, p. 83–126.

22. North Pacific Fishery Management Council, 2009, Fishery management plan for fish resources of the Arctic management 

area: Anchorage, Alaska, North Pacific Fishery Management Council, 158 p.

23. Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in Oceanography, v. 53, no. 2, p. 247–281.

24. Cooper, D.W., Maslenikov, K.P., and Gunderson, D.R., 2007, Natural mortality rate, annual fecundity, and maturity at 

length for Greenland halibut (Reinhardtius hippoglossoides) from the northeastern Pacific Ocean: Fishery Bulletin, v. 105, 

no. 2, p. 296–304.
Greenland Halibut 613

25. Ianelli, J.N., Wilderbuer, T.K., and Nichol, Dan, 2007, Assessment of Greenland turbot in the eastern Bering Sea and 

Aleutian Islands: North Pacific Fisheries Management Council, Bering Sea and Aleutian Islands SAFE Document, 

p. 513–562, accessed April 6, 2014, at http://www.afsc.noaa.gov/refm/docs/2007/BSAIturbot.pdf.

26. Acuna, E.I., Goddard, P., and Kotwicki, S., 2003, 2002 bottom trawl survey of the eastern Bering Sea continental shelf: 

National Marine Fisheries Service, Alaska Fisheries Science Center, Processed Report 

2003-01.

27. Hoff, G.R., and Britt, L.L., 2005, Results of the 2004 Eastern Bering Sea upper continental slope survey of groundfish and 

invertebrate resources: National Oceanic and Atmospheric Administration Technical Memorandum, 300 p. 

28. Gregg, J.L., Anderi, D.M., and Kimura, D.K., 2006, Improving the precision of otolith-based age estimates for Greenland 

halibut (Reinhardtius hippoglossoides) with preparation methods adapted for fragile sagittae: Fishery Bulletin, v. 104, no. 4, 

p. 643–648.

29. Treble, M.A., Campana, S.E., Wastle, R.J., Jones, C.M., and Boje, J., 2008, Growth analysis and age validation of a 

deepwater Arctic fish, the Greenland halibut (Reinhardtius hippoglossoides): Canadian Journal of Fisheries and Aquatic 

Sciences, v. 65, no. 6, p. 1,047–1,059.

30. Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, K., de Lange Wenneck, T., Høines, Å., Bjelland, O., Aglen, 

A., Prokhorova, T., Murashko, P., Prozorkevich, D., Konstantin, Byrkjedal, I., Langhelle Drevetnyak, and G., Smirnov, O., 

2011, Atlas of the Barents Sea fishes: IMR/PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

31. Shuntov, V.P., 1970, Seasonal distribution of black and arrow-toothed halibuts in the Bering Sea: in Moiseev, P.A., ed., 

Soviet fisheries investigations in the northeastern Pacific: Jerusalem, Israel Program for Scientific Translations 1968, 

p. 397–408.

32. Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution in the Kronitsky Bay in relation to abiotic factors: Zoologicheskii 

Zhurnal, v. 63, p. 1,363–1,373.

33. Mueter, F.J., University of Alaska-Fairbanks, written commun., 2010.

34. Vollen, T., and Albert, O.T., 2008, Pelagic behavior of adult Greenland halibut (Reinhardtius hippoglossoides): Fishery 

Bulletin, v. 106, no. 4, p. 457–470.

35. Kim, S.T., and Biryukov, I.A., 1998, Distribution and some features of the biology of Reinhardtius hippoglossoides 

matsuurae (Pleuronectidae) and Sebastolobus macrochir (Scorpaenidae) from the eastern coast of Sakhalin: Journal of 

Ichthyology, v. 38, no. 1, p. 143–146.

36. Zenger, H.H., Jr., 2004, Data report—2002 Aleutian Islands bottom trawl survey: U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS-AFSC-143, 247 p.

37. Junquera, S., Román, E., Morgan, J., Sainza, M., and Ramilo, G., 2003, Time scale of ovarian maturation in Greenland 

halibut (Reinhardtius hippoglossoides, Walbaum): ICES Journal of Marine Science, v. 60, no. 4, 

p. 767–773.

38. Albert, O.T., Harbitz, A., and Høines, Å.S., 2003, Greenland halibut observed by video in front of survey trawl—Behaviour, 

escapement, and spatial pattern: Journal of Sea Research, v. 50, no. 2–3, p. 117–127.

39. Dunn, J.R., and Sample, T.M., 1976, Greenland turbot (family Pleuronectidae) in Pereya, W.T., Reeves, J.E., and Bakkala, 

R.G., eds., Demersal fish and shellfish resources of the eastern Bering Sea in the baseline year 1975: Seattle, Washington, 

National Marine Fisheries Service, Northwest and Alaska Fisheries Science Center, Processed Report, p. 475–487.

40. Cooper, D.W., National Marine Fisheries Service, written commun., 2010.

41. D’yakov, Y.P., 1982, The fecundity of the Greenland halibut, Reinhardtius hippoglossoides (Pleuronectidae), from the 

Bering Sea: Journal of Ichthyology, v. 22, no. 5, p. 59–64.

42. Kihara, K., and Shimada, A.M., 1988, Prey-predator interactions of Pacific cod Gadus macrocephalus and water 

temperature: Nippon Suisan Gakkaishi, v. 54, no. 12, p. 2,085–2,088.
614 Alaska Arctic Marine Fish Ecology Catalog

43. Yang, M.-S., 1996, Diets of the important groundfishes in the Aleutian Islands in summer 1991: U.S. Department of 

Commerce, National Oceanic and Atmospheric Administration Technical Memorandum NMFS-AFSC-60, Alaska Fisheries 

Science Center, 105 p.

44. Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology of groundfishes in the eastern Bering Sea, with emphasis on 

food habits, in Loughlin, T.R., and Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, University of Alaska Sea Grant, 

p. 537–580.

45. McLaren, I.A., 1958, The biology of the ringed seal (Phoca hispida Schreber) in the Eastern Canadian Arctic: Bulletin 

Fisheries Research Board of Canada 118, 97 p.

46. Finley, K.J., and Gibb, E.J., 1982, Summer diet of the narwhal (Monodon monoceros) in Pond Inlet, northern Baffin Island: 

Canadian Journal of Zoology, v. 60, no. 12, p. 3,353–3,363.

47. Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 1985, A natural history of Digges Sound: Report Series Canadian 

Wildlife Service no. 46, Canadian Government Publishing Centre, Canada Communication Group, 63 p.

48. Frost, K.J., and Lowry, L.F., 1980, Feeding of ribbon seals (Phoca fasciata) in the Bering Sea in spring: Canadian Journal 

of Zoology, no. 58, v. 9, p. 1,601–1,607.

49. Harry, G.Y., and Hartley, J.R., 1981, Northern fur seals in the Bering Sea, in Hood, D.W., and Calder, J.A., eds., The Eastern 

Bering Sea Shelf: Oceanography and Resources: Seattle, University of Washington Press, 

p. 847–867.

50. Lang, G.M., Livingston, P.A., Pacunski, R.E., Parkhurst, J., and Yang, M.-S., 1991, Groundfish food habits and predation on 

commercially important prey species in the eastern Bering Sea from 1984 to 1986: U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration Technical Memorandum NMFS F/NWC-207.

51. Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 2001: U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration Technical Memorandum NMFS-AFSC-158, 249 p.

52. Froese, R., and Pauly, D., eds., 2012, FishBase—Global information system on fishes: FishBase database, accessed July 8, 

2012, at http://www.fishbase.org.
Chapter 3 615

Acknowledgments

We wish to acknowledge the technical assistance of 

other project personnel who contributed to the gathering of 

data and information and preparation of maps, graphs, and 

figures in this report. Among those from the U.S. Geological 

Survey’s Western Fisheries Research Center, we thank 

Angela Lind-Null, Michael Hayes, and Steve Rubin for their 

early work in reviews of age-and-growth information. From 

University of California, Santa Barbara, we express our 

special gratitude to Rebecca Bloom and Eddy Peralta for their 

efforts in generating vertical distribution charts for the Arctic 

marine fish fauna. Many individuals are acknowledged for 

their reviews of early drafts of this report. Included in this 

group we appreciate the reviews and comments of Jim Reist 

(Department of Fisheries and Oceans, Canada), Robert Meyer 

(formerly USGS), Larry Moulton (MJM Research LLC), 

David Brown (U.S. Fish and Wildlife Service), Ben Gray 

(University of Alaska Fairbanks), Jefferson Childs (formerly 

BOEM), Nancy Deschu (BOEM), Catherine Coon (BOEM), 

Katherine Wedemeyer (BOEM), Louis Fortier (Universite ́ 

Laval, Quebec, Canada), and Alec Maule (formerly USGS). 

Summary

The species accounts are a major biodiversity milestone 

confirming the presence of 109 marine fishes in 24 families 

in the U.S. Chukchi and Beaufort Seas. One hundred four 

(104) species accounts are presented. Full species accounts 

for the Alaska Skate, Pacific Sleeper Shark, Thorny Sculpin, 

Checkered Wolf Eel, and Pale Eelpout were not prepared 

due to their very recent confirmation and general lack of 

information from the region. Each account describes what 

is presently known about the biology and ecology of each 

species from observations from Alaska, adjacent seas, and 

other locations. A species account summarizes current 

information about geographic distribution, abundance, life 

history and habitats, community relationships and population 

ecology, and ecological roles. The listing of Iñupiaq names 

for some species, but not others, provides an important 

measure of traditional values of these fish. Each species 

account represents an autonomous review of relevant data 

and information; each is scientifically documented with 

key literature sources so that users will have easy access to 

additional detail from the studies reviewed. Collectively, 

this treatment is the most comprehensive inventory of 

species information undertaken for this segment of Alaska’s 

marine fish fauna to date. The species accounts present 

thorough information, some of which reflects quantitative 

depictions of complex ecological concepts and population 

understanding. Information about life cycle requirements, 

trophic position, and population resilience are essential to 

National Environmental Policy Act assessments of potential 

effects from offshore energy development at population and 

ecosystem levels of biological organization.
616 Alaska Arctic Marine Fish Ecology Catalog

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Abstract

The compilation of data and information, its review, 

and the synthesis processes leading to the development of 

individual species accounts focused on descriptive elements 

of Arctic Alaska’s marine fish fauna. The species accounts 

reflect the compilation and review of a large body scientific 

information about the marine fishes off Alaska in United 

States waters in the northern Bering Sea, north of the Bering 

Strait, and throughout the circumpolar Arctic. The purpose 

of this synthesis is to interpret the whole of this biological 

and ecological information in the context of the marine 

biogeography of the Chukchi and Beaufort Seas. This 

interpretive approach provides historical and contemporary 

perspectives to our descriptions of (1) patterns of species 

occurrence, habitat and population relationships, and 

(2) functional ecosystem processes that affect the distribution 

and abundance of marine fishes, and, with respect to the 

present, (3) conceptual understanding and information needed 

for resource management and conservation. The objectives 

of this synthesis focus on environmental relationships 

including physio-chemical factors (for example, temperature, 

salinity, dissolved oxygen, and turbidity), mechanisms 

(for example, currents, migrations, and movements), and 

biological processes (for example, competition, predation, 

colonization, and reproduction) that collectively have limited 

the distribution, abundance, and productivity of marine fish 

populations through adaptations to Arctic conditions.

Introduction—A Biogeographic 

Emphasis 

Individual species accounts present a large amount of 

information about the biology, geography, and environmental 

factors affecting large-scale patterns of distribution and 

abundance. Collectively, this compilation of environmental 

information is foundational for an improved understanding 

of the descriptive and interpretive components of this 

region’s biogeography (Nelson, 2006). Our focus on the 

interpretive component (historical and ecological elements 

of biogeography) provides a meaningful approach for 

explaining the observed diversity relationships, synthesizing 

the information presented, and expressing our impressions 

about outstanding needs. Historical biogeography addresses 

the origins of distributional patterns as determined from 

systematic studies. The paleoceanographic record is not 

well-developed from this part of the Arctic and here 

we explore species origins in light of possible dispersal 

and vicariance events as suggested by paleontology and 

geologic records, climatic histories, and known phylogenetic 

relationships. Ecological biogeography addresses the 

environmental relationships including physiochemical factors 

(for example, temperature, salinity, dissolved oxygen, and 

turbidity), mechanisms (for example, currents, migrations, 

and movements), and biological processes (for example, 

competition, predation, colonization, and reproduction) that 

limit the distribution, relative abundance, and productivity of 

a species. Information about body size and trophic position 

also are reviewed because they are important parameters that 

correlate with metabolic processes and other life history traits 

(Romanuk and others, 2011). 

The descriptive component of biogeography is addressed 

in project tasks leading to an updated checklist of marine 

fishes, presentation of information in the species accounts, 

and special analyses related to (1) new species occurrences 

and range extensions, (2) large-scale patterns of abundance 

from past and ongoing studies, and (3) identification of species 

likely to occur, but yet-to-be confirmed from the Chukchi 

and Beaufort Seas. The visualization of species information 

in new maps and depth profiles provides another dimension 

to evaluation of the status of baseline information than was 

previously available. This is especially relevant as it applies to 

availability of age- or stage-specific information and related 

habitat relationships. Quantitative relationships between 

age-and-length and size-at-length have been described for 

several dominant species, but this information is dated. For 

almost all species, information about stock structure and 

population dynamics (that is, differential mortalities between 

life stages or age classes) remains to be described. Although 

many science gaps exist, compiling and integrating descriptive 

and interpretive information-types allows a more detailed 

examination of linkages between geographic distributions, 

evolutionary processes (patterns of life history variations), 

ecological factors, genetic diversity, and population dynamics 

Chapter 4. Synthesis of Arctic Alaska Marine Fish Ecology

By Lyman K. Thorsteinson1 and Milton S. Love2

1

U.S. Geological Survey.

2

University of California, Santa Barbara.
618 Alaska Arctic Marine Fish Ecology Catalog

for the fauna as an initial approach to estimating effects of 

environmental change (Winemiller, 2004; Benton and others, 

2006). This approach illustrates why the information presented 

in the species accounts is relevant to decision making and, for 

potential indicators species, a rationale for research priorities.

Historical Biogeography

The Chukchi and Beaufort Seas occupy a relatively small 

region within the zoogeographic realm (Mecklenburg and 

others, 2011) that has been used by classical ichthyologists 

to describe the taxonomy and composition of the Arctic 

marine fishes (Briggs, 1974; Andriashev and Chernova 1994; 

Eastman, 1997; and Mecklenburg and others, 2011). In Alaska, 

the Arctic Realm includes the northern Bering Sea and marine 

environments beyond the EEZ3. As such, it does not directly 

correspond to conventional natural resource management areas 

in the Alaskan Arctic. The southern boundary of the province 

in the Bering Sea is ecologically significant because the 

area demarks a sharp gradient in diversity and abundance of 

Arctic and Boreal marine fishes that is defined by temperature 

(<2 °C). As such, a natural rather than administrative faunal 

separation is represented. For example, NOAA collected 86 

species of marine fishes in the northern Bering Sea in 1981 

(Thorsteinson and others, 1984) compared to the 300 species 

reported from the southeastern Bering Sea (Wilimovsky, 

1974). The zonation is important to contemporary Arctic 

issues with respect to extralimital species and northerly 

shifts in distributions and range expansions associated with 

climate change and potential fisheries. The importance of the 

Bering Strait with respect to geologic and climatic history 

and origins and exchanges of fishes, especially during the late 

Pleistocene, has been shown to be critical to understanding 

Pacific influences on Arctic fauna (for example, Mecklenburg 

and others, 2011). 

A systematic comparison of the marine fishes reported 

from the Arctic province, marginal seas of the Arctic Ocean, 

and Bering Sea provides clues and insights about the 

evolutionary processes (origins, rates of endemism) underlying 

current patterns of taxonomic representation (zoogeographic 

patterns, chapter 2). The comparison of marine fish faunas 

from adjacent waters also is instructive with respect to origins 

(distribution centers), expectation of probable occurrence, 

or in some instances, understanding founding sources of 

confirmed species from the U.S. Arctic (table 4.1, additional 

Arctic seas in Christiansen and others, 2013). 

The estimates of marine fish diversity (occurrence) in 

table 4.1, was from published literature or acquired through 

reliable sources of written communications, and represent 

“working” totals from a field of ichthyology that is rapidly 

changing. These changes relate to new discoveries, increased 

sampling, and new genetic tools for identification and better 

understanding of evolutionary relationships. The presence 

of 15 new marine fishes from the U.S. Arctic not reported 

in the Fishes of Alaska (Mecklenburg and others, 2002) 

has been confirmed for this report (table 2.2). Many of the 

authors cited reported significant problems associated with 

species identifications (related these to unresolved taxonomic 

issues), deficiencies in geographic sampling coverage, 

or access to existing data and information. Despite these 

issues, the systematic comparisons help to begin to explain 

large-scale patterns of biodiversity including origins of 

species distributions.

Biogeographic research has shown that fish species in 

the Pacific and Atlantic Oceans have similar zoogeographic 

patterns with respect to latitudinal gradients, but not diversity 

(Christiansen and others, 2008; Christiansen and others, 

2013). In each ocean, species richness peaks in the tropics 

with sharp gradients between tropical and temperate waters. 

There is leveling off in decline of richness toward the poles 

(Roy and others, 1998). Using marine gastropod diversity 

data, Roy and others (1998) examined latitudinal gradients 

in the Atlantic Ocean with respect to ecological traits (that 

is, range size, habitable area, and input of solar energy). The 

greatest correlation reported was between diversity and sea 

surface temperature (a proxy for solar input) suggesting that, 

if the relation was causal, it probably was linked through some 

aspect of production. Because the physical mechanisms of 

dispersal for marine invertebrates and fishes are similar (or 

the same), process effects—such as production cycles and 

events—would similarly influence geographic distribution and 

abundance patterns.

In areas where regional faunas are relatively well known, 

Briggs and Bowen (2012) described a high concordance 

between levels of endemism in fishes, molluscs, and other 

biota. However, Roy and others (1998) could not explain 

latitudinal gradients in the North Atlantic based on recent 

geologic history. By contrast, Vermeij (1991) and Briggs 

(1995, 2003) described the biogeographic consequences 

of the opening of the Bering Strait and the Great TransArctic Biotic Exchange (3.5 million years ago [Ma]) on the 

dispersal of Arctic molluscs and marine fishes, respectively 

(see table 4.2 for geologic periods). Prior to this opening, 

the marine barrier formed by Beringia had isolated Pacific 

and Atlantic faunas for millions of years. Vermeij (1991) 

determined that of the 295 mollusc species that participated in 

the interchange, almost 90 percent (n=261) had Pacific origins 

or were descended from taxa with Pacific origins. Briggs 

(2003) hypothesized an asymmetrical invasion of marine 

fishes from a North Pacific evolutionary center (“centres 

of origin” hypothesis) and that at least one species from 

3

The three provinces described in Sigler and others (2011) relate to Bering, 

Chukchi, and Beaufort Seas and relate more directly to smaller ecological 

units in a conservation context. For this discussion, the classical definition for 

a province is used based on endemism. In this case, the provincial boundaries 

correspond with the geography of shelf areas of the Arctic Region described 

by Briggs and Bowan (2012).
Chapter 4 619

Table 4.1. Systematic comparisons of marine fishes occurring in the Arctic marine and adjacent seas or marine areas bordering the 

U.S. Chukchi and Beaufort Seas.

[Regions: Arctic Realm (as defined in this chapter), Arctic Zoogeographic Region described by Mecklenburg and others (2011). OCS, Outer Continental Shelf; 

~, approximately; –, not applicable]

Regions Geography relative to 

U.S. study area

Estimated total number of 

References

Orders Families Genera Species

Arctic Realm Encompasses circumpolar Arctic 

including the northern Bering Sea 

22 45 126 242 Mecklenburg and others (2011)

U.S. Beaufort Sea Beaufort Sea OCS Planning Area 9 19 44 83 This report

U.S. Chukchi Sea Chukchi Sea OCS Planning Area 11 22 56 97 This report

Chukchi Borderland North U.S. Chukchi Sea 4 7 11 12 Mecklenburg and others (2014); 

Longshan and others (2014)

Canadian Beaufort Sea1 East of U.S. Beaufort Sea 21 48 115 189 Coad and Reist (2004)

Yukon Adjacent to U.S. Beaufort Sea and 

north of Yukon Territory, Canada

9 14 28 44–46 Coad and Reist (2004)

Russian Chukchi Sea West of U.S. Chukchi Sea 11 26 68 112 Parin (2004); Parin and others 

(2014); Datsky (2015)

East Siberian Sea Northern Russia east of Chukchi Sea 9 14 33 65 Pauley and Swartz (2007)

U.S. Bering Sea U.S. territorial marine waters south of 

Chukchi Sea

21 64 190 ~375 Mecklenburg and others (2002); 

Mecklenburg and others (2011 

OR1); Maslenikov and others 

(2013); Love and others, written 

commun. (2015)2

Northern Bering Sea Bering Sea from the Bering Strait to 

the south of St. Lawrence Island

– – – (a) 128

 (b) 136

(c) 165

(a) OBIS database (http:/www.

iobis.org/); 

(b) Chernova (2011); 

(c) Datsky and Andronov (2007)

Bering Sea Entire Bering Sea south of Chukchi 

Sea

– – – (a) 385

 (b) 418

(a) Christiansen and others (2013); 

Mecklenburg and Steinke 

(2015)

(b) Greenwald and others (2006)

1

This estimate does not include brackish water species.

2

Love and others, written commun. (2015) refers to an ongoing updating of Love and others (2005).

Salmonidae, Osmeridae, Hexagrammidae, Cottidae, Agonidae, 

Liparidae, Stichaeidae, and Pholidae contributed to the ArcticNorth Atlantic fauna. The Capelin (Mallotus catervarius) 

is an example of such a Trans-Arctic dispersal during the 

Pleistocene (Dodson and others, 2007). Similarly, the cod 

family, Gadidae, is thought to have developed in the North 

Atlantic and contributed two species to the North Pacific. 

Large segments of an ancient fauna of boreal origins 

are believed to have gone extinct during the late Miocene 

(Mecklenburg and others, 2011). The opening of the Bering 

Strait, prevailing ocean conditions and currents probably 

favored the dispersal of new North Pacific species into 

habitable Arctic shelf areas during the Pliocene (Eastman, 

1997; Briggs, 2003). Ocean connections to the Arctic during 

this period are hypothesized to have been primarily through 

the Fram (North Atlantic) and Bering (North Pacific) Straits. 

The relatively small number of species in the Arctic suggests 

that widespread dispersal processes were constrained by the 

separation of basins and absence of a circumpolar current. 

Christiansen and others (2013, p. 195) noted that (Arctic) 

freshwater and diadromous fishes were “significantly molded 

by glaciation, deglaciation and geological events during the 

late Pleistocene and Holocene epochs (i.e. ~ 126,000 and 

12,000 years ago, respectively)” and that (citing Krylov and 

others, 2008 and Polyak and others, 2010) the evolutionary 

history of the marine fish fauna “dates back to the Neogene 

period as the modern circulation in the Arctic Ocean began 

to form some 14–17 million years ago.” The Great Exchange 

occurred during a period of ice-free, boreal conditions. The 

greater success of Pacific invaders was hypothesized by 

Briggs (1995) to be related to their “superior competiveness” 

(for example, behavioral, reproductive rate, individual 
620 Alaska Arctic Marine Fish Ecology Catalog

size, or vulnerability to predators or parasites) resulting 

from their evolution in a highly diverse marine ecosystem. 

Extended cooling between 2.9 and 2.4 Ma is postulated to 

have led to the extinction of the truly boreal species and given 

rise to the modern Arctic fauna (table 4.2; see Eastman, 1997; 

Mecklenburg and others, 2002, 2011; Briggs, 1974, 1995, 

2003, 2004). This fauna is characterized by relatively few 

(numbers) endemic species when compared to the diversity 

of marine fish fauna from lower latitudes. The low amount of 

endemism has been hypothesized to be related to evolutionary 

effects of ocean continuity and lack of geographic isolation 

over recent time. Briggs (1995) indicated that the Arctic 

climate repeatedly warmed and cooled until about 3 Ma when 

present cold conditions stabilized.

In Alaska, the species richness of the marine fish fauna 

from the high Arctic (includes northern Bering Sea as used 

here) is similarly low when compared to lower latitudes 

(chapter 2). Interestingly, the number of endemic species 

relative to total number of marine fishes reported from 

the U.S. Chukchi and Beaufort Seas is relatively high. We 

propose that this relates to the geographic isolation of this 

region during the last glacial period resulting from a lack of 

connection between Pacific and Arctic oceans through the 

Bering Strait, and the absence of shelf environment between 

openings with the Nordic Seas. 

Table 4.2. Geologic and climatic history of the Arctic.

Era(s) Recent geologic history

(millions of years ago) Geologic and climatic significance

Late Cretaceous 80 Beringia (Bering Land Bridge) forms and separates North Pacific 

and Arctic-North Atlantic faunas.

Paleocene–Oligocene 66–23.03 The Arctic Region was in high latitude position and the climate 

was temperate with water temperatures of 10–15 °C. Water 

temperatures gradually cooled during the Eocene and ligocene 

eras (56–23.03 million years [Ma] ago)

Miocene 23.03–5.3 Arctic land masses reached their present positions and water 

temperatures dropped below freezing (10–15 Ma). The Bering 

Strait may have opened during the Cenozoic (6–12 Ma) 

allowing for limited passage (Briggs, 2003). Additional 

cooling, ice sheet expansion, lower sea levels, and shallow 

Bering Strait, limit exchanges between Arctic and Pacific 

Oceans. Ocean circulation was latitudinal with limited 

Arctic-North Atlantic exchanges. The southward movement 

of the warm-temperate zone and its replacement by the 

cold-temperate zone are hypothesized to have caused mass 

extinctions of boreal families during the late Miocene.

Pliocene 3.5–3 The opening of Bering Strait allowed many Pacific species to 

invade the Arctic. Ocean circulation changed with the closure 

of the Isthmus of Panama (about 3.2 Ma), which strengthened 

the Gulf Stream, increased precipitation at high latitudes, and 

contributed to further glaciation in the northern hemisphere. 

The Atlantic and Pacific Oceans had oceanic connections to 

the Arctic during the late Pliocene and faunal interchange was 

possible. 

Pleistocene 2.588–0.0117

(11,700 years ago)

The early Pleistocene was a time of glacial advances and retreats, 

periodic freshwater inputs into the Arctic Basin, and cooling 

leading to the formation of Arctic sea ice. Permanent ice cover 

has been present for at least 0.7 Ma and possibly intermittently 

as long as 2.0 Ma. Some components of the Arctic marine 

fauna may have persisted from the Miocene and Pliocene eras 

and exchanges with the Atlantic and Pacific were also possible 

as the Bering Strait remained open.
Chapter 4 621

Geographic Perspectives of Marine 

Fish Diversity

Mecklenburg and others (2011) examined the faunal 

composition of marine fishes in the Arctic region and 

confirmed earlier characterizations of the young phylogenetic 

age of dominant families (for example, Zoarcidae and 

Stichaeidae) and the apparent lack of successful invaders 

from the Atlantic (for example, Gadidae and Anarhichadidae). 

Data in this report support these findings as they extend to 

Alaska. As an example, the two largest families (Zoarcidae 

with 17 species and Cottidae with 17 species) were determined 

to contain almost one-third of the total number of species 

reported. They were followed, in order of decreasing species 

numbers, by Salmonidae (12), Pleuronectidae (9), Stichaeidae 

(8), Agonidae (7), and Liparidae (6). Collectively, seven 

(7) families accounted for nearly 68 percent of the region’s 

confirmed marine fish diversity. In the western Chukchi Sea, 

Datsky (2015) noted a similar pattern (without Salmonidae), 

with members of the Cottidae, Zoarcidae, Pleuronectidae, 

Stichaeidae, and Agonidae accounting for 66 percent 

(73 species) of the total number of species identified. 

Regional comparisons of the diversity of fishes reported 

from marine waters adjacent to the U.S. Arctic are qualitative4; 

however, the distributional patterns suggested by presenceabsence data suggest dispersal processes. With respect to the 

U.S. Arctic, the Bering Sea, with more than 400 marine fishes, 

represents the largest regional source of potential colonizing 

species. The same is true of the North Atlantic for the Barents 

and Eastern Siberian Seas. When the species information 

is examined in the context of prevailing ocean currents and 

transport processes, a northerly dispersal of species in Pacific 

Waters, notably in Alaska Coastal Water, seems to be the most 

probable (with respect to likelihood of success) historical and 

contemporary mechanism for large-scale colonization of the 

U.S. Chukchi and Beaufort Seas. The distribution of shared 

species in the Chukchi Sea (United States and Russian), 

Chukchi Borderlands, and southeastern Beaufort Sea (United 

States and Yukon) would seem to reinforce the Pacific 

transport hypothesis (table 4.2). However, Mecklenburg 

and others (2014) noted that all 12 species collected from 

the Chukchi Borderland occur, and many are common in 

northern Atlantic sectors of the Arctic. Their findings support 

an Atlantic transport hypothesis in subsurface currents 

associated with Atlantic water masses. Alternatively, as the 

authors suggest, the species could have been in the region for 

thousands of years but simply not discovered until recently 

because of the previous lack of sampling.

A regional comparison of the presence data indicates 

a high percentage of shared species between United States 

sectors of the Chukchi and Beaufort Seas (table 4.3). Although 

this could be an artifact of existing data collection, it supports 

a major dispersal pathway in Alaska Coastal Water. The 

regional data as a whole further support the existence of an 

inter-regional gradient of declining species richness with 

increasing latitude. This pattern was reported previously for 

the U.S. Arctic (Craig, 1984) and was related to the lack of 

expatriate species from the Bering Sea found in the Beaufort 

Sea (Crawford and others [2012] citing others). In this case, 

the transport of Alaska Coastal Water to the east and north 

of Point Barrow (fig. 1.1) may impede larval dispersal and 

colonization processes. Cold temperatures, availability of 

foods, and other physical and biotic prerequisites of the 

Beaufort Sea coastal habitats could be major constraints to 

successful colonization. 

4

Comparisons are problematic due to differences in sampling, problems 

associated with field identifications and nomenclature, and taxonomic 

uncertainties.

Table 4.3. Estimated numbers of common or shared species in the U.S Arctic and adjacent marine areas.

[Russian Chukchi Sea: Total estimate of 67 assumes Arctic Cod and Pacific Cod occur in the area, but were not reported by Pauly and Swartz (2007). Arctic:

Includes Chukchi and Beaufort Seas in waters of the United States]

Regions Chukchi 

Borderland

Russian

Chukchi Sea

United States

Yukon

 Chukchi Sea Beaufort Sea Arctic

Chukchi Borderland 12 4 7 9 7 3

Russian Chukchi Sea 67 46 42 39 22

U.S. Chukchi Sea 97 68 68 31

U.S. Beaufort Sea 83 68 34

U.S. Arctic 109 30

Yukon 44–46
622 Alaska Arctic Marine Fish Ecology Catalog

Understanding the diversity of marine fishes in adjacent 

areas also provides information about extralimital occurrences 

and a basis for reasoned expectations about probable species 

occurrences (not yet recorded) and possible founding sources 

of new species into Alaska waters (table 4.3). Mecklenburg 

and others (2011) reported the presence of 242 species of 

marine fishes in a circumpolar Arctic study area that, for 

Alaska, extends into the northern Bering Sea. Mecklenburg 

and Steinke (2015) report 122 marine fishes from the Pacific 

Marine Region, a smaller region that includes northern 

Bering, Chukchi, Beaufort, and East Siberian Seas. They do 

not include amphidromous species in their tally of marine 

fishes. The Center of Biodiversity reports the occurrence of 

418 species of marine fish in the Bering Sea (Greenwald and 

others, 2006) and although this number could not be verified, 

it is indicative of the high diversity of fishes found south of the 

Bering Strait. With respect to the U.S. EEZ in the Bering Sea, 

Mecklenburg and others (2002) reported about 350 species. 

As part an ongoing effort to update a checklist of marine and 

estuarine fishes in United States waters by Love and others 

(2005), the current estimate is between 375 and 400 species 

for this part of the Bering Sea.

An examination of regional oceanographic and 

zoogeographic patterns in light of known phylogenetic 

relations provides additional support for hypotheses regarding 

the ancestry of Arctic marine fishes. For the United States, the 

updated list of fishes is indicative of both widespread (Pacific 

and Atlantic) and endemic (Arctic Ocean) origins. The species 

derived from Atlantic and Pacific origins further suggest 

the role of ocean currents on dispersal and the hypothesized 

importance of the continuity of Arctic shelves to colonization 

(Eastman, 1997). To illustrate the concept, Carmack and 

Wassman (2006), estimated that the flow of Atlantic Water into 

the Arctic Ocean is, on average, more than five times larger 

than inflow of Pacific Water. This difference was considered to 

be responsible for the dominance of Atlantic-derived species 

over much of the northern Russian shelf. 

The analysis herein of zoogeographic patterns revealed 

that Atlantic-derived species account for less than 10 percent 

of the diversity (n=9 species) of marine fishes confirmed from 

United States waters. This includes species from Stichaeidae 

(Eumesogrammus praecisus, Leptoclinus maculatus, and 

Stichaeus punctatus), Gadidae (Gadus chalcogrammus and 

Gadus macrocephalus), Psychrolutidae (Cottunculus microps), 

Osmeridae (Hypomesus olidus), Myctophidae (Benthosema 

glaciale) and Pleuronectidae (Reinhardtius hippoglossoides). 

Interestingly, only two stichaeids (E. praecisus and S. 

punctatus) are found in both Chukchi and Beaufort Seas 

suggesting a discontinuous dispersal for members of this 

family. A slightly broader examination of the distribution of 

Atlantic-derived species that included marine areas adjacent to 

Alaska revealed four species from four families in the Russian 

Chukchi Sea (G. chalcogrammus, E. praecisus, L. maculatus, 

and R. hippoglossoides); five species from five families in 

the Chukchi Borderland (G. chalcogrammus, Artedellius 

atlanticus, C. microps, L. maculatus, and R. hippoglossoides); 

and one species from the Yukon (Ammodytes dubius) and the 

possibility of a longitudinal gradient along the shelf break.

Paleontological Connections

Knowledge of the paleoceanographic conditions and 

how these conditions affect the evolution of Arctic marine 

fishes is limited. It seems clear from the inventory of known 

marine fishes from U.S. Arctic waters that endemic species 

(that is, Arctic, predominantly Arctic and Arctic Boreal 

fauna) are significant components of Chukchi (58 percent) 

and Beaufort (>74 percent) assemblages. How the opening 

and closing of the Bering Strait, interglacials, and formation 

of shelf ecosystems affected dispersal and speciation 

processes and faunal distinctions must be related to observed 

differences in regional oceanography, dispersal corridors, 

and energy pathways (Barber and others, 1997; Carmack and 

Wassmann, 2006; Dunton and others, 2006; Cui and others, 

2009; Norcross and others, 2010). For example, Pacific 

water influences on the Chukchi Sea shelf are more direct 

than advection of regional water masses onto the Beaufort 

Sea shelf. The hydrography of the Beaufort Sea shelf is 

affected by winds, upwelling, and river inputs, and presents a 

variable environment for its fishery occupants (Carmack and 

Wassman, 2006). Biogeographic data are indicative of Pacific 

influences on recent (<3 Ma) colonization processes. Regional 

differences in shelf environments (topographic, bathymetric, 

hydrographic, hydrologic, currents, and biologic) correspond 

to broad patterns of observed diversity in adaptive strategies 

between the Chukchi (greater than numbers/abundance of 

marine species) and Beaufort (greater than numbers/abundance 

of amphidromous species) Seas. The latitudinal gradient that 

forms the southern boundary of the Arctic province is evident 

in the data presented. A less pronounced subregional gradient 

is near Point Barrow (Chukchi Sea greater than numbers of 

Bering Sea species less than Beaufort Sea).

Adaptations in Marine Arctic Fishes

Environmental conditions in the Chukchi and Beaufort 

Seas pose numerous substantial challenges to their fish 

populations. As noted by Power (1997, p. 16), “Fish living 

in arctic marine waters have to adapt to low temperature, in 

most places below the usual freezing point of teleost blood; 

seasonally constrained low productivity; perpetual or long 

periods of darkness; and an ice-affected shoreline.” In 

contrast to waters of Antarctica, Arctic waters may also have 

substantial annual changes in salinity (Eastman, 1997). The 

Arctic challenges extend to how, where, and when research 

has been conducted, and the resulting ecological information is 

much better for coastally occurring species and for those taken 

in subsistence fishing, than for truly marine fishes. As a result, 
Chapter 4 623

Arctic marine fishes are a current emphasis of BOEM research 

including physiological studies in laboratory and field studies 

(appendix A) as sea ice retreats and technological advances 

allow greater access to offshore marine environments for 

scientists and industry alike.

Many investigations show how large-scale oceanographic 

processes over coastal, shelf, and slope environments of the 

Arctic Ocean and marginal seas relate to observed differences 

in regional productivity and transfer of energy to benthic 

and pelagic components of the marine ecosystem. These 

affect the form (anatomical and morphological, for example, 

Mecklenburg and others, 2002) and function (ecological 

niche) of individual species and, collectively, the suite of life 

strategies for marine fishes occupying these waters (chapter 3). 

Arctic shelves are shallow and often are characterized by the 

richness and biomass of their benthos. With respect to the U.S. 

Arctic, Carey and Ruff (1977) and Carey (1987) hypothesized 

the predominance of benthic- and pelagic-dominated 

ecosystems in the Chukchi and Beaufort Seas, respectfully, as 

suggested by the distribution and abundance of infaunal and 

epifaunal invertebrates. Subsequent research (for example, 

Grebmeier and others, 2006a) related observed patterns of 

distribution to ocean features and to the cropping efficiency 

of herbivorous zooplankton and transport of organic matter 

to the benthos. The tight coupling (low efficiency) between 

pelagic and benthic ecosystems reported by Grebmeier and 

others (2006a) support the Carey (1987) hypothesis for the 

Chukchi Sea shelf, but the effects of shelf-slope exchange in 

deeper waters and the Beaufort Sea (Forrest and others, 2007) 

are far less clear. The low diversity and biomass of benthic 

invertebrates reported in ongoing benthic surveys (Katherine 

Wedemeyer, BOEM Alaska OCS Region, Anchorage, Alaska, 

oral commun., 2015) may lend further support to the pelagic 

hypothesis especially in eastern sectors of the U.S. Beaufort 

Sea. Coastal processes and food webs in the nearshore marine 

(≤10 m depths) are affected by terrestrial, freshwater, and 

marine influences. Marine exposures and influences generally 

are relatively greater along the U.S. Chukchi Sea coast than 

the Alaska Beaufort Sea.

The role of sea ice as a substrate for algal production 

and marine food webs represents a specialized flow of energy 

in the Arctic marine environment (Alexander, 1992). The 

composition, low relative abundance, and low trophic position 

of the marine fish assemblage reported from the Chukchi Sea 

are indicative of an invertebrate-dominated benthic ecosystem 

over shelf waters. The suspected occurrence of large 

concentrations of Arctic cod in deep waters of the Chukchi 

and Beaufort Seas (Crawford and others, 2012) suggests more 

pelagic processes may be occurring over the slope. Nearer 

shore, the role of terrestrial peat in coastal food webs of the 

Alaska Beaufort Sea represents a conceptually different but 

important source of organic matter in coastal food webs and 

in the adaptation, and interactions of marine fish in ephemeral 

brackish waters that annually form along the coast (Dunton 

and others, 2006). As new data from BOEM studies on the 

distribution and abundance of marine fish and invertebrates 

from offshore marine areas of the Chukchi and Beaufort Seas, 

including food habits and bioenergetics become available, 

greater insights into more specific Arctic adaptive strategies 

will be possible. 

Descriptions of the survival strategies of cold-water 

fishes in the Arctic environment are provided herein along 

with considerations of adaptations gleaned from known 

information about marine fishes living in other polar regions 

and, to a lesser extent, more temperate waters. However, there 

is a shortage of research on the physiology and behavior of 

the fishes of the Chukchi and Beaufort Seas. Although some 

studies were conducted on taxa like Arctic Cod and Shorthorn 

Sculpin in other parts of their ranges, most physiological 

research has been on Antarctic species from a region quite 

different from the Arctic. At high latitudes particularly, 

Antarctic waters have been covered in an ice sheet for 

10–15 million years. Its inshore waters have been isolated 

from lower latitudes for about 25 million years (leading to 

a suite of endemic and closely related species), and it has 

both temperatures and salinities that are quite stable over the 

course of a year (Eastman, 1997; DeVries and Steffensen, 

2005; Verde and others, 2006). Additionally, the water beneath 

this ice sheet is super-cooled, leading to the formation of ice 

crystals to depths of more than 30 m. In contrast, a permanent 

ice cover has existed in the Arctic for only 0.7–3 million years, 

so there are no barriers to immigration into the region (leading 

to a relatively diverse fish assemblage). Water temperatures 

can vary as much as 15 °C during a year, and at least in some 

nearshore environments, the variability in salinity is significant 

(Eastman, 1997; DeVries and Steffensen, 2005; Whiteley and 

others, 2006). There is no super-cooled water in the Arctic. 

As an example, over the course of a year, conditions for the 

nearshore fish assemblage in Simpson Lagoon, Beaufort Sea, 

included temperatures from at least –2 to 14 °C, freshwater to 

marine conditions, and clear to extremely turbid waters (Craig 

and Haldorson, 1981).

Because most of the research on physiological and 

behavioral adaptations of fishes to polar conditions has been 

conducted on Antarctic fishes, some of the results of this 

research should be cautiously extrapolated to Arctic taxa. As 

noted by Wells (2005, p. 302), “There is no a priori reason 

that their [Antarctic fishes] adaptations should parallel those 

seen in the Arctic fishes.”

Physical and Sensory Adaptations

Particularly in comparison to fish physiology and 

biochemistry, physical adaptations to Arctic environments, 

if any, are poorly understood. Seasonal changes in light and 

sea ice effect in concert with cold temperatures, geographic 

vagaries in productivity and coupling of marine pelagicbenthic ecosystems, and the general low abundance of 

marine fishes are evident in their physical structure and 
624 Alaska Arctic Marine Fish Ecology Catalog

functional morphology. These adaptations relate to how 

fishes interact and navigate Arctic marine waters in search 

for prey, potential mates, and optimum habitats. Examples 

of physical adaptations have been described with respect 

to sensory organs such as eyes, lateral lines, and barbels; 

skeletal and musculature structures; egg and larval conditions; 

and patterns of coloration. The physical placement of eyes 

and mouth, shape and size of mouth, and arrangement of 

teeth, reflect examples of morphological adaptations to 

dietary requirements and evolved predator-prey relationships 

(for example, piscivores, plankivores, benthivores, and 

generalists). Collectively, the adaptations relate to various 

search, capture, and ingestion strategies related to life (that is, 

moving, feeding, growing, maturing, and reproducing) in the 

Arctic marine environment.

Little is known about the direct selective effects of cold 

on sensory organs, although there has been some research 

on adaptations of the nervous system to lower temperatures 

(see section, “Physiological and Biochemical Adaptations”). 

Some work has examined the role that low light levels (either 

seasonal or during summer under ice cover) might have 

on sensory system function. McAllister (1975) discussed a 

number of potential sensory adaptations to low light levels in 

Arctic fishes. He compared eye diameters between five pairs 

of closely related fishes— one pair from the genera Eleginus, 

Gadus, Triglops, Aspidophoroides, and Liopsetta). In each 

comparison, one species of the pair was taken from the Arctic; 

the other species were from boreal or subarctic waters. In 

each case, the northern representative had larger eyes. He also 

noted that Arctic-caught Pacific Cod (Gadus microcephalus; 

identified as Greenland Cod, G. ogac) and Arctic-caught 

Atlantic Cod (G. morhua) had barbels that were longer (when 

standardized against orbit length) than those of Atlantic Cod 

taken in boreal waters. Similarly, the barbels of the Arcticdwelling Arctic Alligatorfish (Aspidophoroides olrikii) are 

prominent, whereas those of the congeneric and, arguably 

more boreal-dwelling Alligatorfish are small. McAllister also 

generalized that the lateral line systems of a number of Arctic 

gadids and sculpins are composed of exposed filaments rather 

than filaments inside of canals, a trait found in other members 

of these families that lived in more southerly waters.

Since McAllister’s (1975) review, no research has been 

done in Arctic waters that attempted to verify his hypothesis. 

Work on Antarctic fishes has shown that there are some, 

but not extreme, adaptations to low-light environments 

in visual, lateral line, chemosensory, and tactile systems. 

For instance, and apparently similar to some Arctic fishes, 

many nototheniids have relatively large numbers of free 

neuromasts, perhaps in response to feeding and avoiding 

predation at low light levels (Montgomery and Sutherland, 

1997; Macdonald and Montgomery 2005). However, it may be 

difficult to assign the reason for the evolution of this trait. As 

an example, Coombs and Montgomery (1994) examined the 

frequent occurrence of free neuromasts on the ventral trunks 

of Antarctic notothenioid fishes. They argued that this may 

not be a response to living in low light levels. Rather, it may 

be a non-adaptive, paedomorphic trait, essentially a holdover 

from larval structures. However, it is not clear whether a 

midwater lifestyle or a low-light environment selected for 

this morphology. McAllister also made the generalization 

that, with few exceptions, Arctic fishes tend to be drably 

colored and suggested that the relatively low light levels 

of that environment might favor species that blend in with 

that environment.

Numerous attempts have been made to link increasing 

vertebral number, and its possible ramifications, with 

adaptation to Arctic conditions. This was first presented by 

Jordan (1891) as “in certain groups of fishes the northern or 

cold-water representatives have a larger number of vertebrae 

than those members which are found in tropical regions” and 

is now called “Jordan’s Rule.”(McDowall, 2008, p. 502). 

Nikolsky (1963), McAllister (1975), and other subsequent 

authors have speculated that greater numbers of vertebrae 

would be selected for in cold-water species. They noted that 

the additional myomeres associated with those vertebrae 

would aid in swimming through more viscous cold waters 

and help overcome a putative lowered efficiency of colder 

muscles. However, McDowall (2008) questions whether 

“Jordan’s Rule,” as well as several other generalities (that is, 

“Bergmann’s Rule,” “Lindsey’s Rule,” and “Williston’s Rule”) 

regarding the interrelationships among water temperature, 

size of individuals, and number of vertebrae, have any 

evolutionary significance. McDowall’s (2008) position is that 

(1) there are many exceptions to these “rules,” (2) it remains 

unclear what drives these relationships in an adaptive sense, 

and (3) there is a lack of understanding regarding whether 

variation in vertebrae number is under genetic and (or) 

environmental control.

An additional generalization, summarized in varying 

degrees in Rass (1941, 1989) and Marshall (1953) is that 

species living in Arctic waters have larger and fewer eggs 

(with larger yolks) than do those living in boreal waters. 

Marshall (1953) suggests that larger and yolkier eggs may be 

most adaptive in polar waters, where prey may be available 

only over a short plankton season. The advantages of the 

larger larvae that emerge from these eggs are “the smaller 

food requirements in relation to size combined with increased 

powers of swimming, leading to a widening of the range of 

search for suitable food...Hatching at an advanced stage with 

a shortening of the period leading to metamorphosis is likely 

to be advantageous in high arctic [sic] and Antarctic waters 

with a short seasonal plankton production...an increased 

supply of yolk leads to increased rates of growth of embryos 

and larvae, thus counteracting the retarding influence of low 

temperatures...” (Marshall, 1953, p. 340). Since that time, only 

a few studies have examined this phenomenon. Miller and 

others (1991) determined that among 13 species of flatfishes 

in the Atlantic, the 3 species in Arctic waters had the largest 

eggs. The operative factor(s) selecting for this trend remain 

somewhat unclear. Rass (1941) considered water temperature 
Chapter 4 625

alone responsible, whereas Marshall (1953) considered both 

physical factors (such as temperature) and biological factors 

(such as a short planktonic production season) important. The 

results of several studies imply that, at least in some groups, 

larger eggs are attributable to differences in spawning seasons 

rather than directly to temperature clines (Chambers, 1997). 

Lastly, we note the observations of Miller and Kendall 

(2009, p. 439): “It’s interesting to note that on the Arctic and 

Antarctic shelves, fish eggs are mainly demersal. A review 

of maternal output in polar fishes (Christiansen et al. 1998) 

revealed that while there was a pattern of relatively few and 

large eggs in polar versus boreal fishes this difference did not 

exist in substrate spawning species with large yolk reserves 

and long incubation periods (e.g., Greenland cod [Gadus 

ogac] and arctic flounder [Liopsetta glacialis], whereas 

most other cods and right eye flounders [Pleuronectidae] 

have pelagic eggs. It’s possible that spawning demersal 

eggs may protect the eggs against the risk of freezing and/

or also against the low salinity of the surface water (where 

osmoregulation is difficult) during the melting of the ice.” 

Physiological and Biochemical Adaptations

Adaptations related to living at low temperatures 

generally relate to the “scope for growth” bioenergetics 

processes that occur at molecular and cellular levels of fish 

physiology. Much of the available research has focused 

on antifreeze proteins (AFPs) in fishes from Polar regions. 

These proteins and other properties of blood chemistry are 

important adaptations to how Arctic marine fishes may avoid 

freezing or respond to below freezing conditions (for example, 

specializations in the nervous system).

Below freezing conditions represent a significant 

environmental threat to fish species living in polar regions, 

as the blood plasma of most marine teleosts freezes at about 

-0.7 to -0.9 °C (Holmes and Donaldson, 1969), well warmer 

than the freezing point of sea water (that is, -1.9 °C). In 

response, many fish species that inhabit Arctic waters have 

evolved the ability to lower the freezing point of their body 

fluids. Of particular importance to a fish living in subfreezing 

waters is the ability to prevent the ice crystals that occur in 

these waters from growing larger, either on that fish’s external 

surface or, after ingestion, within the body (DeVries and 

Cheng 2005; Cheng and others, 2006). 

The freezing point of aqueous solutions, such as those in 

fishes, can be depressed through elevating the concentrations 

of dissolved solutes. These solutes include such plasma 

electrolytes as sodium (Na+

) and chloride (Cl–)(O’Grady 

and DeVries, 1982). However, high concentrations of these 

ions can interfere with a range of biological processes, such 

as enzyme activity, protein subunit assembly, and protein 

solubility (Somero, 1992). Fishes also depress their freezing 

points with a number of organic compounds that have less 

disruptive effect on these processes (Raymond, 1997). These 

compounds include glycerol (Raymond, 1992; Driedzic and 

Short, 2007), trimethyamine oxide (Raymond, 1994; Raymond 

and DeVries, 1998), urea (Raymond, 1994; Raymond and 

DeVries, 1998), and glucose (Benziger and Umminger, 

1973). All of these ions and compounds depress the freezing 

point colligatively (acting through the number of ions or 

molecules in a given amount of solvent and not on their size 

or mass). Serum-ion concentrations in marine teleosts living 

in very cold waters generally are relatively high, typically, 

40 percent higher than in fishes from more temperate waters 

(Raymond, 1997).

The concentrations of dissolved solutes in many coldwater dwelling marine fishes (that is, Shorthorn Sculpin, Hew 

and others [1980]; Sea Raven [Hemitripterus americanus],

Fletcher and others [1984]; Atlantic Wolffish [Anarhichas 

lupus], Desjardins and others [2006]; see also O’Grady and 

DeVries [1982]) increase during the winter, probably in 

response to decreasing temperature and photoperiod (Driedzic 

and others, 2006). Dissolved solute concentrations may 

provide sufficient or near-sufficient antifreeze protection 

for only a few species (for example, Navaga [Eleginus 

navaga], Christiansen and others [1995] and Arctic Smelt, 

Raymond [1992]), whereas for all others the amount of 

freezing protection afforded is likely limited (Fletcher and 

others, 1985b). 

All cold-water dwelling fish species exposed to ice 

protect themselves from freezing by synthesizing AFPs. There 

are five distinct molecular groups of antifreeze proteins (AFP 

Types I–IV and antifreeze glycoproteins (AFGPs) (Fletcher 

and others, 2001); for the purposes herein all antifreeze 

proteins are referred to as AFPs. The five AFP types (among 

only distantly related taxa) are sufficiently structurally 

different from each other as to have evolved independently 

(Davies and others, 1988; Fletcher and others, 2001). The 

evolution of AFPs in Northern Hemisphere Arctic teleost 

fishes (table 4.4) likely occurred within the last 3 million 

years. This was the first time that these waters were glaciated 

(ice crystals formed) since the last ice age about 280 million 

years ago before the rise of teleosts (see Kennett, 1982; Scott 

and others, 1986; Fletcher and others, 2001; Davies and 

others, 1988).

Each type of AFP likely is coded for by a gene family 

(Hew and others, 1988; Scott and others, 1988; Shears and 

others, 1993; Desjardins and others, 2006). For instance, the 

number of gene copies of Type III AFP in the Atlantic Wolfish 

and Ocean Pout (Zoarces americanus) is estimated to be 85 

and 150, respectively (Desjardins and others, 2006). Although 

the adult fish ability to inhibit freezing in the presence of ice 

through the production of AFPs is well documented, there 

is growing evidence that this ability also is in embryos and 

larvae of some species (Murray and others, 2002, 2003; 

Robles and others, 2005; Desjardins and others, 2007) and 

perhaps in their eggs (Desjardins and others, 2007).
626 Alaska Arctic Marine Fish Ecology Catalog

AFPs may be produced in several parts of the body. 

In many species, AFPs are primarily (or solely) produced 

in the liver (Davies and others, 1988). However, in at least 

some species (for example, Shorthorn Sculpin, Longhorn 

Sculpin (Myoxocephalus octodemspinosus), Winter Flounder 

(Pseudopleuronectes americanus), and Ocean Pout) AFPs are 

produced in the liver and in the epidermis of structures such 

as the gills, skin, scales, and fins (Gong and others, 1992; 

Fletcher and others, 2001; Low and others, 2002; Murray 

and others, 2002, 2003). The exocrine pancreas appears to be 

the sole site of AFP production in all Antarctic notothenioid 

fishes (Cheng and others, 2006); such pancreatic production 

has not yet been detected in Arctic taxa. Praebel and Ramlov 

(2005) determined that, in some species, AFPs also are 

derived from food sources. They examined levels of AFPs in 

the gastrointestinal tracts of Arctic Cod (Boreogadus saida) 

feeding either on AFP-containing Arctic Cod or on crustaceans 

that did not produce AFPs. They determined that Ice Cod 

(Arctogadus glacialis) that had fed on Arctic Cod contained 

significantly higher levels of AFPs in their digestive fluids. 

Praebel and Ramlev (2005) hypothesized that partly digested, 

but still functioning, AFPs could be absorbed into the bodies 

of Arctic Cod.

AFPs do not prevent the creation of ice crystals within 

the fish; rather they retard the growth of ice crystals that have 

entered the fish (Raymond and DeVries, 1977; Evans and 

others, 2011). Each AFP molecule binds preferentially (and 

likely permanently) to well-defined sites on newly formed ice 

crystals (rather than to the surrounding water), thus altering 

their structure and inhibiting their growth (Fletcher, 1977; 

Fletcher and others, 2001; Davies and others, 2002; Goddard 

and Fletcher, 2002; Howard and others, 2010). The different 

Table 4.4. Species known to have antifreeze-protein or antifreeze-glycoprotein activity in the U.S. Chukchi–Beaufort Sea 

region. 

[All studies listed used fishes taken outside the U.S. Chukchi–Beaufort Sea region. Families are arranged in phylogenetic order (Nelson and 

others, 2004)]

Family Common name Source

Clupeidae Pacific Herring1 Power (1997)

Gadidae Arctic Cod Denstad and others (1987); Enevoldsen and 

others (2003); Nahrgang and others (2010)

Saffron Cod Raymond and others (1975)

Pacific Cod2 Van Voorhies and others (1978); 

Enevoldsen and others (2003)

Osmeridae Arctic Smelt Raymond (1992)

Cottidae Atlantic Hookear Sculpin3 Enevoldsen and others (2003)

Arctic Staghorn Sculpin Enevoldsen and others (2003)

Arctic Sculpin Enevoldsen and others (2003)

Shorthorn Sculpin4 Raymond and others (1975); Hew and 

others (1980); Denstad and others (1987); 

Enevoldsen and others (2003)

Liparidae Kelp Snailfish1 Enevoldsen and others (2003)

Stichaeidae Arctic Shanny Enevoldsen and others (2003)

Zoarcidae Polar Eelpout5 Davenport (1992)

Pholidae Banded Gunnel Enevoldsen and others (2003)

Pleuronectidae Alaska Plaice6 DeVries (1980)

1

Enevoldsen and others (2003) reported antifreeze proteins (AFPs) from “Pygmy Snailfish” now known as “Kelp Snailfish.”

2

As Greenland Cod, Gadus ogac.

3

Scientific name for Atlantic Hookear Sculpin is Artediellus atlanticus.

4

Raymond and others (1975), reported AFPs from “Myoxocephalus verrucosus” from Saint Lawrence Island, Bering Sea. This is a synonym 

of M. scorpius (Mecklenburg and others, 2002).

5

Enevoldsen and others (2003) reported AFPs from “Canadian Eelpout” now known as “Polar Eelpout.”

6

DeVries (1980) reported AFPs from “Arctic Plaice” now known as “Alaska Plaice.”
Chapter 4 627

types of AFP molecules bind to different surface planes on 

an ice crystal (Knight and others, 1991; Fletcher and others, 

2001). In at least some species, “chaperone molecules,” 

such as glycerol, enhance AFP activity (Gong and others, 

2011). Compared to dissolved solutes, AFP molecules act 

noncolligatively; that is they have little effect on the osmotic 

balance within a fish (because the action of AFP molecules is 

not proportional to the number of molecules present).

Preventing the buildup of ice crystals is a two-step 

process involving preventing ice crystals in the environment 

from entering the body through the skin and preventing those 

crystals that do enter the body from growing larger. For fishes 

encountering freezing temperatures and ice, the first barrier is 

through prevention of ice crystals entering the body through 

the epithelium. Fishes living in Antarctic waters (few studies 

have targeted Arctic fishes) have ice on their skin, gills, and in 

their gastrointestinal tract, but not in the body fluids, muscles, 

and most organ systems, except for the spleen (Praebel and 

others, 2009; Evans and others, 2011). Fish skin appears to 

be an effective barrier to ice crystal penetration into the fish, 

with gill epithelium perhaps less so (Valerio and others, 1992). 

How do fish prevent ice crystals that form on the mucous 

covering the skin (Praebel and others, 2009) from gaining 

access to the body interior? Although it is possible that larger 

ice crystals fall off through frictional forces as the fish moves 

through the water or as currents pass over the body, it is more 

likely that these ice crystals are prevented from growing by the 

AFPs in the mucous covering the skin. Ice crystals also have 

been found in the gastrointestinal tract, which were carried 

along with ingested food and as fishes drink seawater (to 

maintain osmotic balance). Here, these crystals are prevented 

from enlarging by AFPs in the esophagus and stomach as 

well as pancreatic secretion into the intestine where the fluid 

becomes hyposmotic to seawater (Cheng and others, 2006).

In some species, these mechanisms are not sufficient 

to completely prevent the entry of all ice crystals into the 

body. When this occurs at least some fish sequester these 

crystals in the spleen. Evans and others (2011) studied the 

process by which ice crystals are lodged in the spleen. Using 

nanoparticles as proxies for ice crystals, they found evidence 

that (1) AFP-coated ice crystals were removed from the 

circulatory system and stored by spleen phagocytes and that 

(2) these phagocytes were able to recognize the difference 

between AFP molecules alone (which they ignored) and 

those molecules attached to ice crystals (a process seen in 

crystallizable [Fc] receptor-mediated phagocytosis induced by 

immunoglobin-bound foreign particles [Swanson and Hoppe, 

2004]). Evans and others (2011) note that there remain several 

areas of uncertainty. First, it is unclear how high levels of 

AFP (all of which are first discharged into the intestinal tract) 

are maintained in the blood and lymph systems. Second, the 

fate of the stored splenic ice crystals is unknown. If fish find 

themselves in waters above freezing temperatures, as happens 

to some Antarctic species (Hunt and others, 2003), these 

crystals could be removed through passive melting.

Seasonal changes in AFPs in Arctic fishes are poorly 

understood and vary among species. For instance, although 

high AFP levels in Arctic Sculpin from the Canadian Arctic 

do not vary appreciably over the course of a year ([Fletcher 

and others, 1982], a pattern was observed in numerous 

Antarctic fishes [DeVries and Lin, 1977]). Arctic Cod 

taken off Svalbard, Norway (Nahrgang and others, 2010) 

and Shorthorn Sculpin (location unknown; DeVries, 1980) 

exhibited reduced AFP activity during summer and increased 

activity during winter. Generally, seasonal changes in AFP 

production seem to be widespread in fishes exposed to 

freezing winter temperatures and warmer summer waters 

(for example, Winter Flounder [Fletcher, 1981]; Atlantic Cod 

[Fletcher and others, 1987]; Ocean Pout [Fletcher and others, 

1985b]; Eelpout [Lycodes spp.; Sorenson and Ramlov, 2001]; 

and Atlantic Wolffish and Spotted Wolfish [Anarhichas minor; 

Desjardins and others, 2006]), although there is considerable 

interspecific variation in the timing of this cycle (Fletcher and 

others, 2001). Given that Arctic waters, particularly relatively 

nearshore, exhibit considerable summer warming, it might be 

expected that many species will exhibit seasonal changes in 

AFP production. However, the degree of seasonal variation of 

AFP produced by skin cells is much less than that produced by 

liver cells (Low and others, 2002).

Two environmental parameters, photoperiod and water 

temperature (sometimes working together), help control 

annual cycles in AFP production, although the importance 

of each parameter may vary with species. Photoperiod 

(mediated through the hypothalamus), for instance, is likely 

a major controlling factor in Winter Flounder, although 

water temperatures must be less than 8 °C for a buildup of 

AFP mRNA in the liver and subsequent production of AFP 

(Fourney and others, 1984; Davies and others; 1988, Fletcher 

and others, 1989). However, even in Winter Flounder, water 

temperature does influence the maintenance of AFPs in the 

body through controlling its removal from the blood stream 

and controlling the levels of AFP mRNA in the liver (Davies 

and others, 1988). On the other hand, Fletcher and others 

(1987) determined that changes in water temperature alone 

were primarily responsible for boosting AFP production in 

Atlantic Cod, because manipulating day length did not affect 

antifreeze production, whereas lowering water temperature to 

0 °C did. Finally, based on laboratory studies, it is likely that 

both seasonal changes in water temperature and photoperiod 

play a role in the timing of increases and decreases in AFP 

production in Atlantic and Spotted Wolffishes (Desjardins 

and others, 2006; Duncker and others, 2006). The timing 

of AFP production seems to be at least partially genetically 

fixed (Fletcher and others, 1985a), although these previously 

mentioned environmental cues, working through the central 

nervous system and the pituitary gland (particularly through 

the release of growth hormone), act to “fine tune” the precise 

timing of the onset and decrease of AFP production (Davies 

and others, 1988; Fletcher and others, 2001). 
628 Alaska Arctic Marine Fish Ecology Catalog

Within a species, concentrations of AFP may vary 

considerably, even among individuals living close together. 

As an example, juvenile Atlantic Wolffish hatched together 

from eggs collected in a single bay in Newfoundland, varied 

in their blood plasma AFP concentrations between a high of 

18.1 mg/ mL and a low of 8.5 mg/mL (Desjardins and others, 

2007) and similar between-individual variability was observed 

in sympatric adults of that species (Dejardins and others, 

2006). In these two wolffish studies, juveniles and adults 

harbored similar concentrations of AFPs.

Conspecifics taken from geographically separated areas 

commonly produce different amounts of AFP. For instance, 

Ocean Pout living off Newfoundland contained higher levels 

of AFP than did their conspecifics living off warmer-water 

New Brunswick, differences linked to the New Brunswick fish 

having only one-quarter the number of AFP genes than did 

those from Newfoundland (Hew and others, 1988). Similarly, 

in a comparison of plasma AFP from Ocean Pout from 

Denmark and Newfoundland, fish from both areas reportedly 

had high levels during the winter. However, the warmer-water 

Danish fish lost most of their plasma AFP during the summer, 

whereas levels in fish from Newfoundland remained relatively 

high (Fletcher and others, 1985b; Sorenson and Ramlov, 

2001). Similar variability occurs in, among other species, 

Atlantic Cod and Winter Flounder (Fletcher and others, 1985a; 

Goddard and others, 1999).

There are differences in AFP production within closely 

related species. Desjardins and others (2006) compared AFP 

production in the Atlantic Wolffish and Spotted Wolffish. They 

determined that although both species produced AFPs only 

the Atlantic Wolffish produced sufficient AFP to protect an 

individual down to the freezing point of seawater. Likely of 

significance is that, although both species encounter subzero 

water temperatures, the more shallow-dwelling Atlantic 

Wolffish has more of a chance of encountering ice and 

freezing conditions.

Even at temperatures greater than freezing, AFPs may 

protect cells (including perhaps fish cells) against damage. 

As an example, cells require the ability to expel calcium 

ions through active transport (thus balancing the passive ion 

transport that is continually occurring). It is thought that low 

temperatures may damage a cell’s ability to actively transport 

these ions out of the cell. Studies by Negulescu and others 

(1992) determined that AFPs “inhibited passive Ca entry 

across the cell membranes” without inhibiting such normal 

cell functions as active transport. Antifreeze proteins may also 

interact with various constituents of cell membranes to inhibit 

cell leakage and death (Rubinsky and others, 1990; Wu and 

Fletcher, 2000; Wu and others, 2001).

Lastly, the mechanism(s) of freezing protection in fish 

eggs are poorly understood. Davenport (1992) reported 

that the eggs of Capelin can survive as much as 6 hours at 

temperatures as low as -5.2 °C. He noted that the chorion of 

these eggs had a “sticky, thick extra outer layer, not found 

in other species, which appears to have the dual effect of 

slowing ice crystal penetration” thus implying that there is a 

morphological component to fish egg protection. However, 

Goetz and others (2005) determined evidence of what they 

believed to be an antifreeze protein in the eggs of Atlantic 

Cod and significant amounts of AFPs exist (and significant 

resistance to ice propagation exists) in the newly spawned 

eggs of Antarctic notothenioids (for example, Ploughfish 

(Gymnodraco acuticeps, Cziko and others, 2006).

Cellular and Metabolic Adaptations

Evidence for one long-proposed adaptation, called 

Metabolic Cold Adaptation (MCA), has been effectively 

undercut by a relatively recent series of studies. MCA holds 

that fishes adapted to cold waters appear to have evolved 

a higher metabolic rate “when corrected for differences 

in measurement temperature” that is 2–4 times that of 

temperate species (Steffensen and others, 1994, p. 49). The 

hypothesis was based on fish respiration experiments by 

Ege and Krogh (1914) and Krogh (1916) and later followed 

by those of Scholander and others (1953) and Wohlschlag 

(1960). However, studies on a number of Arctic and Subarctic 

species (for example, Arctic Cod, Arctic Staghorn Sculpin 

(Gymnocanthus tricuspis), Spatulate Sculpin (Icelus spatula), 

Shorthorn Sculpin, Fish Doctor (Gymnelus viridis), Greenland 

Cod [now known to be Pacific Cod]) strongly imply that MCA 

is almost certainly an artifact caused by insufficient laboratory 

acclimation time for fishes in respiration experiments 

(Holeton, 1974; Mikhail and Welch, 1989; Steffensen and 

others, 1994; Hop and Graham, 1995; Steffensen, 2002; 

Karamushko and others, 2004).

Much of the experimental research on the cold-adapted 

performance of fishes has focused on the selective forces 

on energy production, particularly on how these forces have 

influenced mitochondria. Cold temperatures affect the aerobic 

capacity of mitochondria; the respiration rates of isolated 

mitochondria generally decrease with decreasing temperature 

(Johnston and others, 1998). In response, and to maintain 

sufficient aerobic activity, cold-water species harbor higher 

volumes and densities of mitochondrial clusters than do more 

temperate taxa; these may reach 50 percent of the muscle fiber 

volume in the Antarctic Icefish ([Champsocephalus esox] 

see Johnston and others, 1998; Pörtner and others, 2005). 

However, several factors limit the amount of mitochondria 

produced by Arctic fishes. First, there is a maximum limit 

to mitochondrial densities beyond which there is not 

space for myofibrils, thus muscle function suffers, and the 

greater volumes of mitochondria in cold-water fishes only 

partially compensate for the reduced oxidative capacity the 

mitochondria afforded at low temperatures (Johnston and 

others, 1998). Second, an increase in mitochondrial density 

causes an increase in oxygen demand even under resting 

conditions and, as noted by Pörtner and others (2001, p. 1,992) 

this particularly becomes “detrimental during warming 

when [oxygen demand] can no longer be covered by oxygen 

uptake through ventilation and circulation.” As an example, 
Chapter 4 629

an increase in ambient water temperatures of only 2.3 °C 

results in large increases in oxygen consumption in Arctic 

Cod (Hop and Graham, 1995). Additionally, a comparison 

of the mitochondrial enzyme capabilities (citrate synthase, 

cytochrome c oxidase, and the electron transport system) of 

Arctic and boreal populations of Atlantic Cod demonstrate that 

there also have been cold adaptations in these systems (Lannig 

and others, 2003).

With greater mitochondrial densities in Arctic fishes has 

come a shift from carbohydrate to lipid (fatty acid) catabolism 

by these mitochondria. Although the rationale for this shift 

and the process by which it occurs remains somewhat obscure, 

one possible reason is that fatty acids are transported across 

cell membranes composed of fatty acids at a lower energy 

cost than those with high levels of carbohydrates (Pörtner, 

2002). As a consequence, lipid accumulations in Arctic fish 

tend to be higher than in many temperate taxa. Additionally, 

the mitochondrial membranes of Arctic fishes tend to have 

elevated levels of polyunsaturated fatty acid phospholipids, 

probably because this type of fat supports efficient energy 

release rates (proton leakage) through the inner mitochondrial 

membrane (Brand and others, 1992; Pörtner and others, 

2005). This increase in polyunsaturates, combined with the 

aerobic generation of high-energy phosphates, increases 

the opportunity for damage by free radicals and, likely in 

response, Antarctic fishes have high levels of the antioxidants 

vitamin E and C (Ansaldo and others, 2000; Gieseg and 

others, 2000; Davison, 2005).

In cold-water fishes, mitochondrial proliferation and 

associated adaptations come with physiological tradeoffs, as 

energy demands for these adaptations can be relatively high 

and would be shifted away from growth and reproduction 

(Lannig and others, 2003). Pörtner and others (2001) gave an 

indication of what these tradeoffs might entail. Their work 

was conducted on Atlantic Cod and Ocean Pout at a number 

of sites ranging from the European Arctic through the more 

temperate Baltic and North seas. They determined that in both 

species growth rates and fecundity declined with increasing 

latitude (that is, decreasing ambient water temperatures). 

Selection for greater metabolic efficiency at fluctuating 

lower temperatures also may have led to intraspecific 

polymorphisms in the hemoglobin molecule. Functional 

differences in polymorphic hemoglobin molecules are 

known from Arctic Cod (Boreogadus saida) and Ice Cod

(Arctogadus glacialis) (Verde and others, 2006) as well as 

in Spotted Wolffish, Ocean Pout, and in several Antarctic 

taxa (Hjorth, 1974; Verde and others, 2002). As an example, 

some hemoglobin molecules of both Boreogadus saida and

Arctogadus glacialis show a low oxygen affinity and a low 

Root effect, probably linked to the high concentrations of 

oxygen in cold marine waters (Verde and others, 2006). 

Polymorphic hemoglobins, labeled HbI(I) and HbI(2), have 

been most intensively studied in the Atlantic Cod. First 

described by Frydenberg and others (1965) and Sick (1965), 

the frequency of the two alleles shows a north-south cline 

along the Norwegian and eastern North American coasts. 

In both instances, a greater proportion of fishes living in 

cold more northerly, waters carry the HBI(2) alleles. Since 

that time, a range of studies has demonstrated that the 

two polymorphisms function most efficiently at different 

temperatures (Karpov and Novikov, 1981; Pörtner and others, 

2001; Brix and others, 2004). For instance, HbI(2) is better 

fitted to cold temperatures through its ability to transport more 

oxygen to the tissues by superior oxygen binding capabilities 

at low temperatures. This, in turn, seems to be responsible 

for differences in a number of life history parameters. For 

instance, Atlantic Cod living off northern Norway and 

carrying the homozygotic HBI(2) allele exhibit faster growth, 

earlier maturation, and earlier spawning season than do fish 

carrying the homozygotic HBI(1) allele (Mork and others, 

1983). Laboratory studies have also shown that Atlantic Cod 

held at low temperatures grew faster when carrying the HBI(2) 

gene (Imsland and others, 2004). Additionally, fish behavior 

also may be influenced by hemoglobin type as juvenile cod 

that carry the HbI(2) form seem to preferentially inhabit 

significantly lower temperatures than do fish carrying the 

HbI(1) variant (Petersen and Steffensen, 2003). Although few 

studies have examined the handful of cartilaginous species 

that inhabit polar waters, the hemoglobin of the Arctic Skate 

(Amblyraja hyperborea) and the Antarctic Eaton’s Skate 

(Bathyraja eatonii) seem to be (1) similar to one another and 

(2) functionally different from those of temperate cartilaginous 

species (Verde and others, 2005).

Hibernation

Small to moderate levels of reduced activity, feeding, and 

growth in winter are widespread among fishes, largely because 

metabolism is directly coupled to environmental temperature. 

However, true hibernation among fishes has been perhaps 

best documented in the Antarctic species Black Rockcod 

([Notothenia coriiceps]; Johnston and Battram, 1993; Coggan, 

1996; Campbell and others, 2008). These studies determined 

that in natural habitats during winter (or in testing situations 

that simulated winter conditions) Black Rockcod had 

(1) greatly reduced feeding rates (even when additional food 

was available) and increased mobilization of lipid reserves, 

(2) a negative growth rate (the loss of body mass), (3) a 

58 percent suppression in total metabolic rate and 29 percent 

in standard metabolic rate, and (4) a 20-fold reduction in 

activity and a 6-fold reduction in home range. Thus, despite 

living in a thermally stable environment where food was 

readily available throughout the year, N. coriiceps exhibits 

a number of the characteristics of terrestrial hibernators. It 

is likely that the trigger for hibernation in this fish species is 

decreasing light levels rather than decreasing temperatures. As 

Campbell and others (2008, p. 7) note: “The winter dormancy 

we have documented...is distinct from the facultative 

dormancy observed in temperate fish species by the levels, and 

duration of the reduced physiological state...”
630 Alaska Arctic Marine Fish Ecology Catalog

Hibernation in fishes is likely a continuum (Campbell 

and others, 2008) ranging from small amounts of metabolic 

depression in some species to the extreme examples of 

physiological and behavioral adaptations noted in Black 

Rockcod. What evidence for hibernation is there in Arctic 

fishes? To date, no studies have directly addressed this issue 

in fishes living in the Chukchi and Beaufort Seas. However, 

results from other research imply that hibernation, or at least 

dormancy, is a likely strategy for many Arctic species. For 

example, in the Subarctic waters of Newfoundland Cunner 

(Tautogolabrus adspersus) display numerous attributes 

suggestive of dormancy (Green and Farwell, 1971; Green 

1974; Valerio and others, 1990). At temperatures less 

than 5 °C, Cunner become extremely torpid and shelter 

in the rocks and boulders of shallow, nearshore waters (at 

water temperatures as low as -1 °C). Here they remain, 

without feeding, throughout winter and spring, until water 

temperatures increase in May or June. Additionally, Cunner 

produces a thick mucus coat that likely hinders contact 

between ice crystals and skin and that skin produces 

antifreeze. Unlike Antarctic fishes, however, these survival 

mechanisms are imperfect, as extremely stormy winter 

conditions can drive ice crystals with sufficient force against 

sheltering fishes that they freeze to death. In the cold waters 

of laboratory experiments, a North Atlantic species, Winter 

Flounder, burrow into soft bottom, perhaps to avoid ice 

crystals (Fletcher, 1975).

Hibernation has not been directly investigated in Arctic 

fishes. However, the winter behavior of young Arctic Cod 

in Franklin Bay, southeastern Beaufort Sea off Canada, led 

Benoit and others (2010) to speculate that these fish were 

hibernating. During this hibernation period, these fish form 

extremely dense midwater schools and make only occasional 

vertical excursions, likely for feeding. Noting the slow 

digestion rates in this species (about 17 days), Benoit and 

others (2010) speculated that fish remained for the most part in 

a torpid state, leaving that state only when sufficiently hungry. 

Amphidromous species, such as Arctic Cisco (Coregonus 

autumnalis) and Least Cisco (C. sardinella), overwintering in 

the Sagavanirktok River and Colville River deltas in Alaska 

may become torpid during winter, as feeding ceases or is much 

reduced (Schmidt and others, 1989). However, during winter 

these fishes will migrate upstream away from the more saline 

conditions that occur as winter progresses, so hibernation is 

not complete. On the other hand, the marine Fourhorn Sculpin 

(Myoxocephalus quadricornis), a dominant benthic species 

in the nearshore of the Beaufort and Chukchi Seas, shows no 

sign of hibernation, as it actively feeds throughout the year 

(Schmidt and others, 1989). Fishes including Arctic Cisco and 

Least Cisco, Humpback Whitefish (C. pidschian) and Broad 

Whitefish (C. nasus), Arctic Cod, Saffron Cod (Eleginus 

gracilis), Fourhorn Sculpin (M. quadricornis), and Arctic 

Smelt (Osmerus dentex) are all captured in winter fisheries, 

implying enough individual activity for them to become 

entangled in gillnets or to strike hooks (Craig, 1989b).

Other Functional Adaptations

As water temperatures decrease, blood viscosity 

increases in Antarctic fishes. For instance, blood viscosity 

in the Emerald Rockcod (Trematomus bernacchii) at 0 °C is 

about 40 percent higher than at 10 °C, thus increasing cardiac 

workload (Axelsson, 2005). To compensate for this added 

viscosity, a major adaptation in the circulatory system of 

Antarctic fishes in extremely cold water has been a reduction 

in haematocrit (packed red blood cell volume) and mean cell 

hemoglobin concentration (Egginton, 1996; Davison and 

others, 1997). Interestingly, there is no evidence for increased 

flexibility of red blood cells, although this would also reduce 

the effect of increased viscosity (Lecklin and others, 1995). 

Additionally, the gross anatomy of Antarctic fish hearts 

appears to be similar to those of more temperate-water taxa, 

although the diameter of Antarctic fish heart myocytes (muscle 

cells) are slightly larger, due to an increased number of 

mitochondria (Axelsson, 2005). There is no evidence for cold 

adaptation of vascular control as measured by branchial artery 

contractility in polar compared with temperate fishes (Hill and 

Egginton, 2010).

Most research addressing possible adaptations to the 

nervous system of Arctic fishes also has studied Antarctic 

species. These studies demonstrate that polar fishes have 

evolved a number of adaptations to this cold environment. 

Among these, alterations in the cell membrane to maintain 

fluidity is one of the most important and is accomplished 

through an increase in unsaturated fatty acids, thus lowering 

viscosity that would normally attend ambient lower 

temperatures (Morris and Schneider, 1969; Macdonald, 1981). 

Work on Shorthorn Sculpin from the northern Bering Sea 

demonstrates that at low temperatures this trend of increasing 

membrane fluidity declines or ceases (Cossins and Prosser, 

1978). Conductivity within the axoplasm of nerve cells also 

has adapted to lower temperatures through an increase in 

ion concentrations within and around these cells (Dobbs and 

DeVries, 1975; MacDonald, 1981). The brain gangliosides 

(molecules in the cell plasma membrane that influence 

membrane fluidity and are involved in cell signal transduction) 

of Antarctic fishes seem to be cold adapted through an increase 

in their concentrations and polarity (Becker and others, 1995). 

However, this variety of adaptations leads to only a partial 

compensation for low temperatures. Overall, the nervous 

systems of Antarctic fishes transmit impulses at a slower rate 

than in fishes living in warmer environments (Macdonald, 

1981; Pockett and Macdonald, 1986), a phenomena also found 

in the peripheral nerves of Arctic species such as Arctic Cod, 

Snakeblenny (Lumpenus lampretaeformis), and an unidentified 

eelpout (Lycodes sp.) (Moran and Melani, 2001). 

Similar to the nervous system, the musculature of 

cold‐water fishes has seen a series of adaptations to maximize 

function under Arctic conditions. These adaptations include 

(1) changes in myosin structure (Johnson and Johnston, 1991) 

and the abilities of cross bridges to generate force (Johnston, 
Chapter 4 631

1985; Johnston and Altringham, 1985), (2) increases in rates 

of Ca+ accumulations in sarcoplasmic reticulum (McArdle and 

Johnston, 1980), (3) and changes in the activation enthalpies 

of the sarcoplasmic reticulum ATPase (McArdle and Johnston, 

1980). All these adaptations have helped bolster the force 

generation, muscle contraction speed, and twitch duration 

of cold muscles beyond what might be expected. Within the 

Antarctic notothenioid fishes, there has also been a marked 

trend toward large white and red muscle fibers (Davison, 

2005). However, this may represent an ancestral trait within 

the group, rather than an adaptation to cold water. As with the 

nervous system, these compensations (including the changes 

in aerobic capacity due to increases in mitochondrial densities 

are relatively modest and muscle function of fishes living 

in Arctic conditions is significantly impaired compared to 

those of fishes living in warmer waters (Montgomery and 

Macdonald, 1984; Johnson and Johnston, 1991). For instance, 

power output for the Antarctic Icefish at -1 °C is about 

60 percent that of the Pacific Blue Marlin (Makaira nigricans) 

at 20 °C (Johnston and Altringham, 1985). Together, this 

translates to generally lower maximum swimming speeds and 

low maximum tail-beat frequencies in polar fishes (Johnson 

and Johnston, 1991) and to relatively poor burst swimming 

performance in at least some of these species (Archer and 

Johnston, 1989).

Despite a range of compensatory adaptations, coldwater dwelling fishes remain incapable of maintaining the 

metabolic rates and active life styles of most temperate species 

(Macdonald and Montgomery, 1982; Johnson and Johnston, 

1991; Lannig and others, 2003). This is true both for the 

Antarctic (Steffensen, 2005) and for northern Arctic and 

subarctic fishes, which have the additional burden of coping 

with significant annual changes in temperature (Lannig and 

others, 2003). Inevitably, many Arctic species have a relatively 

sluggish (that is, lowered mobility) lifestyle and shift a 

significant amount of their energy budgets away from growth 

and reproduction (summarized in Steffensen, 2005). 

Life Strategy Adaptations

Arctic fishes have evolved numerous life strategies for 

coping with low temperatures, long periods of diminished 

light conditions, and brief periods of marine production 

that generally favor benthic-dwelling species (for example, 

Mecklenburg and Steinke, 2015). These strategies relate 

to a fish’s environmental requirements for food, growth, 

and reproduction. These requirements involve multiple life 

history stages, habitats, and behaviors that optimize life 

processes and opportunities for successful reproduction. 

Diadromous (migratory) patterns that involve amphidromy 

or anadromy are common life strategies that link the fish to 

stable conditions in freshwater and marine environments 

and, at the species level, are characterized by species that 

vary widely in their tolerance to salinity, exhibit variable 

freshwater residencies in immature fish, older ages at first 

reproduction, and widespread differences in their fidelity, 

or suspected fidelity, to known spawning and overwintering 

areas (for example, DeGraaf and Machniak,1977; and Craig, 

1989a). The amphidromous species (for example, charr and 

whitefishes) are iteroparous and generally long-lived, typically 

living 20 years or more. They tend to mature at 7–8 years and, 

after their first spawning event, tend to spawn every other year 

thereafter. Anadromous species (for example, Pacific salmon 

and lampreys) are semelparous, spending most of their lives at 

sea. The anadromous fishes tend to live between 2 and 7 years 

and die shortly after spawning. These species tend to be much 

larger and more fecund than the amphidromous fishes. Both 

are conspicuous members of the Chukchi and Beaufort Sea 

nearshore fish assemblages during summer months. Existing 

information for many marine fish species suggests shorter life 

cycles, earlier maturation, and greater relative investments of 

energy in the production of young (for example, large numbers 

of eggs). There are exceptions such as sharks, rays, and 

Pacific Halibut. The location of important seasonal habitats, 

reproductive ecologies, and life histories of most species from 

Arctic Alaska remain largely undescribed. It is known that 

many populations, or segments of populations have important 

life history linkages to coastal waters for food, reproduction, 

and migration between seasonal habitats. Examples include 

all of the amphidromous and anadromous species as well as 

other coastal dominants such as Arctic Cod, Arctic Flounder, 

Capelin, Arctic Smelt, and several sculpin species such as the 

Fourhorn Sculpin. Generally, onshore-offshore movements 

such as those related to ontological development of species 

like Pacific Herring and Arctic Smelt, or feeding or spawning 

of Arctic Cod remain to be described because winter surveys 

are lacking.

Reproductive Ecology

Although patterns of reproductive ecology (for example, 

timing of reproduction) vary considerably among species, 

the spawning seasons and incubation periods of some Arctic 

fishes, particularly anadromous or amphidromous species, 

are timed to take advantage of the late spring-early summer 

breakup of ice and the concomitant increases in photoperiod 

and temperatures and in primary and secondary production 

of prey. This synchrony is a “hedge betting” strategy that has 

evolved to increase the probability for early recruitment. For 

instance, although many amphidromous species (for example, 

charr, ciscoes, and other whitefishes) spawn under ice in 

autumn (Craig, 1989a), the eggs remain in gravel beds for 

7–8 months and larvae emerge from April to July (McCart 

and others, 1972; Yoshihara, 1973; Craig and McCart, 1974; 

Griffiths and others, 1975; Burns, 1990). In the Beaufort and 

Chukchi Seas, both juvenile and adult Arctic Smelt overwinter 

under ice in brackish river deltas and coastal waters; many 

of the river mouths along the Beaufort and Chukchi Seas 

harbor overwintering populations (Bond, 1982, Schmidt 

and others, 1987; Bond and Erickson, 1989, Craig, 1989a; 
632 Alaska Arctic Marine Fish Ecology Catalog

Sekerak and others, 1992). As winter progresses, fish gather 

near spawning grounds (Haldorson and Craig, 1984) and 

spawning occurs in spring, just prior to ice break-up with most 

spawning occurring in the lowermost, but still fresh, parts of 

rivers (Belyanina, 1968; Burns, 1990). Compared to autumnspawning fishes, however, Arctic Smelt eggs hatch quickly, in 

10–30 days depending on water temperature (Belyanina, 1968; 

Morrow, 1980; Gritsenko and others, 1984; Burns 1990). 

Saffron Cod spawn under ice from December to at least May 

(Andriyashev, 1954; Stewart and others, 1993) and Arctic 

Flounder from March to June (Ratynski, 1983; Lawrence and 

others, 1984). However, the spawning seasons of other marine 

fishes appear to be less tightly linked to seasonal productivity 

and are perhaps tied to pelagic-benthic coupling of energy to 

benthic organisms that comprise the diets of the juveniles of 

some species. For instance, in the Chukchi and Beaufort Seas, 

Pacific Herring spawn from June to September (Ratynski, 

1983; Gillman and Kristofferson, 1984; Lawrence and others, 

1984; Stout and others, 2001), Capelin primarily in July and 

August (Kendel and others, 1975; Fechhelm and others, 1984; 

Stewart and others, 1993), and Polar Eelpout (Lycodes polaris) 

likely in autumn or early winter (Andriyashev, 1954; Frost 

and Lowry, 1983). In some instances, at least part of the adult 

population spawns during a season where few larvae survive. 

As an example, Fortier and others (2006) demonstrated that 

most of the Arctic Cod larvae produced during the relatively 

cold spring die soon after. However, early spawning persists 

in this species because the few larvae that survive are larger 

and better able to survive the following winter than the larvae 

produced later in the year.

Synchrony

The regional bathymetry of the Chukchi and Beaufort 

Seas indicates small-scale local variability in the physical 

location of the shelf break relative to average conditions across 

larger scales. In many areas, the gradient between 75- and 

200-m is relatively sharp, occurs over a short distance, and 

the corresponding surface area and volume is relatively small. 

The effects of small-scale physical and biological features 

resulting from shelf-slope interactions along a shallower, but 

sharper gradient, especially in the northeastern Chukchi and 

western Beaufort Seas, could be seasonally important to many 

species having evolved life history and reproductive ecologies 

linked to, or concentrated in these environments (Crawford 

and others, 2012). For marine fishes occupying shelf and slope 

habitats, this is important because the ecological processes 

operating over short times and small spatial scales, such 

as food or prey concentrations, may be especially critical 

to early life history success (for example, match-mismatch 

survival of larval fish predators). For instance, it is common 

that wind forcing at the shelf break of both seas frequently 

drives upwelling events that lift nutrient-rich water from 

the depths to the shelves (from Crawford and others [2012, 

p. 180], citing Weingartner [1997] and Carmack and Kulikov 

[1998]). We hypothesize that shelf-slope processes in areas 

of sharp gradients are important oceanographically to life 

history processes. However, from a practical view, the spatial 

extent of existing data relative to this narrow depth interval is 

coarse at best. For analytical purposes, this scale of resolution 

limits our predictive capability and, although we know from 

many studies (for example, Norcross and others, 2009) that 

temperature and depth are important determinants of fish 

distribution, only broad hypotheses are currently possible 

regarding the relative importance of habitats near shelf break 

and slope areas in the U.S. Arctic (for example, Crawford 

and others, 2012). At the population level (for example, 

reproduction, immigration, and emigration processes), the 

importance of shelf-slope processes are further suggested by 

newer biogeographic information presented herein.

Life History and Behavioral Adaptations

Unlike the small number of well-studied nototheniid 

species of Antarctica that live in a stable environmental 

regime, the behavioral adaptations of Chukchi and Beaufort 

Seas fishes to seasonally fluctuating temperature, salinity, 

turbidity, and variable food conditions are quite poorly 

understood. Much research, especially in the coastal waters 

of the Beaufort Sea indicates the existence of many adaptive 

responses to seasonally dynamic hydrographic and biological 

properties of nearshore ecosystems and coastal lagoons. 

Examples include fish migration (for example, bathymetric, 

between freshwater and marine), reduced activity and potential 

hibernation, and reproductive specialization (for example, 

Arctic Cod and sea ice). How reproduction and early life 

history in marine fishes (that is, timing, habitat locations, 

and spawning behaviors) have evolved with respect to being 

in synchrony with biological production events, sea ice, 

hydrological and meteorological conditions remains to be 

described. For most species, studies of genetic diversity and 

population structure are needed to understand the variability 

in patterns of age and growth that have been observed and 

genetic relationships in the regional fauna.

Migration is an evolved process that reflects the regular 

movement of fish populations between habitats important 

to the completion of their life cycle. Movement, although 

related, generally describes the immediate tactical responses 

of fish to environmental surroundings. During winter, ice 

covers the marine waters of the Chukchi and Beaufort Seas 

and in some coastal areas freezing occurs throughout the 

water column. In response, fishes move into unfrozen parts 

of rivers (that is, amphidromous and anadromous taxa), into 

river mouths (euryhaline species), into unfrozen, near-shore 

marine refuges, or perhaps into deeper, more-offshore marine 

waters. Arguably, seasonal-linked coastal migrations of many 

amphidromous species (whitefishes) are reasonably well 

understood. However, there have been few winter studies of 

Arctic nearshore or, particularly offshore, marine fish species. 

Thus, the behavior of the generally sluggish, benthic species 

that occupy these waters is little understood.
Chapter 4 633

With that caveat, examples of life strategy models of 

known or inferred seasonal movement patterns of some 

of the most conspicuous or ecologically and traditionally 

important species inhabiting the Chukchi and Beaufort seas 

are presented. This focus, illustrated with several prominent 

species, or groups of species, serves to model key features of 

the life history strategies and regional variations in migratory 

patterns in order to explain the range of behaviors and habitat 

dependencies that have evolved. The models described reflect 

adaptive strategies for life processes that occur in Arctic 

marine fishes across the continuum of freshwater, estuarine, 

and marine habitats:

Amphidromous/Anadromous Model.—This segment 

of the nearshore fish community is important in subsistence 

fisheries throughout Arctic Alaska. Prior to the 1990s, most 

technical reports and publications did not distinguish between 

the two life strategies and referred only to anadromous 

species. Craig (1989a) examined the evolution of anadromy 

in the Arctic as an adaptation related to differences in food 

quality and availability and energetic benefits conferred by 

coastal migrations. Most young fish reside in fresh water for 

1–3 years prior to their first coastal excursion, the exception 

being the Arctic Cisco in the Alaska Beaufort Sea. However, 

although the species undertakes a coastal migration during its 

first year of life, once re-entering freshwaters in and around 

the Colville River, they resume the variable residency strategy. 

Overwintering habitats in freshwaters may be limiting and, 

where studied, strong density dependence is evident in their 

population dynamics.

Amphidromy Example: Dolly Varden.—All Dolly 

Varden (Salvelinus malma) overwinter in fresh water habitats. 

Amphidromous Dolly Varden must return to rivers and 

streams during summer or early autumn and timing or return 

often varies with whether they will spawn that year (Craig and 

McCart, 1975; Griffiths and others, 1975). Except for the rare 

and small habitat areas kept free of ice by perennial springs 

(taliks), the upper parts of rivers draining into the Beaufort 

Sea and southeastern Chukchi Sea freeze solid during winter. 

Perennial springs are located in rivers and streams flowing 

from the eastern Brooks Range into the Beaufort Sea (Craig 

1989a; Wiswar, 1994). Both spawning and overwintering 

occurs in those tributaries where perennial springs prevent 

freezing throughout the water column (Craig and McCart, 

1974; Griffiths and others, 1988; Burns, 1990), whereas in 

rivers flowing to the Chukchi Sea, the lower parts of the 

waterways also may be partially ice-free (DeCicco,1996; 

DeCicco, 1997). It is likely that spawning does not occur in 

low-lying coastal plain rivers of the southwestern Beaufort Sea 

west of the Colville River because they tend to be connected 

to lakes and lack the perennial springs that prevent freezing 

(Burns, 1990; Gallaway and Fechhelm, 1997). Thus, in some 

waterways, a large segment of a population may be found in a 

small stretch of water (Burns, 1990).

In Beaufort Sea drainages, most Dolly Varden are thought 

to overwinter in their natal drainages. For instance, in the 

Sagavanirktok River, one study determined that an estimated 86 

percent of overwintering fish had originated in that river (Crane 

and others, 2005). In the rivers flowing into the southeastern 

Chukchi Sea, fish do not show overwintering fidelity to natal 

waterways and stocks from various rivers share ice-free zones 

(Armstrong and Morrow, 1980; DeCicco, 1997; Crane and 

others, 2005).

As they make their return migrations into fresh waters, 

the timing and behavior of Dolly Varden is quite complex and 

varies with location, reflective of variable water temperature 

regimes and states of maturity (DeCicco, 1997; Crane and 

others, 2005). Spawning occurs in autumn. During those 

years when a fish will spawn, some spawners will remain 

in freshwater; others will undertake an abbreviated coastal 

migration and return to their spawning habitats as early as 

late June (Armstrong and Morrow, 1980). The lack of, or 

short-term nature of, the coastal migration of an individual may 

be an adaptive strategy that allows these fish to occupy and 

defend premium spawning sites. Additionally, size and bright 

coloration helps them attract mates and, by remaining resident, 

or almost resident, allows them to spawn during most optimal 

times. However, although many Dolly Varden begin returning 

in July, in most instances most Dolly Varden are thought to 

return to freshwaters from mid-August and early-September. 

The environmental trigger for this migratory behavior is 

unknown but is thought to relate to changing light and 

hydrographic properties. A return migration may occur as late 

as October (after ocean freeze-up) before fish move into river 

systems (Yoshihara, 1973; Kendel and others, 1974; Bendock, 

1977; Craig and Haldorson, 1981; Griffiths and others, 1988; 

Burns, 1990; Thorsteinson and others, 1990). 

The migratory behaviors of fish in the southeastern 

Chukchi Sea drainages are more complex than in the Beaufort 

Sea drainages and regional differences in summer and autumn 

spawning populations may be related to differences in thermal 

regimes and summer durations of open water periods (DeCicco, 

1997). There is apparently a greater incidence of summer 

spawning in Chukchi Sea drainages. Summer spawning fish 

do not seem to migrate to sea during the spawning year. 

Rather, in June and early July, they ascend rivers, spawn, and 

then descend to lower river over-wintering grounds. Summer 

spawning fish that have overwintered in non-natal rivers move 

into the Chukchi Sea in June, migrate directly to the spawning 

grounds, and descend to lower-river overwintering areas. In 

both instances, fish do not feed in the sea during the spawning 

year. Fish that spawn in autumn behave more like their Beaufort 

Sea congeners, migrating to sea in summer, feeding, and then 

returning to fresh waters in autumn. Recent satellite telemetry 

studies of Dolly Varden from the Wulik River reconfirm the 

extensive ocean migrations by adult fish (Courtney, 2014), 

whereas previous, limited tagging indicated transboundary 

movements between Alaska and Russia (DeCicco, 1992).
634 Alaska Arctic Marine Fish Ecology Catalog

Amphidromy Example: Whitefishes.—The seasonal 

movements of all of the whitefishes of the U.S. Arctic (Arctic 

Cisco, Bering Cisco, and Least Cisco, Broad and Humpback 

Whitefish) have much in common. The degree of tolerance for 

salinity varies by species and affects the extent of intrusions 

into marine waters. The Least Cisco life cycle model is 

broadly representative of most whitefishes and generally 

illustrates the migratory pattern of this group.

Juvenile Least Cisco may remain in rivers for several 

years before entering the sea (variable residence). During 

feeding migrations in coastal waters, Least Cisco schools 

can maintain this integrity for months (Craig and Haldorson, 

1981). Juvenile fishes do not migrate as far away from natal 

origins as larger adults and tend to remain in brackish habitats 

(5–10 °C, 10–25 practical salinity units [psu]). Older fishes 

are more tolerant of both colder temperatures and increased 

salinity (27–32 psu), and disperse farther along the coast. Most 

Least Cisco stay within the brackish water lens, although they 

have been found in river plumes as much as 20–30 km (12–19 

mi) off shore (Kendel and others, 1975; Griffiths and others, 

1988; Burns 1990, Schmidt and others, 1991; Philo and others, 

1993a, 1993b). Most Least Cisco found along Alaska’s south 

central Beaufort Sea are thought to originate from spawning 

and over wintering grounds in the Colville River (Griffiths 

and others, 1988). Generally, the extent of coastal migrations 

is affected by prevailing summer winds (for example, wind 

direction and speed). Least Cisco from the Colville River 

migrate farther east in years of persistent westerly winds. 

Similarly, fish from the Mackenzie River benefit from 

wind-aid migrations and move farther to the west in years 

of prevailing easterly winds (Griffiths and others, 1988; 

Fechhelm and others, 1996; Gallaway and Fechhelm, 1997). 

Along Alaska’s North Slope, larger fish can swim faster 

and reach distant feeding grounds before smaller individuals 

(Fechhelm and others, 1996). Given the short summer season, 

Least Cisco can travel substantial distances when conditions 

are favorable. For instance, a fish tagged in Simpson Lagoon 

was recaptured off Barrow, 300 km to the west (Craig and 

Haldorson 1981). However, fish leaving the Colville River 

generally migrate eastward along the coast of the Alaska 

National Wildlife Refuge because marine waters bathing the 

nearshore between Smith Bay and Cape Halkett appear to 

retard westward movement (Gallaway and Fechhelm, 1997; 

L. Moulton, MJM Research, oral commun., 2012). Most of 

the fish emigrating from the Mackenzie River travel westward 

perhaps as far as Phillips Bay (Kendel and others, 1975) and 

eastwards to the Tuktoyaktuk Peninsula (Percy, 1975).

Between July and September, Least Cisco begin to 

return to freshwater spawning and overwintering grounds 

(Jones and Den Beste, 1977; Craig and Haldorson, 1981; 

Griffiths and others, 1988; Fechhelm and others, 1996). At 

least in some areas (for example, Prudhoe Bay), the initiation 

of this migration may be linked to the onset and intensity 

of the coastal marine water intrusion or to food availability 

(Gallaway and Fechhelm, 1997). Several researchers have 

noted that the fish that will spawn in a given year are the first 

to leave coastal waters, followed by non-spawning adults and 

juveniles (Mann, 1974; Kendel and others, 1975; Jones and 

Den Beste, 1977). Both juveniles and adults overwinter in the 

brackish waters of river deltas (the Colville and Mackenzie 

Rivers are the largest over wintering areas along the Beaufort 

Sea), open coast (for example, Tuktoyaktuk Harbor), and the 

freshwater lakes of the Tuktoyaktuk Peninsula, Canada (Percy, 

1975; Bond, 1982; Fechhelm and others, 1996). Except for 

the Sagavanirktok River (where some overwintering occurs 

in delta waters), Least Cisco do not seem to either spawn 

or overwinter in any waterway between the Colville and 

Mackenzie Rivers (Fechhelm and others, 1996). Although 

water temperature and food availability likely trigger most 

of the movements in this species, Schmidt and others 

(1989) observed that during their under-ice sojourn in the 

Sagavanirktok River Delta these fish also moved upstream 

when saline waters increased over the course of the season.

Anadromy Example: Pacific Salmon.—Although all 

five species of Pacific salmon spawn in rivers entering the 

Chukchi Sea (Craig and Haldorson, 1986; Craig, 1989b; 

Healey, 1991, Mecklenburg and others 2002; Stephenson, 

2006), currently only Pink Salmon and Chum Salmon are 

likely to spawn in significant numbers in the rivers of the 

Beaufort Sea (Stephenson, 2006; Irvine and others, 2009). All 

of the salmon species spend at least one winter at sea. Given 

the harsh conditions in the marine nearshore, the location 

of where Beaufort Sea salmon overwinter while at sea is 

of increasing interest. Irvine and others (2009) posit three 

options: (1) Bering Sea Refuge—during their marine sojourn, 

young salmon migrate westward and live in the Bering Sea 

and perhaps Gulf of Alaska; (2) Atlantic Layer Beaufort 

Refuge—salmon remain in the Beaufort Sea, but winter deep 

under the pack ice; (3) Freshwater Beaufort Refuge—salmon 

remain in the Beaufort Sea region, overwintering in river 

plumes. Irvine and others (2009) examined the strontium 

(Sr):calcium (Ca) microchemistry signatures of otoliths of 

Chum Salmon from the Colville River (emptying into the 

Beaufort Sea) and the Tanana River, a tributary of the Yukon 

River, which drains into the Bering Sea. Although the results 

were somewhat ambiguous, Irvine and others (2009) reported 

that “the most parsimonious explanation is that Arctic chum 

salmon live in the North Pacific for most of their marine 

life...[but] may spend their first winter deep within the 

Beaufort Sea.”

Nearshore—Marine Model

The movements of the vast majority of species 

inhabiting the waters of the U.S. Chukchi and Beaufort 

Seas are not known. However, seasonal data are available 

for a small number of species, especially those occurring 

in coastal waters during summer months. The brackish 

water conditions, especially along the Alaska Beaufort Sea, 

provide important nursery habitats for Arctic Cod, Capelin, 
Chapter 4 635

Pacific Herring, snailfish, and probably many other marine 

fishes that are not adequately represented in the sampling 

of existing research (Thorsteinson and others, 1991). The 

(1) roles of deltas, lagoons, and other nearshore habitats for 

reproduction, incubation, and rearing of early life stages; (2) 

relative importance of such coastal areas with respect to fish 

production; and (3) evolutionary significance of these sites 

with respect to freshwater inflows, substrate, or wind- and 

current-aided transport or retention of early life stages are 

expected to be the subject of future studies.

Linked Estuarine—Marine Example: Forage 

Species.—The spawning, overwintering, and migration 

patterns of Pacific Herring are highly variable. For example, 

Bond (1982) reported that many Pacific Herring remained 

in Tuktoyaktuk Harbor (Canadian Beaufort Sea) for most 

of the year, leaving the harbor only for a few months during 

the summer to feed. Of the 10 known wintering sites in the 

Tuktoyaktuk Peninsula region, 8 were in estuarine coastal 

habitats, 1 was in the lower Mackenzie River, and 1 in the 

saline waters of Tuktoyaktuk Harbor (Sekerak and others, 

1992). At the other extreme, many herring in the eastern 

Bering Sea overwinter hundreds of kilometers offshore (at 

depths of 110–130 m [361–426 ft]) and move into nearshore 

waters in spring preparatory to summer spawning (Dudnik 

and Usol’tsev, 1964). It is not known if Pacific Herring use 

offshore waters in the U.S. Chukchi and Beaufort Seas. 

Capelin and Arctic Sand Lance (Ammodytes hexapterus) 

are seasonal dominants of the Chukchi Sea nearshore and are 

important in the diets of some fish, seabirds, and mammals. 

Like the Pacific Herring, reproduction, especially in Capelin 

is linked to inshore and coastal waters. Their distribution 

over shelf waters and seasonal use of shelf habitats, including 

onshore-offshore linkages are not well understood. Arctic 

Sand Lance burrow into sandy substrates of the shelf and 

are known to spawn coastally. In the Beaufort Sea, Capelin 

reproduction occurs in the delta habitats of the larger rivers 

such as the Colville and Sagavanirktok Rivers near Prudhoe 

Bay. Thorsteinson and others (1990) hypothesized that the 

successful reproduction of Capelin in these areas was spatially 

and temporally linked to the first marine intrusion of the 

year. That is, the success of Capelin spawning, incubation, 

and offshore transport and recruitment of young fish to other 

nearshore habitats is related to strong westerly winds and 

oceanographic conditions that push inshore waters carrying 

larvae and early-age juveniles offshore along the central 

coast. The hypothesized wind-driven transport in relatively 

fresh surficial waters was further suggested to span a critical 

period for capelin growth and physiological development 

in which juvenile transformation from living in brackish to 

marine conditions occurs (Thorsteinson and others, 1990). 

These authors noted a strong association between young 

Capelin and Arctic Cod, a predator, in surface catches 

taken off the Sagavanirktok River, which suggests another 

adaptive mechanism linking, in this case, brackish and 

marine environments.

Nearshore—Marine Example: Demersal Species 

and Other Coastal Dominants.—If the water column does 

not completely freeze, some nearshore species remain in 

shallow waters throughout the year. An example of such 

residency is the Fourhorn Sculpin. In the Beaufort Sea, these 

fish overwinter along shallow-bottom coastlines (Kendel 

and others, 1975; Percy, 1975; Jones and Den Beste, 1977; 

Craig and Haldorson, 1981) and in river deltas (for example, 

Colville and Sagavanirktok River deltas, see Craig [1989a] 

and Schmidt and others [1987]). A few winter surveys have 

determined that at least some individuals of the following 

species also overwinter more-or-less where they reside during 

warmer months: Arctic Smelt, Saffron Cod, and unidentified 

snailfish (Craig and Haldorson 1981, Craig 1989b). The Arctic 

Flounder, another shallow-water species, however, is known to 

undertake a bathymetric migration and move slightly offshore 

in autumn. In the Beaufort and Chukchi Seas, the extent of 

this winter migration is not known and may not be extensive 

(Andriyashev, 1954, Craig and Haldorson, 1981). For 

instance, in Tuktoyaktuk Harbor, fish overwinter in the deep 

parts of the harbor, at depths of 12 m or less (Bond, 1982). 

As waters warm in spring, fish move into shallow waters in 

large numbers, where they remain until autumn (Bond, 1982). 

Spawning of at least some of these species (that is, Fourhorn 

Sculpin, Arctic Flounder) occurs in winter and spring under 

ice in shallow marine waters. Arctic Smelt spawn in fresh or 

brackish waters under ice in spring, just prior to breakup.

Marine Example: Arctic Cod.—The Arctic Cod is an 

excellent example of a truly marine, although euryhaline, 

species. Arguably, this is the most widespread and abundant 

fish species in the U.S. Arctic. However, although it is 

clear that these are highly mobile and migratory fish, their 

movements, particularly seasonal movements, throughout 

much of the Arctic are poorly understood (Ponomarenko, 

1968; Craig and others, 1982; Benoit and others, 2008). 

Throughout their range, fish are abundant in nearshore waters 

in summer, the precise time varying between locations and 

between years at the same location. In the nearshore, schools 

may move quickly through an area (Craig and Haldorson, 

1981; Craig and Schmidt, 1985) or may reside in the same 

location for weeks at a time (Welch and others, 1993). How 

much of the overall population migrates into shallow waters 

is not known, although many fish may not migrate to the 

shallowest waters (Fechhelm and others, 1984; Frost and 

Lowry, 1983; Hop and others, 1997). In the Alaskan Beaufort 

Sea, Arctic Cod tend to be relatively scarce immediately after 

ice-out, but move into shallow waters as the season progresses 

especially after the first marine intrusion that typically occurs 

in mid-July following a west windstorm (Craig and others, 

1982; Palmer and Dugan, 1990; Wiswar and others, 1995). 

The high abundance of colonizing epibenthic invertebrates in 

lagoonal waters provides a rich prey base for cod and other 

marine fishes. The species also has been shown in recent 

studies to be abundant over shelf and slope environments. 
636 Alaska Arctic Marine Fish Ecology Catalog

Ecological Theory

Adaptations (anatomical, molecular, physiological, 

and behavioral) of marine fishes to life in Arctic waters are 

evidenced in strategies (genetically determined life history 

or behaviors [Gross, 1987]) and demographic characteristics 

of each population. The strategies relate to how resources are 

allocated to maximize fitness (in individuals and populations) 

and are tightly linked to life history parameters (Stearns, 

1977). The primary life history traits—size at birth; growth 

pattern; age and size at maturity; number, size, and sex ratio 

of offspring; age- and size-specific reproductive investments; 

fecundity; age- and size-specific mortality schedules; and 

life span—are poorly known for Arctic fish species. These 

parameters inform the population dynamics (survival, 

recruitment, growth rate) of each species and vary in response 

to temporal variations in ecological traits (for example, 

habitat, diet, home range, and other ecological traits) and other 

environmental factors.

Population Theory (r-K Selection)

Life history information provides important insights 

about selective forces operating on fish populations and 

their evolution to niches in marine ecosystems. In the 

absence of reliable population parameters, general studies 

of environmental effects on species adaptations are possible 

within an r-K conceptual framework5. Considering the 

postulated recency of marine ecosystem stability and related 

phylogenetic age of the Arctic fish families, r and K selection 

models are instructive. At one evolutionary extreme, the 

amphidromous fish (that is, Salmonidae) are unambiguous 

K-strategists (for example, Craig, 1989a). Amphidromy, like 

anadromy, involves the alternate occupation of freshwater 

and sea habitats by fishes. It involves complex migratory 

behaviors that ensure population fitness through predictable 

migrations of populations, or segments of populations, 

between stable habitats in freshwater, brackish, and marine 

environments. At the other end of the continuum, r-strategists, 

like the Pacific Herring and Capelin (for example, published 

values for instantaneous rate of natural mortality [M] and 

gonadosomal index [GI] from Gunderson [1980] and Rose 

[2005]) and Arctic Cod are adapted to seasonal periods of 

high food abundance followed by periods of relative scarcity. 

These small pelagic species allocate a greater proportion 

of their resources to reproduction, produce larger numbers 

of young, and have short life spans and wide niche widths 

(unspecialized). Although population sizes are unknown for 

Capelin and Arctic Cod, it is likely that they are prone to 

fluctuating stock abundance as is the pattern for similar forage 

species elsewhere. Life history information presented in the 

species accounts shows that the vast majority of Arctic species 

display mixes of r and K traits (for example, Cottidae and 

Agonidae). The mix of life history traits reported, although 

mostly incomplete, is thought to have adaptive significance 

with respect to ecological trade-offs (for example, production 

of young and longevity) and changes in M (Gunderson, 1980; 

Power, 1997).

Environmental Stability

The Arctic marine ecosystem is a relatively young system 

that is characterized by both stable (predictable) and variable 

(seasonal) environmental features in time and space. The 

small size and low densities of many species would suggest 

that many of the marine fishes are living at the abiotic and 

biotic extremes of their ranges (for example, temperatures 

between -2 and +2 °C). How body size and abundance 

co-varies with other traits (for example, temperature tolerance 

and foods) is an area of increasing interest given the rapidity 

and potential effects of changing Arctic ecosystems. The 

relative vulnerabilities (extinction selectivity, see McKinney, 

1997) of marine species to climate effects or other large-scale 

perturbations is presently difficult to assess on the basis of life 

history and ecological traits not only because information is 

lacking (that is, environmental tolerances and preferences), 

but because non-linear threshold effects (population and 

ecosystem levels) are impossible to estimate. Linked 

biological and physical models are needed to inform research 

and assessment processes.

The biological mechanisms linking environment, through 

the life history, to the population dynamics are complex. 

As harsh as Arctic environmental conditions may seem, the 

Arctic marine fishes are adapted to life therein. The presence 

of sea ice and cold water temperatures are stable features in 

deep waters and the life histories of some species, such as 

the Arctic Cod, are intricately linked to sea ice habitats. How 

variations in environmental conditions (abiotic and biotic 

factors) influence population demographics is not known for 

most species and is an important topic for monitoring longterm change. Benton and others (2006) described the potential 

interplay of these factors in terms of differential mortality and 

5

The r-K conceptual framework is useful for understanding adaptations 

to life in the Arctic marine environment. The underlying theory concerns 

population growth in stable and unstable environments and the respective 

roles of density-dependent and density-independent factors on population 

dynamics in each. Carrying capacity (K) is an attribute of the ecosystem 

(habitat) and the selection of K attributes (long-life spans, slow growth, large 

size, delayed maturity, iteroparity, few offspring, and stable mortality rate) 

reflects population growth near in stable environments. At the opposite end of 

the r-K continuum, r-selected species are adapted (short life expectancy, small 

size, early maturity, reproduces once, many offspring are produced, variable 

recruitment rate) to living in unstable, fluctuating environments and densityindependent factors (abiotic factors) affect population processes such K values 

are low and the exponential growth rate (intrinsic rate of growth rate [r]) is 

characteristic of the population. Density-dependent factors seldom come into 

play in r-selected species. The r/K concept represents a continuum and many 

species display a mix of both traits.
Chapter 4 637

survival for different life stages and resulting effects on age 

structure and abundance of populations (that is, developing 

life cycle understanding). These authors note the importance 

of population structure for two primary and related reasons: 

(1) the life cycle takes time to complete and (2) different ages 

or stages may be affected differently by environmental effects 

(that is, potential for cascading effects).

Oceanographic influences on ecosystem conditions 

and productivity are critical determinants of the distribution 

and abundance of marine fishes. Marine production cycles 

in the Chukchi Sea lead to a tightly coupled benthopelagic 

system supporting a biotic community dominated by benthic 

invertebrates and small marine fishes (Grebmeier and others, 

2006b). It is not known whether inefficient cropping of 

algal production produced beneath sea ice or in open water 

environments leads to a similar benthic pattern across the 

Beaufort Sea shelf and slope. The brackish habitat that forms 

along the north coast of Alaska (within 10–15 km of the shore) 

in early summer is often characterized as a wind-driven system 

subject to rapid changes in temperature and salinity. The 

brackish conditions are ephemeral and by late summer marine 

conditions are widespread nearshore. Terrestrial sources 

of organic matter can fuel invertebrate-driven (epibenthic 

crustaceans) food webs that are seasonally established along 

the Beaufort coast (Dunton and others, 2006; 2012). Winter 

ice, as deep as 2-m coastally, and hypersaline conditions 

in unfrozen waters, render nearshore and lagoon waters 

inhospitable to most marine fishes. The temporary conditions 

of many Arctic habitats makes them “ecological vacuums” 

for r strategists (Gunderson, 1980) and apparently many 

mixed-trait species (for example, those favoring a strategy 

of high investment in reproductive output, short generation 

times, etc.). The information presented about food habits, 

physical habitats, size, and reproductive ecology suggests 

that many species are non-specialists (opportunists), lacking 

competitive capacity (high dietary overlap in invertebrate 

foods), and expend large amounts of energy on gonadosomatic 

development at the expense of large body size.

The combinations of life history and abundance traits 

conceptually extend to fish assemblages associated with 

pioneer (developmental) and mature (climax) communities 

in Arctic marine ecosystems (Vadstein and others, 2004). 

Pioneer communities have low stability against perturbation 

and low biological control (that is, r strategists). Other 

characteristics include low diversity, wide niche width, 

and low specialization. Mature communities are typically 

dominated by K-strategists (for example, amphidromous 

species in Arctic rivers and lakes (for example, Johnson, 

1997) in stable systems with high biological control and 

high resistance (below a specific threshold) to perturbation. 

The effect of high or low diversity, in the relative sense 

(that is, generally low across the Arctic) is not known. Other 

comparisons indicate niche width is wide and specialization is 

high in mature systems.

Ecological Biogeography 

Correlative information about the relations between sea 

temperature, growth, life span, and marine fish diversity in 

Arctic conditions (generally -2 to > 6 °C) exists for many 

species. Much of this information was obtained for marine 

species studied outside the Chukchi and Beaufort Seas. 

Winter data regarding habitat requirements and fish behavior 

is generally lacking. Generalizing from the data presented in 

the species accounts (chapter 3) and section, “Adaptations in 

Marine Arctic Fishes” (this chapter), marine fish communities 

occurring in colder regimes have characteristic tendencies 

for slow growth (populations), small size, short food chains, 

and low diversity. Arctic conditions favor large species, 

populations, and predators in the marine invertebrates. 

Few species are considered apex predators (for example, 

sharks) and relatively few species are characterized by large 

populations. Arctic Cod is the most conspicuous exception 

and other commonly reported marine species (for example, 

Capelin, Arctic Sand Lance, and Fourhorn Sculpin) and 

amphidromous fishes (for example, Dolly Varden, Least 

Cisco, Arctic Cisco, and Broad Whitefish) also may have large 

population sizes. Large predators (for example, Pacific Halibut 

(Hippoglossus stenolepis), Pacific Salmon, Walleye Pollock 

(Gadus chalcogrammus), Inconnu (Stenodus leucichthys), 

and at least one shark species) are most abundant in the 

southeastern Chukchi Sea. Fewer species, such as Dolly 

Varden and possibly Arctic and Greenland flounders fit this 

description from the Beaufort Sea. No species from these high 

latitude seas is strictly piscivorous, which is indicative of the 

length of food chains and relative importance of invertebrate 

foods (including fish larvae in the zooplankton community) in 

energetic pathways.

Environmental Relations

Certain life history parameters (for example, growth 

rate, age at maturity, and fecundity) are sensitive to changes 

in temperature and food (that is, ecological conditions). This 

makes interpretation of field data difficult with respect to 

causal mechanisms and a requirement for laboratory work 

necessary (Stearns, 1977). Ongoing fieldwork on marine 

fish populations in the Chukchi and Beaufort Seas will 

develop much new information about population dynamics, 

structure, and community functions. New information about 

habitat associations will represent snap shots of ecosystem 

condition and laboratory experiments still will be needed 

to fill information gaps remaining about the physiological 

tolerances and environmental preferences of Arctic marine 

fish. Empirical data will be critical to evaluations of potential 

effects of climate change and human activities on this fauna 

(that is, natural and anthropogenic effects).
638 Alaska Arctic Marine Fish Ecology Catalog

Temperature effects are especially important as they 

directly affect growth and survival of individuals and thereby 

recruitment at the population level. In many analyses of 

field data, thermal conditions often are used as a proxy 

for mechanisms affecting survival such as enhanced prey 

production or advection of young fishes in warmer water 

masses to nursery or feeding habitats. Because thermal 

relations are often indirect, temperature is not the only 

factor affecting recruitment, and correlations often break 

down. Well-designed experiments addressing the effects of 

multiple stressors may reduce the uncertainty associated with 

population forecasts.

Regarding environmental preferences, the species 

accounts support previous finding about marine fishes 

(excluding amphidromous and anadromous forms) and 

their apparent preferences for colder, more saline waters 

(for example, Logerwell and others, 2011). In the summer, 

this includes marine waters lying seaward, or underlying, 

nearshore brackish waters along the Alaskan Beaufort Sea. 

As summer wanes, the nearshore becomes more saline due 

to decreased freshwater input from rivers and streams and 

marine intrusions associated with summer storms. During 

this time, marine fishes often share nearshore brackish waters 

with diadromous fishes, primarily to feed on the abundant 

epibenthic fauna or possibly to spawn (for example, Capelin 

and Arctic Smelt). In autumn, after most amphidromous fishes 

have returned to freshwater systems to spawn and overwinter, 

the marine fishes remain in the nearshore area to feed. 

Onshore-offshore dimensions of their migratory or movement 

behaviors are generally unknown, but for important forage 

species like Capelin and Arctic Cod could be important to 

ecosystem function.

The use of coastal habitats by Arctic Cod in Simpson 

Lagoon (near Prudhoe Bay) follows a pattern of increased 

nearshore abundance with the first major marine intrusions 

(usually mid-July), seasonal residence through early winter 

months, followed by an occupation of offshore waters during 

winter months. How much of the Arctic Cod population 

undertakes such a bathymetric migration, how it relates to 

life history patterns (spawning times and locations), and 

locations of seasonal habitats remain largely unknown for 

both the Beaufort and Chukchi Seas. There are suggestions 

that large numbers of fish remain over shelf and slope 

habitats and, given their importance in regional food webs, 

there remain important questions to be addressed. Because 

of reports of nearshore spawning (for example, summaries 

by Thorsteinson and others [1990] and Pirtle and Mueter 

[2011]), important questions about reproductive ecology and 

stock structure remain for Arctic Cod and other dominant 

species of ecological or subsistence importance (for example, 

Saffron Cod).

Concern about the potential effects of North Slope oil 

and gas development on amphidromous species (ciscoes, 

whitefish, and charr) and coastal habitats have a long-standing 

history. Because of their subsistence, and until recently 

commercial values, Arctic Cisco have received special 

research and monitoring attention. Coastal monitoring data 

from Prudhoe Bay and harvest records from the Colville 

Delta extend to the early 1980s. During the late 1980s and 

early 1990s, the effects of West Dock and Endicott gravel 

causeways in Prudhoe Bay on physical habitat conditions, 

especially temperature and salinity properties, and effects 

on the migrations and movements of amphidromous fishes, 

were a focus of many fishery studies. More than a decade 

later, in an innovative research approach, BOEM and its 

investigators worked with Nuiqsut residents (experts) in 

2003 to identify new and continuing information needs 

relative to Arctic Cisco (Murphy and others, 2007). During 

the process of expert engagement, they identified issues and 

developed study hypotheses (n = 27). Within the constraints 

of existing information, each question was addressed and 

considered regarding future priorities. A summary of the 

issues identified from local observations included: fewer fish 

available to subsistence fishery; reduced size and (or) weight 

of fish; changes in distribution of fish in the Colville River; 

changes in the quality of fish (taste, texture, color); deformed 

fish; food chain effects; and energetics stress. These changes 

were thought to be related to changes in habitat associated 

with local industrial developments such as seismic noise, 

causeways and coastal siltation, ice roads, contaminants, 

and dredging, or larger scale processes such as ocean and 

climate changes. The possibility of over harvesting also was 

considered. Although causeway effects were still of concern, 

the study was done after a period of oil and gas development 

in the Colville Delta, and this explains the expanded list of 

local concerns.

The Murphy and others (2007) study was unique in 

its comprehensive evaluation of environmental influences 

on Arctic Cisco and the participation of Nuiqsut residents 

from beginning to end. Environmental influences were 

geographically partitioned into three regions including the 

MacKenzie River Region, Coastal Beaufort Region, and 

Colville-Prudhoe Bay Region to investigated population 

effects on immature ciscoes (age 0–5 and 5–7). Environmental 

factors analyzed included: MacKenzie River—river discharge, 

air temperatures, regional sea surface temperatures (SSTs), 

spring ice concentration, and a combined climate index 

(temperature, ice, discharge); Coastal Beaufort Region—

winds, regional SSTs, and ice concentrations; and ColvillePrudhoe Bay Region—air temperature, river discharge, 

SSTs, sea surface salinities, autumn salinities (delta), ice 

concentration, Arctic Oscillation (AO), flow through the 

Bering Strait, and the combined climate index. 

The results of the analysis reconfirmed, in part, the 

importance of east wind conditions on the recruitment 

of young-of-the-year Arctic Cisco to the Colville Delta. 
Chapter 4 639

Murphy and others (2007) reported that 80 percent of the 

natural variation in recruitment could be explained by 

these winds. Regional changes in sea ice conditions, winds, 

coastal circulation, and upwelling are known to affect the 

geographic extent and duration band and the quality and 

possibly coastal access to habitat by amphidromous fish; these 

relationships were examined by Murphy and others (2007). 

A weak correlation was established between survival rates 

(age 0–5) and the AO, possibly related to changes in regional 

circulation. Warming influences on the timing of breakup 

and river discharges were studied in the Colville-Prudhoe 

Bay Region and a relation between increased discharge and 

decreased survival in age 5–7 ciscoes was described. Changes 

in hydrology and changes in regional upwelling and nearshore 

prey conditions and competition were suggested as causes 

of the increased mortality and decreased fish condition. 

Insufficient information was available to evaluate the effects 

of siltation effects in coastal habitats in the West Dock area, 

effects of contaminants (for example, heavy metals), effects 

of autumn salinities on cisco distributions in delta habitats, 

or effects of seismic activities (noise and vibration) on fish 

distribution and abundance. A key finding for Arctic Cisco, 

and other marine fishes, is the demonstration of how physical 

drivers interact to affect coastal habitats at different time and 

spatial scales with differential effects to species throughout 

their life cycles.

With respect to Arctic Cisco, Murphy and others (2007) 

noted the lack of population information for this species 

from the MacKenzie River and need for greater resolution 

of the age structure of the population in the Colville River. 

Murphy and others (2007) also noted the need for increased 

understanding about AO effects on regional marine 

ecosystems. In response, the BOEM and USGS began 

researching the genetics and age of fishes in the Colville Delta 

and a broader study on the influences of AO on recruitment 

processes in young-of-the-year fish (von Biela and others, 

2011; Zimmerman and others, 2013). The genetics research 

confirmed the MacKenzie River origins of Colville River 

ciscoes and the aging studies indicated the presence of small 

numbers of older-aged fishes than previously reported (that 

is, the presence of fishes to age 11). Large-scale effects of 

AO on the MacKenzie River focused on annual variability in 

coastal discharge and interannual effects of marine ecosystem 

conditions on the abundance and condition of euphausiid 

prey in coastal plumes. With respect to the latter, von Biela 

and others (2011) hypothesized that prior year conditions (to 

the arrival of age-0 migrants), are critical to the growth and 

condition of euphausiids and prey concentrations in the next 

year. The temporal connection between euphausiid ecology 

and early marine survival and growth in Arctic Cisco adds 

significant new information about another source of natural 

variation in the recruitment of these fish into Alaskan waters.

Fish Assemblages

DeGange and Thorsteinson (2011) used an earlier 

classification scheme (Mecklenburg and others, 2008; 

Minerals Management Service, 2008) to describe primary 

and secondary marine fish assemblages in the Chukchi 

and Beaufort Seas. The DeGange and Thorsteinson (2011) 

classification scheme is consistent with the life history profiles 

presented herein (for example, vertical distributions) and with 

respect to partitioning the primary assemblage (all species) 

into ecological categories. The ecological categories broadly 

define fish assemblages based on existing knowledge of 

fish behavior and ecology and associations with prominent 

oceanographic features (for example, seafloor, shelf break, 

pack ice, and other features). In this analysis, the ecological 

categories used by DeGange and Thorsteinson (2011) 

were re-defined to include regional distinctions in habitat 

and dominant fauna associated with nearshore (<5 m) and 

offshore areas (>5 m) of the continental shelf. This separation 

of inshore and offshore communities is more consistent 

with hydrographic properties (water mass distributions) of 

the Chukchi (for example, Norcross and others, 2009) and 

Beaufort Seas (for example, Craig, 1984; Logerwell and 

others, 2011). A nearshore assemblage was added to the 

classification and subdivided into three subregions to capture 

faunal differences in Kotzebue Sound, in the Barrow area, and 

in brackish coastal waters along the Alaska North Slope to the 

east of and including Harrison Bay. 

The definitions of regional assemblages now includes: 

(1) nearshore assemblages (in shallow bays and lagoons 

and along open exposed coastlines); (2) neritic-demersal 

assemblage (at or near the seafloor of the continental shelf); 

(3) neritic-pelagic assemblage (in the water column over the 

continental shelf); (4) oceanic-demersal assemblage (living 

on or close to the bottom beyond the continental shelf break at 

200 m); (5) oceanic-pelagic assemblage (inhabiting the water 

column of oceanic waters seaward of the continental shelf 

break); and (6) cryopelagic assemblage (inhabiting neritic 

or oceanic waters, but during their life cycle, are associated 

with sea ice). The dominant species associated with these 

assemblages are listed in table 4.5.

A cryopelagic assemblage was not included in table 4.5

and is considered separately due to the special role of Arctic 

Cod in sea-ice community dynamics. The cryopelagic 

assemblage and role of the Arctic Cod in the marine 

ecosystems are unique to the Arctic (Bradstreet, 1982). The 

term “cryopelagic” describes fishes that actively swim in 

neritic or oceanic waters but, during their life cycle, are 

associated with drifting or fast ice. Both young and adult 

fishes can be associated with ice or water immediately beneath 

the ice. These relationships can be trophic in nature or related 

to refuge and possibly reproduction and nursery for young. 
640 Alaska Arctic Marine Fish Ecology Catalog

Table 4.5. Marine fish assemblages in the Chukchi and Beaufort Seas.

[Marine fish assemblages: Cyropelagic assemblage is described in text. Neritic-pelagic: Salmon, Pacific Herring, and Yellowfin Sole from southeastern 

Chukchi Sea]

Community 

dominants

(family/species)

Marine fish assemblages

Nearshore Neritic-Demersal Neriticpelagic

Oceanicdemersal Southeast Northwest North Slope Chukchi Beaufort

Petromyzontidae Arctic Lamprey

Squalidae Spiny Spotted 

Dogfiish

Rajidae Arctic Skate Arctic Skate1

Clupeidae Pacific Herring Pacific Herring

Osmeridae Arctic Smelt,

Pacific Capelin

Arctic Smelt,

Pacific Capelin

Arctic Smelt,

Pacific Capelin

Pacific Capelin,

Arctic Smelt

Pacific Capelin 

Salmonidae Chum Salmon,

Pink Salmon,

Least Cisco,

Inconnu,

Dolly Varden

Pink Salmon,

Least Cisco,

Dolly Varden

Dolly Varden,

Arctic Cisco,

Least Cisco,

Bering Cisco,

Broad 

Whitefish,

Humpback 

Whitefish

Chum Salmon,

Pink Salmon

Gadidae Arctic Cod,

Saffron Cod,

Walleye Pollock

Arctic Cod Arctic Cod Arctic Cod,

Saffron Cod,

Walleye Pollock

Arctic Cod,

Saffron Cod,

Ice Cod,

Walleye Pollock

Arctic Cod,

Saffron Cod,

Ice Cod

Arctic Cod,

Ice Cod

Gasterosteidae Ninespine 

Stickleback

Hexagrammidae Whitepotted

Greenling

Cottidae Hamecon,

Arctic Staghorn 

Sculpin,

Fourhorn 

Sculpin,

Ribbed Sculpin

Antlered 

Sculpin,

Fourhorn 

Sculpin

Fourhorn 

Sculpin,

Twohorn 

Sculpin

Hamecon,

Arctic Staghorn 

Sculpin,

Arctic Sculpin,

Shorthorn 

Sculpin,

Ribbed Sculpin,

Eyeshade 

Sculpin

Hamecon,

Arctic Staghorn 

Sculpin,

Spatulate 

Sculpin,

Ribbed Sculpin,

Eyeshade 

Sculpin,

Fourhorn 

Sculpin

Polar Sculpin,

Bigeye 

Sculpin,

Atlantic 

Hookear 

Sculpin1 

Agonidae Arctic 

Alligatorfish

Veteran 

Poacher

Arctic 

Alligatorfish

Arctic 

Alligatorfish

Liparidae Nebulous 

Snailfish,

Variegated 

Snailfish,

Kelp Snailfish

Gelatinous 

Snailfish,

Variegated 

Snailfish,

Kelp Snailfish,

Nebulous 

Snailfish

Gelatinous 

Snailfish,

Variegated 

Snailfish

Slender 

Eelblenny,

Sea Tadpole
Chapter 4 641

Community 

dominants

(family/species)

Marine fish assemblages

Nearshore Neritic-Demersal Neriticpelagic

Oceanicdemersal Southeast Northwest North Slope Chukchi Beaufort

Zoarcidae Marbled 

Eelpout,

Saddled Eelpout,

Polar Eelpout,

Fish Doctor

Marbled 

Eelpout,

Polar Eelpout,

Fish Doctor

Adolf’s 

Eelpout,

Longear 

Eelpout

Stichaeidae Slender 

Eelblenny

Arctic Shanny,

Slender 

Eelblenny,

Stout Eelblenny,

Fourline 

Snakeblenny

Arctic Shanny,

Fourline 

Snakeblenny

Daubed 

Shanny

Ammodytidae Arctic Sand 

Lance

Arctic Sand 

Lance

Arctic Sand 

Lance

Arctic Sand 

Lance

Arctic Sand 

Lance 

Pleuronectidae Yellowfin Sole,

Starry Flounder

Longhead Dab Arctic 

Flounder

Bering Flounder,

Yellowfin Sole,

Alaska Plaice

Bering 

Flounder,

Greenland 

Halibut,

Arctic Flounder

Bering 

Flounder,

Yellowfin Sole

Greenland 

Halibut

1

Fishery data suggest occurrence in assemblage.

Table 4.5. Marine fish assemblages in the Chukchi and Beaufort Seas.—Continued

[Marine fish assemblages: Cyropelagic assemblage is described in text. Neritic-pelagic: Salmon, Pacific Herring, and Yellowfin Sole from southeastern 

Chukchi Sea]

Andriyashev (1970) was among the early investigators to find 

Arctic Cod in abundance in marine habitats associated with 

broken ice or near the ice edge. In spring, as ice thaws and 

breaks up, phytoplankton blooms and zooplankton colonize 

these habitats and provide food for Arctic Cod. The substrate 

formed by the underside of sea ice supports an amphipoddiatom ice community that is believed to provide an important 

source of food for cod. Other species, such as Capelin and 

Pacific Herring have been mentioned in this assemblage, and 

this relates to limited observations in broken-sea ice conditions 

or near the ice edge. Ecological linkages for these fish species 

with sea ice, and others, and the importance of polynyas 

(biological hotspots), are not known.

Diadromous fishes (migratory fishes), especially 

salmonids, are prominent members of nearshore fish 

assemblages and represent an important source of traditional 

foods for Arctic villagers. Diadromy involves regular 

occupation of fresh, brackish (5–10 °C, <15 psu), and marine 

waters. Fish migrations to, or movements between, reflect the 

response of a population (or segment of the population) to 

various biological stimuli, such as feeding or reproduction, 

or ecological factors, such as temperature, oxygen level, or 

specific spawning-habitat need. Diadromous fishes display 

a wide range of migration behaviors including once-ina-lifetime events, repetitive multiyear events, spawning 

migrations, feeding migrations, and seasonal movements 

between environments. Diadromous fishes inhabit various 

habitats including many of the lakes, rivers, streams, 

interconnecting channels, and coastal waters of the North 

Slope. Common amphidromous species include Arctic, Least, 

and Bering Ciscoes; Arctic Smelt; Humpback and Broad 

whitefishes; Dolly Varden; and Inconnu. Common anadromous 

species include Pink and Chum salmon, Ninespine 

Stickleback, and Arctic Lamprey. 

The highest densities and diversity of amphidromous 

fishes occurs in river-deltas, such as the Colville and the 

Sagavanirktok Rivers along Alaska’s North Slope. Pacific 

Salmon and Arctic Lamprey are in greatest abundance south of 

Point Lay and in Kotzebue Sound in the southeastern Chukchi 

Sea. Lakes that are accessible to the sea typically are inhabited 

by diadromous fishes and many species have evolved resident 

forms (for example, Dolly Varden and Arctic Lamprey). The 

Least Cisco is the most abundant species in sea-connected 

Arctic lakes. With the exception of the Arctic Cisco, the 

amphidromous species are characterized by a variable 

freshwater residence for juvenile fishes that lasts 1–3 years 

before they undertake their first migration to sea. Fish and 

invertebrate prey are seasonally more abundant in coastal 

waters than in freshwater habitats; therefore, amphidromy 

represents a physiological adaptation to food availability 

(Craig, 1989a).
642 Alaska Arctic Marine Fish Ecology Catalog

The primary marine fish assemblage found over the 

continental shelf is composed of relatively few species of 

small size and biomass when compared to similar ecosystem 

attributes of lower latitudes (Irvine and Meyer, 1989). 

As an example, most species are less than 15 cm (FL). Small 

schooling species such as the Arctic Cod can be abundant 

over shelf waters and near the shelf break (Crawford and 

others, 2012). Ecological processes (for example, Smetacek 

and Nicol, 2005; Grebmeier and others, 2006a, 2006b; Jay 

and others, 2011) and associations of fishes in Alaska coastal 

water, Pacific summer water, and Pacific winter waters (for 

example, Crawford and others, 2012) provide stable habitat 

conditions for shelf-affiliated assemblages, especially adult 

fishes, in the Chukchi Sea. The intermingling of these water 

masses with Atlantic water along the shelf break, slope, and 

deep waters of the Chukchi and Beaufort Seas results in 

conditions that are more inhospitable and fewer fish. The use 

and importance of fish habitats in river deltas and coastal 

lagoons have received the most research attention. The relative 

importance of other habitats such as those along exposed 

coastlines, near the ice edge and in polnynas, or located 

at special features, such as the Hannah Shoal, for feeding, 

reproduction, and other life activities have been postulated, but 

in most instances, remains to be described. 

Trophic Relations

The available food habits data (stomach contents, fatty 

acids, isotopes) from across all habitats reviewed in chapter 3

suggest that most Arctic marine fishes are primary carnivores 

that feed across trophic levels T2 and T3. Invertebrates 

are significant in their diets with pelagic species feeding 

on zooplankton herbivores compared to demersal species 

preying more frequently on early life history stages of benthic 

detritivores. Freshwater invertebrates swept downstream 

can be an important source of food for anadromous and 

amphidromous species as they move downstream into the 

coastal sea.

Trophic level status is an important indicator of diversity 

or ecosystem change (that is, changes in community, animal 

size, and consumption in pelagic and benthic environments) 

and has been used to study fishing and other effects on 

species abundance in the northeast Pacific Ocean. More than 

90 percent of Arctic marine fauna are secondary consumers. 

Secondary consumers feed on various fish and invertebrate 

prey by feeding across trophic levels. The trophic position 

thus occurs at intermediate levels of the marine ecosystem. 

The average trophic level position for all Arctic marine fishes 

(juveniles and adults combined) is 3.27 (range: 2.99–4.6)6

. 

Of the tertiary consumers, only two species—Arctic Lamprey 

and Pink Salmon—are common in their occurrence in the 

southeastern Chukchi Sea. 

The trophic positions of each species were estimated 

in FishBase (Froes and Pauly, 2012) and many assumptions 

relative to availability and recency of data are involved. 

Despite these limitations, the mid-level positions indicated 

are considered to be reasonably accurate. This determination 

is based on the widespread consumption of small invertebrate 

prey, benthic and pelagic species, almost universally 

reported in most species diets. The stomach contents indicate 

significant dietary overlaps in co-habitating species in marine 

and nearshore habitats. The abundance of invertebrate foods 

in each habitat type is related to carbon cycling associated 

with the delivery of marine-derived organic matter (primary 

production) to benthic systems offshore and transport of 

terrestrially derived carbon to nearshore systems. Of interest, 

the different life strategies that have evolved in the Arctic 

for marine fishes (anadromy, amphidromy, and marine) are 

adapted to an invertebrate food base and many species seem 

to lack specialization with respect to feeding behavior and 

diet. The generalist strategy may be an effective adaptation 

of marine fishes living in the invertebrate-dominated shelf 

systems of the Chukchi and Beaufort Seas. Although most 

marine fishes are lower-order carnivores, some of the larger, 

especially pelagic, predators are highly piscivorous during 

adult stage phases of their lives (for example, Pacific salmon 

and Inconnu). The absence of exclusive tertiary consumers 

and relatively few secondary consumers that cross over the 

T4 level (feed upward) are indicative of the pelagic-benthic 

coupling and energy pathways characteristic of invertebratedominated shelf environments.

Most Arctic marine fishes appear to feed at similar 

trophic levels and on similar prey comprising the trophic 

spectrum of available food and prey items available to 

them in near- and off-shore habitats. It is hypothesized that 

competition for food resources with invertebrate competitors 

on the shelf may, in concert with environmental temperature, 

contribute to observed abundance patterns and the 

hypothesized ecological redundancy of many cryptic species. 

In brackish marine environments, such as the open and pulsing 

lagoons along the North Slope, epibenthic foods (for example, 

amphipods and mysids) are seasonally abundant, tend to drive 

coastal food webs, and are not considered to be limiting. 

Competition between fish competitors in the nearshore marine 

such as between Arctic and Saffron cods may be significant 

in some locales and intensified in others through shifts in 

distribution or changes in ecosystem function brought forth 

by climate change. Predation pressures on the slope by marine 

fish, especially on benthos, were not historically considered 

great (for example, Carey and Ruff, 1977). However, recent 

data suggesting large standing stocks of Arctic Cod on the 

slope suggest a need for a re-examination of this hypothesis.

6

According to Romanuk and others (2011) omnivores, which consume 

plants or detritus as well as animals, have trophic levels between 2.2 and 2.79 

and carnivores have trophic levels greater than 2.8. Secondary consumers 

(carnivores) have trophic levels between 2.8 and 4.0 and tertiary consumer 

levels are greater than 4.0.
Chapter 4 643

Relative Abundance

Spatial patterns of relative abundance for nearshore and 

shelf fish assemblages can be generalized from a compilation 

of regional fishery data and related to the diversity estimates 

(table 4.1), characterization of marine fish assemblages 

(table 4.5), and advection model developed by Grebmeier and 

others (2015, presented in chapter 1 of this report) (table 4.6). 

The species composition of the nearshore was historically 

known from fish catches in passive gear collections (that is, 

fyke nets and gill nets) from inshore habitats and expressed 

in indices of catch-per-unit-effort (CPUE). Sampling over 

the shelf has involved active sampling methods including 

bottom trawling (near the bottom), seining, and plankton tows 

(surface and water column). The fish catches reported from 

active sampling gears is expressed in densities of fish (in 

numbers or biomass) per area trawled or volume sampled. The 

various sampling and enumeration methodologies required 

development of a simple, unifying classification of abundance 

(rare, uncommon, and common) to describe regional patterns 

in the marine fish assemblage (chapters 2 and 3). 

The diversity of Salmonidae species throughout the 

Pacific Arctic Region (table 4.6) reflects the plasticity of this 

group and its adaptive linkage to freshwaters. All five species 

of Pacific salmon are abundant in the northern Bering Sea 

having important spawning grounds in the Yukon and Anadyr 

Rivers. In the west, sockeye salmon are more abundant than in 

Norton Sound, USA in the east relating to spawning habitats 

in the Anadry River, Russia, and its tributaries. North of the 

Bering Strait, Pink and Chum salmon are the abundant salmon 

species although successful colonization has not demonstrated 

in Alaskan habitats north of Kotzebue Sound. Coho Salmon 

are present in Kotzebue Sound drainages and along the 

Chukotka coast.

Many of the common species, or functional groups of 

species, are benthic, or demersal, in their habitat orientation 

(table 4.6). Densities are low and small-sized fishes such 

as snailfish, poachers, and pricklebacks predominant over 

shelves. The small size and low densities of many species 

suggests that they may be living at the abiotic and biotic 

extremes of their ranges (for example, temperatures between 

-2 and +2 °C). How body size and abundance co-varies with 

other traits (for example, temperature tolerance and foods) is 

of increasing interest given the rapidity and potential effects 

of changing Arctic ecosystems. The relative vulnerabilities 

(extinction selectivity, see McKinney, 1997) of marine species 

to effects of climate or other large-scale perturbations are 

presently difficult to assess on the basis of life history and 

ecological traits not only because information is lacking (that 

is, environmental tolerances and preferences), but because 

non-linear threshold effects (population and ecosystem levels) 

are not possible to describe. 

These abundance patterns, and the classification used, 

are consistent with published literature and, the high number 

of rare and uncommon species reported raises questions 

about their functional roles in the marine ecosystem. 

Biodiverse environments were described by Mouillot and 

others (2013) as having large numbers of rare species whose 

functional importance in ecosystems is largely unknown. 

When the diversity of marine invertebrates is considered, 

the U.S. Arctic waters can be characterized as biodiverse. 

The occurrence of roughly 30 percent of the marine fishes 

from catches in the Beaufort Sea and 20 percent from the 

Chukchi Sea is described as rare. Even greater numbers are 

considered uncommon. Only 41 percent of the marine fishes 

found in the Beaufort Sea, and 53 percent in the Chukchi 

Sea, are described as common. According to Mouillot and 

others (2013), rare species in species-rich areas are generally 

considered to have little influence on ecosystem functioning 

and may fulfill the same ecological roles as common species. 

Because of their low abundance, they have less effect, a 

phenomenon known as functional redundancy (Mouillot and 

others, 2013). The redundancy hypothesis suggests that rare 

species serve as an insurance policy for the ecosystem in the 

event of severe disruption or regime shift and ecological loss. 

Thus, the ecological role of the individual species, especially 

in benthic ecosystems in the Arctic where invertebrates 

predominate and many marine fish are rare or uncommon, 

is probably best assessed for the demersal assemblage as 

a whole, rather than single species contributions to food 

webs and energy flows. Disruptions, such as the cascading 

effects of ecosystem change reported by Grebmeier and 

others (2006a; 2006b) related to climate change, would tend 

to favor pelagic species (for example, salmonids and cods) 

and potential switch from an invertebrate-dominated system 

to a gadid-dominated system. In this scenario, the pelagic 

community would be enriched through increased productivity 

and immigration at the potential expense of abundant 

invertebrates such as snow crabs. The potential effect on 

rare and uncommon fishes could be positive (reduced 

competition for foods) or negative (less food available). If 

the hypothesized effects of the cold water pool in the Bering 

Sea are true, this thermal barrier may protect many of the low 

density, benthic fishes living in the Arctic from extinction or 

northward shifts in the decades ahead. 

The disruption of the marine food chain could 

lead to reduced growth rates, lower abundance of some 

marine fish populations, and restructuring of the fish 

assemblages. Regime shift effects on pelagic and benthic 

ecosystems, as hypothesized by Grebmeier and others 

(2006a; 2015), are driven by decoupling mechanisms 

(for example, influx of heat in Pacific waters, changing 

sea ice conditions, reduced transport of organic matter 

to the benthos, and remineralization), which result in 

the destabilizing of shelf environments (for example, 

increased use of pelagic habitats by salmon, reduced benthic 

productivity and abundance of invertebrate foods, changes in 

animal distribution relative to prey conditions) from current 

conditions. In an analysis of potential effects of climate 

change, Mueter and others (2009, p. 108) concluded that 
644 Alaska Arctic Marine Fish Ecology Catalog

Table 4.6. Most abundant species of Arctic marine fish in northern Bering Sea, Chukchi Sea, East Siberian Sea, and Beaufort Sea 

regions of the Pacific Arctic Region.

[Pacific Arctic Region is as described by Grebmieir and others (2015). Habitat/shelf region: Pelagic refers to species occurrence in water column; demersal 

refers to species occurrence at or near the seafloor]

Northern Bering and Chukchi Seas

Arctic Sea Habitat/shelf region

Fish abundance

East Community dominants and 

species groups West

Nearshore Offshore Nearshore

Northern Bering Pelagic Pacific Salmon, Dolly 

Varden, Pacific Herring, 

Arctic Smelt, Capelin

Pacific Salmon, Pollock, 

Pacific Herring

Pacific Salmon, Dolly 

Varden, Pollock, Arctic 

Smelt 

Demersal Arctic Lamprey; Starry 

Flounder and other 

flounders; Pollock, 

Arctic and Saffron Cods; 

pricklebacks

Pollock; Pacific Cod; 

Pacific Halibut; Bering 

Flounder, Yellowfin 

Sole, Alaska Plaice and 

Northern Rock Sole; 

Capelin; Pacific Herring; 

eelpouts; snailfish

Pollock, Pacific Cod, 

Saffron Cod, flounder, 

sculpins

Chukchi South Pelagic Chum and Pink Salmon; 

Pacific Herring; Capelin, 

Arctic Smelt; Least 

Cisco; Inconnu 

 Chum and Pink Salmon; 

Dolly Varden; Pacific 

Herring

Chum and Pink Salmon, 

charr, Least Cisco

Demersal Arctic Lamprey; Arctic 

Cod; Saffron Cod; 

Pollock; Hamecon, 

Arctic Staghorn and 

other sculpins; Arctic 

Sand Lance; Yellowfin 

Sole; Starry Flounder; 

Whitespotted Greenling

Spiny Dogfish; Arctic 

Skate; Arctic and 

Saffron Cods; Arctic 

Sand Lance; Bering 

Flounder; Yellowfin Sole, 

Alaska Plaice and other 

flounders; numerous 

species of eelpouts, 

pricklebacks, poachers, 

and snailfish

Chum and Pink Salmon, 

charr, Least Cisco

North Pelagic Pacific Herring, Capelin Arctic and Saffron Cods

Demersal Arctic Cod, Four Horn 

Sculpin, Veteran Poacher, 

Arctic Sand Lance, 

Arctic Flounder

Arctic Skate; Arctic and 

Saffron Cods; Arctic 

Sand Lance; Bering 

Flounder; Yellowfin Sole, 

Alaska Plaice and other 

flounders; numerous 

species of eelpouts, 

pricklebacks, poachers, 

and snailfish

Chum and Pink Salmon, 

charr, Least Cisco
Chapter 4 645

East Siberian and Beaufort Seas

Arctic Sea Habitat/shelf region

Fish abundance

East Community dominants and 

species groups West

Nearshore Offshore

Beaufort Pelagic Dolly Varden; Arctic and Least Ciscoes, 

Broad Whitefish; Pacific Capelin; 

Arctic Smelt; Ninespine Stickleback; 

Pacific Herring

Arctic Cod; larval and juvenile forms of 

demersal fish species

Demersal Arctic Cod; Fourhorn and Twohorn 

Sculpins; Arctic Flounder; Slender 

Eelblenny; snailfish

Arctic Skate; Pacific Capelin; Arctic 

Smelt; Arctic Cod; Saffron Cod; Arctic 

Sand Lance; Greenland Halibut; Arctic 

Flounder; Hamecon, Fourhorn, Arctic 

Staghorn, and other sculpins; snailfish, 

eelpouts, blennies

East Siberian Sea Pelagic Char, Arctic and least ciscoes, broad 

whitefishes, Arctic smelt 

Meroplankton (fish larvae)

Demersal Saffron and Arctic Cods, Arctic Flounder, 

sculpins

Bering Flounder and other founders, 

sculpins, eelpouts, snailfish

Table 4.6. Most abundant species of Arctic marine fish in northern Bering Sea, Chukchi Sea, East Siberian Sea, and Beaufort Sea 

regions of the Pacific Arctic Region.—Continued

[Pacific Arctic Region is as described by Grebmieir and others (2015). Habitat/Shelf region: Pelagic refers to species occurrence in water column; demersal 

refers to species occurrence at or near the seafloor]

“...further shifts in spatial distribution and northward range 

extensions are inevitable and that the species composition of 

the plankton and fish communities will continue to change 

under a warming climate. While overall productivity is 

likely to increase in the northernmost systems as the region 

changes from Arctic to subarctic conditions, some (primarily 

subarctic) species will increase, while other (primarily 

Arctic) species will decrease or retreat northward.” These 

analyses demonstrate linkages between climate and species 

distributions and, when considered in light of sea-ice retreat 

and marine ecosystem effects toward the poles (for example, 

Doney and others, 2012), suggested a heightened vulnerability 

of the pelagic fish community over the Chukchi shelf to 

widespread environmental change.

The life cycles of marine fishes (eggs, larvae, juveniles, 

and adults) variously link pelagic and benthic systems and 

nearshore and offshore through evolved life strategies and 

behavioral patterns. Many shelf species have a benthic life 

history stage and depend on the benthic environment for some 

part, or all of their lives. Cascading effects from pelagic to 

benthic ecosystems, without immediate additions of demersal 

fishes from the Bering Sea, are possible, and already may be 

occurring. Changes in productivity, competition, and predation 

as noted by Mueter and others, (2009) would accompany 

changes in habitat and affect population processes including 

growth and survival. The different species (for example, 

salmonids, forage fishes, flatfishes, and cods) will be affected 

differently by changes in the pelagic community. For instance, 

flatfishes live on the ocean bottom and typically release eggs 

into the plankton drift. Most eelpouts and sculpins dwell on 

the bottom and their eggs, larvae, and juveniles develop on the 

bottom. Saffron Cod adults occur at various depths; however, 

they lay their eggs on the bottom and the eggs develop there. 

Key information gaps about the abundance, life history, 

migrations, and reproductive ecology (for example, spawning 

times and locations) of Arctic Cod, Capelin, and other forage 

species will be needed to inform assessments of trophic 

cascades and effects of ecosystem change.

The fish assemblages of nearshore and shelf waters 

of the Chukchi and Beaufort Seas are under current fishery 

investigation (appendix A). Previous research has shown 

that, although many marine species such Capelin and Arctic 

Sand Lance commonly are detected in nearshore and offshore 

habitats, Arctic Cod are by far the most common marine 

species in the U.S. Chukchi and Beaufort Seas. In reviewing 

the life history and ecology of Arctic Cod, Crawford and 

others (2012) reported temperature preferences of early life 

history stages to be the same as those associated with cold 

marine waters (that is, larvae and fry development between 

1.5 and 3.0 °C with highest survival rates between 0.5 and 

3.0 °C). Juveniles and adults were tolerant of a wider range 

in temperatures conditions. Adults have been sampled in 

abundance in coastal waters >13 °C and in marine habitats 

typically no warmer than 6–7 °C. Although specific physical 

and biological information about migratory behaviors, 

reproductive ecology, and importance of offshore habitats for 

this species is lacking, its keystone role is supported by its 

widespread occurrence, dominance in fish assemblages, and 

importance in regional food webs. 
646 Alaska Arctic Marine Fish Ecology Catalog

Population Dynamics

Estimating how marine fish populations respond to 

changing environments requires an understanding of the 

mechanisms by which the environment (its mean and 

variability) creates changes in population dynamics. A 

life cycle approach that includes a numerical assessment 

of differential effects at different ages or stages enhances 

predictive capability and understanding of the contribution of 

individual effects to the population level (Benton and others, 

2006). Environmental effects are mediated through changes 

in demographic rates and population size, which may result 

from functional responses at molecular and genetic levels 

of organization. In population ecology, the most important 

environmental factors are those that have the greatest effect 

on mortality, growth, and spawning and recruitment rates 

(Reist and others, 2006; Allen and Hightower, 2010). In the 

U.S. Arctic, research has addressed environmental effects 

on physiology (food and growth)7

, dispersal (active and 

passive transport mechanisms), and predation mortality

(predator-prey). How physical influences (for example, sea 

ice, light, temperature, salinity, currents, pH, and dissolved 

oxygen [DO]) affect these processes (directly or indirectly) 

and ultimately the distribution and abundance of fishes is an 

increasingly important part of ecosystem-based management 

and a focus of integrated science. The role of disease and 

parasitism in Arctic marine fish population dynamics, 

although thought to be significant, are unknown and baseline 

information has not been included. At present, the effects of 

fishing mortality on Arctic marine fishes are of negligible 

effect to their population dynamics.

Population Parameters and Models

Population size, the number of individuals within a 

geographic range, is a fundamental demographic parameter. 

In surveys of marine fishes, it is not possible to count all 

individuals in a census and therefore a number of techniques 

have been developed to estimate abundance and monitor 

change. These methods commonly include estimates of 

population density, the size of the population in relation 

to the space it occupies. Population density is affected by 

density-dependent factors (competition, predation, migration, 

and disease) and density-independent factors (for example, 

weather, storms, currents, temperature). The structure 

of populations often is studied with respect to age- or 

size-related cohorts (for example, juveniles or adults) and 

related distributions of gender (sex ratios) to describe the 

reproductive potential of the population and its current and 

future growth. Genetic information is commonly obtained 

to further characterize population structures and elucidate 

evolutionary relationships. Age- and stage-structured analyses 

show how some individuals will have a greater effect on 

population-level processes. In this context, fecundity is an 

important parameter that describes the numbers of offspring an 

individual or population is able to produce in a given amount 

of time. In marine fishes, fecundity (number of eggs) varies by 

species and age/size of fish with younger smaller fish typically 

producing fewer eggs, and larger, older fish producing more 

eggs. Environmental conditions often affect the physiological 

condition of a fish and the number and quality of eggs 

produced, and for some species whether or not spawning 

occurs. Another individual trait that affects population growth 

is mortality, the measure of deaths per unit of time. Life tables 

are regularly used to display age-specific statistical patterns of 

a population’s survival patterns.

Many different kinds of models (for example, exponential 

and logistic) have been developed to study growth in fish 

population to aid fisheries management, environmental impact 

assessment, or restoration of stocks that have been exploited 

or somehow reduced. Stock models range from surplus 

production models (for example, Schaefer, Fox, and Pella 

and Thomlinson models) to more detailed and age-structured 

demographic models (for example, yield-per-recruit, delay 

difference, and Beverton and Holt stock-recruitment models. 

The most common uses of population models have been to 

estimate recruitment (young fish entering the population) 

and yield (harvestable amount) in managed fisheries. The 

greatest challenge in these model systems is in their ability to 

realistically capture the “complex causation driving population 

dynamics”; the greatest value may be in the identification 

of the general mechanisms, which map the environment, 

through the life history, to the dynamics (from Benton and 

others, 2006, p. 1,173). In most instances, the scattered and 

fragmented nature of the existing Arctic data sets and lack 

of time series does not allow quantitative approaches to 

population modeling. 

The ideal is to have the capability to estimate 

(1) long‐term fluctuations in abundance and biomass 

production due to human perturbations or environmental 

variations, and (2) temporal-spatial responses of fish 

populations to change (Monterio, 2002). Surplus production 

models generally require much less biological information 

than age-structured models. In data-rich regions, like the 

southeastern Bering Sea, the role of the physical environment 

in predictions of resource availability is increasingly the focus 

of ecological (coupled physical and biological processes) 

models (for example, Miller, 2007). 

7

The studies stop short of linking bioenergetics (energy expenditures) and 

population dynamics (mortality, survival, and population growth).
Chapter 4 647

Measures of growth, such as age-length and lengthweight relationships are often used in stock assessments, in 

models that estimate habitat or ecosystem productivity, and 

in assessments of the health of individuals. However, it is 

not presently possible to model the population dynamics of 

most Arctic marine fishes with actual data collected from 

the Chukchi and Beaufort Seas. Regional-scale resource 

assessment surveys have been recently designed to collect 

data for the purpose of parameter estimations (that is, BOEMsponsored NOAA and UAF surveys). Earlier baselines surveys 

produced information about species occurrences, relative 

abundance, and habitats. The collection of population data 

has, in the whole, been somewhat limited to length-weight 

and in some instances, age-length (size-at-age) relationships. 

In appendix B, age-length and length-weight relationships are 

presented for 19 nearshore species of Arctic fishes. In fishes, 

both growth rates and length-weight relationships are highly 

variable reflecting differential changes in population responses 

to environmental parameters such as water temperature, prey 

quality, population density, and direct human interaction, 

such as fishing (Courcelles, 2011; Loher, 2012). Species 

specific and interspecific differences in population growth 

rates and age and weight relationships are evident within 

and across years and by geography from various locations in 

Chukchi and Beaufort Sea study areas (appendix B). These 

differences demonstrate the plasticity in age and growth noted 

by Courcelles (2011) and Loher (2012) in Arctic marine 

fishes. The sources of age-length data and regression analysis 

are appended to this report (appendix B) and the species and 

locations of data collection from outside the U.S. Chukchi and 

Beaufort Seas are listed in table 4.7.

The length-weight regressions are dated, but at the time 

of their collection, allowed the estimation of fish condition 

and growth rates at scattered locations around the Arctic 

coast. Their historical value will be in comparison with 

contemporary estimates of these parameters and a potential 

benchmark to evaluate the effects of changing climate 

and environmental conditions (for example, Chen and 

Sakurai, 1993 for Saffron Cod). Given the status of existing 

information, a structured information management process, 

such as the Bayesian modeling approach used by Jay and 

others (2011) for Pacific walrus, could be used to evaluate 

needs and priorities for population dynamics modeling relative 

to offshore oil and gas development, climate change, and 

ecosystem-based management.

Current Understanding of Population Dynamics

There are typically four reasons to study population 

dynamics. First, population effects are often considered 

the most relevant response to environmental (ecosystem) 

perturbation. Second, population viability analyses (time to 

extinction) are used to manage endangered species. Third, 

they are critical determinants of sustainable yields in fishery 

management. Fourth, population processes are critical to 

understanding ecosystem dynamics and ecological processes 

(for example, regime shifts and trophic cascades). The first 

and fourth reasons are the most germane to the present Arctic 

management need and qualitative approaches to each are 

possible. Considerable research attention is being applied to 

developing quantitative understanding of population dynamics 

in light of changing Arctic conditions and potential fisheries.

Knowing how fish respond to variable environmental 

conditions and demographic changes in their populations 

is central to effective environmental assessments, potential 

fishery management, and informed Arctic policy. Knowledge 

about the demographics of marine fish population in the U.S. 

Arctic, while sparse, varies by season and distance from 

shore. More population data and information are available 

from nearshore areas during ice-free months simply because 

this is where the most research has been conducted. Offshore 

data collection has been of a reconnaissance nature and has 

allowed useful descriptions of the distribution and abundance 

of species for given areas and times. In the nearshore, the 

greatest amount of population information is available from 

inshore waters (<2 m) of the Alaska Beaufort Sea. This relates 

to the seasonal habitats for iconic species (Arctic Cisco and 

Least Cisco, Broad Whitefish, and Dolly Varden), logistical 

constraints of sampling, and coastal locations of industrial 

developments. Age-and-growth and size-at-age relationships, 

sex ratio, age- and size-at-maturity, fecundity, and other life 

history traits (for example, gonadosomal indices) have been 

described for Dolly Varden, Arctic and Least Ciscoes, Broad 

and Humpback Whitefish, Arctic Cod, Arctic Flounder, 

and Fourhorn Sculpin with most data collections occurring 

between the mid-1980s and early-2000s8

. Life history 

characteristics of Arctic Smelt sampled in the Colville River 

delta were summarized by Haldorson and Craig (1984).

The nearshore emphasis noted for the Beaufort Sea 

also extends to the amount of process-oriented research in 

Simpson Lagoon and Prudhoe Bay. For fishery components 

of the ecosystem, process studies have examined population 

movements and migrations with respect to prey availability, 

trophodynamic relationships, and coastal hydrodynamics. 8

Population information obtained through NOAA and MMS sponsored 

studies for the Alaska OCS Region; British Petroleum (Alaska), Inc. and the 

North Slope Borough fish monitoring in Prudhoe Bay; and U.S. Fish and 

Wildlife coastal resource inventories of the Arctic National Wildlife Refuge.
648 Alaska Arctic Marine Fish Ecology Catalog

Table 4.7. Age-length and length-weight relationships of fishes in the U.S. Arctic, with available data only from outside the U.S. 

Chukchi and Beaufort Seas.

Age-length relationships

Species Location Source

Acantholumpenus mackayi Tuktoyatuk Harbour, Yukon Territory, Canada Hopky and Ratynski (1983)

Ammodytes hexapterus Lower Cook Inlet, Alaska, USA Robards and others (2002)

Gadus chalcogrammus Eastern Bering Sea, Alaska, USA J. Ianelli, oral commun. (2010)

Gadus macrocephalus Bering Sea, Alaska, USA

Gulf of Alaska, Alaska, USA

Stark (2007)

Gymnelus viridis Northwest Territories, Canada Green and Mitchell (1997)

Hippoglossus stenolepis Northeast Pacific Forsberg (2011)

Limanda aspera Eastern Bering Sea, Alaska USA Wilderbuer and others (1992)

Lumpenus fabricii Tuktoyatuk Harbour, Yukon Territory, Canada Hopky and Ratynski (1983)

Pleuronectes quadrituberculatus Eastern Bering Sea, Alaska, USA Zhang and others (1998)

Squalus suckleyi Hecate Strait, British Columbia, Canada

Georgia Strait, British Columbia, Canada

Georgia Strait, British Columbia, Canada

Ketchen (1975)

Ketchen (1975)

Saunders and McFarlane (1993)

Triglops pingelii Kamchatka, Russia Tokranov (1995)

Reinhardtius pingeli Eastern Bering Sea, Alaska, USA Ianelli and others (2007)

Zaproa silenus Kuril Islands/Kamchatka, Russia Tokranov (1999)

Northeast Pacific, Alaska, USA Smith and others (2004)

Weight-length relationships

Species Location Source

Acantholumpenus mackayi Tuktoyatuk Harbor, Yukon Territory, Canada Hopky and Ratynski (1983)

Ammodytes hexapterus Lower Cook Inlet, Alaska, USA Robards and others (2002)

Gadus chalcogrammus Eastern Bering Sea, Alaska, USA J. Ianelli, oral commun. (2010)

Gadus macrocephalus Eastern Bering Sea, Alaska, USA Thompson and others (2007)

Gymnelus viridis Northwest Territories, Canada Green and Mitchell (1997)

Hippoglossus stenolepis Northeast Pacific, Alaska, USA Sadorus and others (2012)

Limanda aspera Eastern Bering Sea, Alaska, USA Wilderbuer and others (1992)

Lumpenus fabricii Tuktoyatuk Harbor, Yukon Territory, Canada Hopky and Ratynski (1983)

Pleuronectes quadrituberculatus Eastern Bering Sea, Alaska, USA Zhang and others (1998)

Reinhardtius pingeli Eastern Bering Sea, Alaska, USA Ianelli and others (2007)

Squalus suckleyi Washington, USA Wildermuth (1983)

Triglops pingelii Kamchatka, Russia Tokranov (1995)

Zaproa silenus Northeast Pacific, Alaska, USA Smith and others (2004)
Chapter 4 649

Field and laboratory experiments have focused on 

physiological responses to natural variability in the physical 

conditions of habitats in bays, lagoons, and exposed coastlines 

as well as changes in conditions resulting from coastal 

construction. Years of research and monitoring in the vicinity 

of Prudhoe Bay has yielded a wealth of information about the 

life histories, habitats, and harvests of the coastal fishes and 

increasingly in the area around Point Barrow. Information 

about genetic relationships (diversity and structure) is more 

limited with respect to population dynamics. Genetic studies 

have been conducted on Arctic Cisco, Dolly Varden, Arctic 

Cod, and there is growing interest in Saffron Cod. The 

importance of these species in subsistence fisheries, possible 

effects of offshore oil and gas development, and possible 

effects of changing climate conditions, have, or currently are, 

responsible for these studies.

Salmon, charr, and whitefishes are regionally significant 

as traditional use species in subsistence fisheries and are of 

increasing value as recreational resources in Arctic watersheds. 

However, among these species, especially for those stocks 

occurring in North Slope drainages, harvest rates have not 

historically warranted management concerns about fishery 

effects on population sustainability. As an example, Arvey 

(1991) estimated sport fishery harvests to be 1,000–3,000 fish 

annually, and subsistence catches to be two to six times this 

amount. Freshwater habitats, especially overwintering sites, 

are thought to be most limiting to this species, and thus the 

possible effects of climate change have raised concerns about 

their management and conservation (Viavant, 2001; Crane and 

others, 2005; Greiner, 2009). These concerns have translated 

into efforts to identify and map overwintering locations and 

feasibility studies for enumeration of Dolly Varden at major 

overwintering sites including evaluation of the genetic stock 

composition at key index areas (for example, Hulahula River, 

Saganavirktok River, and tributaries). Population monitoring 

methods include aerial counts of overwintering abundance 

with possible validation by mark-recapture studies (that 

is, Ivishak, Anaktuvuk, Canning, and Hulahula Rivers, see 

Viavant, 2009; Rat River, see Harwood and others, 2009) 

and use of Dual Frequency Identification Sonar (DIDSON) 

technology and various species validation techniques (for 

example, underwater camera and hook and line sampling) in 

the Hulahula River (Greiner, 2009). 

Information about the demographic traits for most marine 

fish populations in the Chukchi Sea is lacking largely due to 

the absence of large-scale fisheries operating in the region and 

corresponding lack of resource assessments and demographic 

data. Some population information is available from surveys 

sponsored by BOEM during the early 1990s. As an example, 

demographic information was developed for Bering Flounder 

and Arctic Staghorn Sculpin in the northwestern Chukchi 

Sea by Barber and others (1994). New research being 

conducted and planned by NOAA and the University of 

Alaska is expected to address many of the gaps in life history 

and demographic traits for marine fishes in the Chukchi and 

Beaufort Sea shelf habitats.

Modeling Environmental Effects on 

Population Dynamics

The quantitative approaches to population ecology 

studies of Arctic fish, although limited, are most appropriately 

considered in light of traditional research associated with 

physiological responses, dispersal processes, and food web 

relations.

Physiological research in the nearshore Alaska Beaufort 

Sea has focused on seasonal growth rates (VBGM growth 

parameters), body condition (Anderson and Gutreuter, 1983), 

as part of long-term monitoring of coastal habitats (<2 m) 

most affected by oil and gas development9. The monitoring 

was designed to obtain biological information about key 

indicators (notably Arctic Cisco and Broad Whitefish), their 

use of habitats (generally in the area between the Colville 

and Saganavirktok Rivers), and population responses to 

variable hydrographic conditions. Shallow coastal waters are 

responsive to wind-driven circulation, regional upwelling, 

and, in the Prudhoe Bay area, localized changes in temperature 

and salinity associated causeway construction (for example, 

breaching and wake eddy effects at West Dock and Endicott 

projects). The long-term monitoring provided seasonal indices 

of fish abundance and samples for age and growth analysis 

and trophic studies for indicator species across a large segment 

of the coast. Physical data collections included observations 

about wind and weather conditions and hydrographic profiles 

associated with fish collection efforts. Fish catch records 

from the Colville River delta are another source of long-term 

population data. Genetic samples (mtDNA) were obtained 

for Broad Whitefish from freshwater habitats in the Colville 

and Saganavirktok Rivers to determine if they were distinct 

populations (Patton and others, 1997).

The monitoring in Prudhoe Bay was designed to 

evaluate potential causeway effects on coastal habitats 

and amphidromous fish populations and related effects to 

subsistence and commercial fisheries. In a 5-year study, 

Griffiths and others (1992) studied growth patterns in young 

(age-1 and age-2) Arctic Ciscoes and Broad Whitefish 

collected around the causeways (Fechhelm and others, 1995a; 

1995b). Growth (increase in length) in each species was 

positively correlated with temperature and body conditions. 

Negative correlations between condition and salinity were 

reported for larger Broad Whitefish and Griffiths and others 

(1992) suggested that the availability of prey and food benefits 

associated with marine waters moved inshore, outweighed 

9

A listing of scientific publications associated with long-term 

monitoring studies in the Prudhoe Bay area, Alaska, is available from LGL 

Limited (2014).
650 Alaska Arctic Marine Fish Ecology Catalog

negative effects on their growth due to temporary exposures 

to colder-higher salinity conditions. Reduced growth in about 

4–6 percent of each was attributed to causeway effects (for 

example, thermal barriers and wake eddies).

Numerous ichthyoplankton surveys have been done in 

the Chukchi and Beaufort Seas. The studies were designed as 

reconnaissance surveys rather than dispersal studies although 

the results, as reviewed in this report, shed considerable light 

on diversity patterns and processes. Research directly focused 

on dispersal processes was used to study population trends and 

origins of Arctic Cisco in northern Alaska. A delay-difference 

population model (Deriso, 1980) was used by Gallaway 

and others (1983) to estimate biomass production of Arctic 

Ciscoes targeted in a local fishery in the Colville River10. 

Harvest records for the autumn gillnet fishery have been 

available since they were first collected in 1967 (Fechhelm 

and others, 2007; Moulton and others, 2010). The Deriso 

model, classified as a partially age-structured model, was 

used to investigate annual abundance patterns in the fishery 

relying on data obtained from (1) 15 years of catch records, 

(2) CPUE obtained in coastal surveys, and (3) population 

information acquired from tagging studies associated with 

those surveys. The model was parameterized with biological 

information about estimated growth, recruitment, survival and 

age structure, of fish captured in the Colville River fishery. In 

this case, the modeling allowed for time lags in the dynamics 

of the stock for Arctic Cisco (stocks are known for their slow 

growth and late age of entry to the fishery) and accounted 

for observed fluctuations in annual abundance as reflected 

in the catch. Importantly, the model generated a hypothesis 

about Mackenzie River origins for this Alaskan stock and 

annual transport of young-of-the-year ciscoes (Gallaway and 

others, 1983). A dispersal mechanism in wind-aided currents 

was described by Fechhelm and Fissel (1988) and its effects 

on recruitment processes in the Colville River fishery by 

Fechhelm and Griffiths (1990). The interaction of fish, winds, 

and currents on young-of-the-year (YOY) migration patterns 

was numerically modeled by Colonell and Gallaway (1997) 

and results considered in light of interannual fluctuations in 

abundance of fish in coastal monitoring in Prudhoe Bay and 

catch rates in the fishery11. A simulation model was developed 

to investigate wind effects on interannual variations in summer 

dispersal patterns of juvenile Least Cisco in the Prudhoe Bay 

area (Fechhelm and others, 1995a). The study of age structure 

in Colville River harvested Arctic Cisco (Zimmerman and 

others, 2013) may result in the improved population modeling 

recommended by Murphy and others (2007).

Predator-prey studies, mostly in the form of stomach 

content analyses and more recently using stable isotope 

and fatty acids (for example, Pirtle and Mueter 2011) are 

important components of habitat and community analyses 

including understanding ecological relations with respect 

to the maintenance of important populations or evaluations 

of potential cascading effects of regime shifts (for example, 

Suryan and others 2009; Carey and others, 2013). Many 

investigations in the Chukchi and Beaufort Seas have 

indicated the critical role that Arctic Cod have in Arctic 

marine food webs (for example, Bradstreet and others, 1986; 

Springer and others, 1986; Pirtle and Mueter, 2011) and their 

intermediate position in top-down and bottom-up energy 

pathways (Hunt and McKinnell. 2006). The results of marine 

fishery surveys invariably list this species, regardless of depth, 

season, or year, as the most abundant; usually far exceeding 

all other species reported in survey catches (Pirtle and Mueter, 

201112). Given this abundance and its central role in energy 

pathways, Arctic Cod must be viewed as a keystone species. 

This does not detract from the ecological significance of lipidrich Capelin (Carscadden and Vilhjálmsson, 2002; Rose, 2005) 

and Arctic Sand Lance (for example, Bluhm and Gradinger, 

2008) and, to a lesser extent Pacific Herring (for example, 

Mueter and others, University of Alaska-Fairbanks, oral 

commun., 2013) and Arctic Smelt (Craig and others, 1984a). 

These populations are subject to greater interannual changes 

in abundance, but have been shown to be locally abundant 

at various coastal locations and to be important in regional 

food webs. 

The role of invertebrate prey in carbon cycling in Arctic 

marine ecosystems (Dunton and others, 2006) and in the 

diets of marine fishes and many higher-level consumers is 

significant (Craig and others, 1984b). In the Beaufort Sea, 

especially, the annual colonization of coastal waters by 

epibenthic amphipods and mysids, pelagic copepods, and 

other species (Craig and others, 1984b) forms the basis of 

the nearshore food web. Craig (1989a) and Craig and others, 

1984b) discussed the availability of food in coastal habitats 

to standing crops of macroinvertebrate prey in freshwater 

habitats as being a significant factor in the evolution of 

anadromy (amphidromy) in the Arctic. The role of the 

epibenthic invertebrate food base is a key premise of timetested conceptual models of fish habitat use in the Beaufort 

10

Arctic Cisco are harvested between October and November each year. The 

availability of fish varies annually and Gallaway and others (1983) reported 

total catches ranging between 9,268 (1979) and 71,575 (1973) fish per year. 

The size of individuals captured in the fishery ranged from 240 to 380 mm 

(FL). From 1985 to 2004 about 38,600 Arctic Cisco have been harvested 

annually in commercial and subsistence fisheries (Least Ciscoes averaged 

about 18,600 annually during the same period). The average annual removal 

rate for Arctic Cisco was 8.9 percent of the available fish; the average annual 

removal rate for Least Cisco was 6.8 percent

11

Zimmerman and others (2013) re-examined the Mackenzie River 

hypothesis in light of new genetics data, otolith microchemistry, and potential 

climate change effects on the coastal migration of YOY fish. Their analysis 

did not refute Mackenzie River origins for Arctic Cisco found in the Colville 

River.

12

Pirtle and Mueter (2011) report peak densities of adult Arctic Cod at 

150,000 fish per ha-1 at bottom depths of 100–350 m, and peak densities of 

YOY Arctic Cod at 160,000 fish per ha-1 at bottom depths of 20–75 m in the 

U.S. Beaufort Sea.
Chapter 4 651

Sea (Craig and others, 1982; Craig, 1984) and bioenergetics 

of coastal fishes in Prudhoe Bay (Fechhelm and others, 

1995a; 1995b). Marine food webs for fishery resources in the 

offshore Beaufort Sea are less studied, but likely are similar to 

shelf and slope areas of the Chukchi Sea. Planktonic species 

and under-ice fauna (phytoplankton, amphipods, copepods, 

euphaussiids, and larval fish) are important food resources 

for marine fishes in pelagic systems (for example, Gadidae 

Salmonidae, and others). Invertebrate species associated with 

the seafloor (for example, shrimp, amphipods, polychaetes, 

and benthic fauna) for marine species occupying benthic 

systems (for example, Pleuronectidae, Liparidae, Sticheaidae, 

and others). 

In both nearshore and offshore environments of the U.S. 

Beaufort and Chukchi Seas, a broad generalization can be 

made that fish are also important in the diets of many fish 

species (for example, Gadidae, Salmonidae, Pleuronectidae, 

and others), particularly older, larger-sized individuals. Like 

other gadids, the Arctic Cod is cannibalistic. Thorsteinson 

and others (1990) hypothesized that the observed differences 

in coastal habitat used by YOY Arctic Cod and older cod 

in Camden Bay, Alaska, were an adaptive response to 

cannibalism. Similarly, some species, such as the Dolly 

Varden, are much more piscivorous than other dominant 

members of the coastal assemblage, but like most other 

species are opportunistic in their selection of foods. Although 

the food habits of some marine fishes are better known 

than others, the application of bioenergetics approaches to 

population dynamics modeling suffers from a (1) quantitative 

data about predator-prey relationships; (2) lack of seasonal 

information (spring and winter months especially); and 

(3) lack of long-term population data for most species of 

interest. For a small number of species in which appropriate 

data are available (for example, Broad Whitefish and Arctic 

and Least Ciscoes), bioenergetics applications to population 

ecology studies in coupled models may be possible. In the 

absence of data, other approaches to reduce uncertainties 

associated with selecting population models to evaluate 

responses to environmental influences (Katsanevakis, 2006), 

such as changes in recruitment and growth, have been done 

for Arctic Cisco (for example, von Bielea and others, 2011). 

In the latter instance, YOY recruitment into the Colville River 

was used examine the long-term response (1978–2004) of the 

population to climate change using multi-model inferences 

to study relations between trends in abundance and YOY 

growth and various environmental parameters (AO Index, 

air temperature, east wind speed, sea-ice concentration, and 

MacKenzie River discharge) and their influences of primary 

production and life stages of important cisco foods.

Biological Interactions 

Estimation of ecosystem functions (for example, 

primary, secondary, and tertiary productivity; nutrient fluxes; 

carbon fixation; and organic matter mineralization and 

suspension removal) are feasible measures of biodiversity 

importance (Weslawski and others, 2006). The ecological 

effects of marine production cycles on the distribution and 

abundance of the Arctic marine fishes has received relatively 

little attention in the United States Arctic. In a large-scale 

analysis, Smetacek and Nicol (2005) compared the biological 

productivity reported in the Barents (Russia) and Chukchi 

Seas to the Siberian and Canadian shelf seas. The much 

higher productivity in the Barents and Chukchi Seas—

gateways to the Arctic—were related to the advection and 

retention of nutrients in these systems. With respect to the 

Chukchi Sea, currents flowing through Bering Strait are rich 

in nutrients which, when mixed with iron (Fe) in shelf water 

and melting ice, fuels the high primary productivity of the 

region. Because of the Chukchi Sea’s shallow depth and lack 

of a well-developed pelagic community, much of the bloom 

biomass (organic matter) settles out on the sediments and 

supports a rich, especially invertebrate, benthic fauna (Jay and 

others, 2011). The carbon sink and intense nutrient recycling 

occurring near the seafloor are critical processes supporting 

the region’s benthic communities. In contrast, the Barents 

Sea is deeper and a greater proportion of phytoplankton 

production is retained in the water column. “The exceptionally 

high copepod biomass of the western Barents Sea, largely 

attributable to advection from the adjoining Norwegian Sea, 

supports, or has supported, huge stocks of planktivorous fish 

including capelin and herring and their predators, particularly 

Atlantic cod” (from Smetacek and Nicol, 2005, p. 364).

Trophic Linkages

Knowledge about ecosystem functions is foundational to 

understanding biological interactions of marine fishes. Fishes 

and zooplankton not only are important prey, but also are 

critical conduits of energy in regional food webs (for example, 

Chukchi Sea—Piatt and others, 1989). The large-scale animal 

distribution patterns (fishes, birds, and mammals) are driven 

by food availability, not the absence of predators (Hunt and 

McKinnell, 2006). These authors noted that at regional scales, 

current predation or past predation events likely have shaped 

local distributions, at least in marine birds and pinnipeds. 

Wasp-waist control occurs when one of the intermediate 

trophic levels is dominated by a single species, which may be 

the case with Arctic Cod in the Chukchi and Beaufort Seas. 
652 Alaska Arctic Marine Fish Ecology Catalog

Considerable research underscores the critical function of 

Arctic Cod in Arctic marine ecosystems because no alternate 

food source of equivalent trophic value exists. Processes in 

these ecosystems may have features that result in a switch 

from bottom-up to top-down control (Hunt and McKinnell, 

2006) given the variable abundance of small pelagic 

species in response to changing environmental conditions. 

A moderating influence of sea ice in the population dynamics 

of Arctic Cod is hypothesized; however, loss of sea ice due 

to climate warming and unknown associated effects on 

the ecology of this species is of environmental concern13. 

Similarly, the potential effects of large oil spills and related 

clean-up activities on Arctic Cod could affect marine 

ecosystem functions.

The Arctic Cod is a key prey of many marine mammals 

and seabirds as evidenced by their occurrence in the diets 

of belugas and ringed and bearded seals, Pacific walruses 

(occasionally), Thick-billed (Uria lomvia) and Common 

Murres (U. aalge), Black Guillemots (Cepphusgrylle), 

Black-legged Kittiwakes (Rissa tridactyla), Northern Fulmars 

(Fulmarus glacialis), Arctic Terns (Sterna paradisaea 

Pontoppidan), and Glaucous (Larus hyperboreus), Sabine’s 

(Xema sabini), Ivory (Pagophila eburnea), and Ross’s Gulls 

(Rhodostethia rosea) (Hunt and McKinnell, 2006). Arctic 

Cod also are of indirect importance to Polar bears (Ursus 

maritimus) and Arctic foxes (Vulpes lagopus), because their 

primary marine food, the ringed seal, also relies on them 

as food. 

Other fishes are important in regional food webs. Pirtle 

and Mueter (2011) summarized the known trophic linkages 

for seven species of marine fish (Arctic Cod, Capelin, Arctic 

Sand Lance, Pacific Herring, and Saffron Cod, and Arctic 

and Least Ciscoes) in the U.S. Beaufort Sea. Some aspects 

about trophic linkages between marine fish and invertebrate 

communities are described in detail in the species accounts 

(chapter 3). Depending on season and location, many of 

the marine species are major food resources for regional 

populations of seabirds, marine mammals, other fishes, 

and humans. As an example, the largest seabird colony in 

the North American Arctic is located at Capes Lisburne 

and Thompson in northwestern Alaska. Arctic Sand Lance, 

Capelin, and Arctic Cod are important prey for kittiwakes and 

murres at the colony during the breeding season (including 

pre-breeding and post-fledging periods). Pacific Herring and 

Arctic Cod are important foods of other fish (Inconnu, Chum 

and Pink Salmon, Dolly Varden), birds, and marine mammals 

in the Bering Strait and southeastern Chukchi Sea. Least 

Cisco is an important summer food of the Red-throated Loon 

in coastal habitats along the Chukchi Sea. Arctic Cod and 

Saffron Cod are notably important in the diets of most marine 

animals common to the Chukchi and Beaufort Seas including 

many seabirds, cetaceans, seals, polar bears, and Arctic foxes. 

The amphidromous fishes (whitefishes and charr) in the 

nearshore Beaufort Sea are preyed upon by Beluga whales 

(Delphnapterus leucas) and Ringed Seals (Pusa hispida). 

Marine Fish Disease Ecology

In spite of obvious effects to humans and captive 

animals, fisheries managers often ignore disease as a 

significant factor affecting the abundance of wild populations 

because the effects are difficult to observe and quantify. 

Historically, most fish-health research has been directed 

toward identification, treatment, and prevention of diseases 

of hatchery fishes; however, recent studies (for example, 

Hershberger and others, 2013) from marine, freshwater, and 

terrestrial environments indicate that infectious and parasitic 

diseases can be responsible for population oscillations, 

extinction of endangered species, reduced host fitness, and 

increased susceptibility to predation as well as an important 

component of natural mortality. The recognition of disease as 

a population-limiting factor for wild fish is partly the result 

of the emergence of high profile pathogens and changes in 

environmental conditions that shift the host-pathogen balance 

in favor of disease. Among such environmental changes, 

global warming associated with climate change is seen to be 

a particularly important threat for poikilothermic vertebrates, 

such as fish, for which environmental temperature is a 

controlling factor in their physiology and immune response. 

The emergence of Ichthyophonus infections14 in adult Yukon 

River Chinook Salmon, Pacific Halibut in the northeast 

Pacific, and Atlantic herring (Clupea harengus) off Iceland, 

are examples of disease conditions with links to global 

warming and possible transmission through trophic linkages 

(Woodson and others, 2011; Dykstra and others, 2012; Jim 

Winton, U.S. Geological Survey, oral commun., 2013). The 

disease is associated with adverse flesh quality and possible 

pre-spawning losses.

13

Arctic Cod are the most abundant forage fish and play a central role in 

the transfer of energy from plankton to higher-level consumers like ringed 

seals and polar bears (Hunt and McKinnell, 2006). As warming alters sea 

ice conditions, northward shifts in the distribution of marine fishes are 

expected. Such food-web impacts would propagate through the ecosystem, 

from sea-floor organisms to their predators, and, ultimately, to the subsistence 

users whose livelihoods largely depend on having reliable access to marine 

mammals, fish, and other wildlife. Shifts in marine biodiversity will partly 

depend on whether species are associated with the open ocean or with 

seasonal sea ice (Hunt and others, 2013).

14

Phenotypic (Hershberger and others, 2008) and genotypic (Criscione and 

others, 2002; Rasmussen and others, 2010) differences have been identified 

among isolations of Ichthyophonus hoferi from the northeastern Pacific, 

suggesting that there are multiple sympatric species in the region. Due to this 

taxonomic uncertainty, the parasite is referred herein by its generic name.
Chapter 4 653

Human Interactions

Subsistence Fisheries

Generally, subsistence is considered hunting, fishing, 

and gathering for the primary purpose of acquiring, sharing, 

or bartering traditional foods. Craig (1987; 1989b), Braund 

and Kruse (2009), and DeGange and Thorsteinson (2011) 

summarized the relative importance of marine fishes in 

the traditional economies of 11 Arctic communities using 

data obtained from the Alaska Department of Fish and 

Game15 and others. Depending on village location, Pacific 

salmon, Dolly Varden, Arctic and Least Ciscoes, Broad 

and Humpback whitefishes, and Inconnu were the major 

fish species harvested. Arctic Cod, Saffron Cod, flounders 

(for example, Arctic Flounder), and sculpins (for example, 

Fourhorn Sculpin) and other marine fishes, such as Capelin 

and Pacific Herring, that occur in inshore waters, are of more 

limited, but still important usage. The subsistence survey data 

summarized by DeGange and Thorsteinson (2011) that Alaska 

Native Arctic residents are actively engaged in subsistence 

fishing and that fish are important foods (5–10 percent or 

more of traditional diet); fish is consumed on a daily basis in 

many villages, and in some cases provides more of a dietary 

contribution than any other food source. In the Kotzebue 

Sound area, fully one-third to one-half of the total subsistence 

harvest by weight consists of fishes. Although the inhabitants 

of the North Slope often depend much less on fishing and 

more on marine mammal and caribou (Rangifer tarandus) 

hunting, significant fish harvests are still made. The fact that 

fishes comprise more than 10 percent of the total subsistence 

harvest of Point Barrow is remarkable, considering the 

number and biomass of bowhead whales (Balaena mysticetus) 

harvested there annually (DeGange and Thorsteinson, 

2011). At a larger scale, communities north of Bering Strait 

harvested 23 bowhead whales (about 750 tons) in 2008; the 

subsistence catch of marine fishes for the same area in Alaska 

was estimated at 60–70 percent of this amount. At Nuiqsut 

(Colville River), fish are the largest single contributors to the 

subsistence economy at nearly 40 percent of the total harvest.

According to an Alaskan catch reconstruction study 

(Booth and others, 2008), the total yearly subsistence harvest 

of fishes from Wales to Kaktovik in 1950 was approximately 

the same as in 2006 (450–500 tons). Changes in subsistence 

use at Wainwright, Barrow, Nuiqsut, and Kaktovik pre- and 

post-2000 were reported by Braund and Kruse (2009) and, 

within the constraints of existing data, show consistency over 

time for most species, and increasing catches of Pacific salmon 

(Carothers and others, 2013). Fish landings in the Chukchi 

Sea are primarily for local consumption (North Pacific Fishery 

Management Council, 2009). The most important species are 

salmonids, especially Chum Salmon and Inconnu, although 

15 species of marine fishes are used. In Kotzebue Sound, 

Dolly Varden, Pacific Herring, and several species of flounder 

also are regularly harvested. About 90,000 fish are harvested 

annually and, in recent years, about 25,000 salmon were 

captured (Magdanz and others, 2011). Small numbers of five 

Pacific salmon found in Arctic Alaska are occasionally caught 

as far to the east as Amundsen Gulf, Canada. Generally, the 

closer a community’s proximity to the more temperate and 

productive Bering Sea, the greater the number of salmon 

species caught and the greater the contribution of salmon 

to the local population’s diet. This corresponds directly to 

the distribution of spawning populations of various salmon 

species. In the U.S. Arctic, Chum and Pink Salmon have the 

widest range and subsistence values.

The whitefish species are significant subsistence fishery 

resources in the 11 coastal villages where data are available 

(DeGange and Thorsteinson, 2011). Georgette and Sheidt 

(2005) documented their importance in Kotzebue Sound 

including the seasonality and methods of catch and complexity 

of Iñupiaq taxonomy. Farther north and east of Kotzebue 

Sound, various whitefish and cisco species and Dolly Varden 

comprise the marine basis of regional subsistence fisheries. 

Inconnu are important in the area of the central Kotzebue 

Sound and in the Mackenzie River Delta, Canada, but 

nowhere between. Least Cisco are important in local fisheries 

east of the Saganavirktok River, Alaska, but are little used 

by villagers at Kaktovik. Capelin, Arctic Smelt, Arctic Cod, 

other gadids, and some sculpins are eaten at Point Barrow. A 

small commercial and subsistence fishery for Least and Arctic 

ciscoes occurred until recently in the Colville River Delta, 

near Prudhoe Bay. In 2010, the Helmerick family terminated 

the commercial fishery for Arctic Cisco, but subsistence 

fishing continues by villagers from Nuiqsut. The Arctic 

Ciscoes captured in this fishery originate in the MacKenzie 

River, Canada; the Colville River, by virtue of its size, is the 

major overwintering site for Arctic Cisco in Arctic Alaska. 

The subsistence gill net fishery is size selective for 5- and 

6-year-old fish, which are used by residents of Nuiqsut and 

Barrow, Alaska. Long-term fishery monitoring in Prudhoe Bay 

indicates increasing abundance of some marine fishes, such 

as Arctic Smelt and Arctic Flounder (Fechhelm and others, 

2006), and suggests the possibility of their increased use by 

coastal residents in Nuiqsut and Barrow.

Dolly Varden is an important subsistence resource 

throughout the coastal Chukchi-Beaufort Sea region from 

the Seward Peninsula east to Kaktovik. Reliance on this 

species by humans is greatest in the villages of Wainwright 

and Kaktovik and in Kotzebue Sound. Located in the eastern 

Alaska Beaufort Sea, Kaktovik is situated near Dolly Varden 

spawning grounds in the Brooks Range (Craig, 1989a). 15

See http://www.subsistence.adfg.state.ak.us/CSIS/.
654 Alaska Arctic Marine Fish Ecology Catalog

The timing and location of subsistence fisheries are 

intimately linked to the life cycle and habitats of targeted 

fishes. Environmental conditions, such as temperatures 

for drying and freezing of the fish flesh also are important. 

Marine and estuarine species are vulnerable to coastal fishing 

and the amphidromous/anadromous species are captured in 

both freshwater and coastal sites. Overwintering species, 

such as Arctic and Least ciscoes are targeted in autumn and 

winter fisheries with gillnets fished under ice. Coastally, 

during summer months, many species are captured in hookand-line fisheries. Other species, such as Dolly Varden and 

Pacific Salmon can be captured in these fisheries. The marine 

connections of some lagoons, such as those in Kotzebue 

Sound, often are blocked by storm-induced gravel barriers 

near the end of summer, forming natural traps, or ditching, for 

desired species, such as Least Cisco (Georgette and Shiedt, 

2005). Under certain conditions, gill nets are seined and fish 

weirs (traps) are used in Kotzebue Sound. 

Commercial Fisheries

Commercial fishing in the U.S. Arctic is currently 

prohibited by Federal and international regulations (for 

example, National Pollution Funds Center (NPFC) and 

International Halibut Commission) because most fish 

stocks in the Chukchi and Beaufort Seas cannot support 

commercial harvesting (Wilson and Ormseth 2009). Three 

stocks (snow crab [Chionoecetes opilio], Arctic Cod; and 

Saffron Cod) were identified as sufficiently abundant to 

support potential fisheries. However, additional research 

(life history, interactions, and population dynamics) and 

resource assessments (monitor abundance and productivity) 

are needed to design sustainable harvest strategies (Levin 

and others, 2009; Fluharty, 2012). The current knowledge 

and understanding of fishery managers is that Arctic Cod 

and Saffron Cod are too important in regional food webs 

(for example, marine mammal and seabird predators) to be 

the targets of new, offshore commercial fisheries. Ongoing 

research by BOEM, NOAA, and NPRB involves ecological 

baseline surveys that may provide the basis for age- or 

length-based stock assessments for management of marine 

fish and shellfish resources in the Arctic. Oceanographic 

measurements and data on abundance, stock structure, growth, 

food habits, and energetics data, are being collected to develop 

quantitative information about the populations and responses 

to environmental changes including those resulting from 

human interventions.

Policy Interface—Ecological and Biologically 

Sensitive Areas

The Convention of Biological Diversity considered the 

ecological significance of Arctic marine fishes in defining 

Ecological and Biologically Sensitive Areas (EBSAs; Speer 

and Laughlin, 2011). Seasonally important habitats for many 

fishes, birds and mammals were identified along the ice edge, 

in polynyas, in coastal lagoons and on barrier islands, and 

in bays and river deltas. In the U.S. Pacific Region (Arctic 

Monitoring and Assessment Programme [AMAP]), the 

EBSAs include St. Lawrence Island, Bering Strait, Chukchi 

Beaufort Coast, Wrangell Island and polar pack ice. The 

global significance of these EBSAs is based on ecosystem 

functioning and with respect to marine fishes in the U.S. 

Chukchi and Beaufort Seas includes the Bering Strait, 

Chukchi Beaufort Coast, and Polar ice pack.

Bering Strait

The Bering Strait16 exhibits the highest levels of fish and 

wildlife productivity and diversity in the Arctic and is the only 

connection between the Pacific and Arctic Oceans. Arctic Cod

and other species of forage fishes are abundant and important 

to many marine predators, and the region supports populations 

of whitefishes and charr, which are important seasonally for 

native community subsistence. 

Chukchi Beaufort Coast

The Chukchi Sea has massive phytoplankton blooms, 

which along with annual sea ice algae production, cannot be 

fully exploited by the zooplankton communities resulting 

in an impressively high biomass of benthic infauna and 

epifauna. Capelin occurs along the sandy seaward beaches 

of barrier islands in the area of Points Lay and Barrow. 

Coastal waters provide whitefish nursery areas and migration 

corridors for whitefish, ciscoes and Dolly Varden. The spring 

lead and Hanna Shoal (Chukchi Sea) retain sea ice, making 

it an important area for seabirds in spring and pinnipeds 

in late summer when sea ice is absent over the rest of the 

continental shelf. 

16 The importance of the Bering Strait, as a gateway to the Arctic is widely 

recognized in national science policy (Speer and Laughlin, 2011; Wilson 

and Ormseth, 2011; Clement and others, 2013; National Ocean Policy 

Implementation Plan, 2013; U.S. Arctic Research Commission, 2013) and 

marine research associated with the Northern Bering Research Area (NPFMC) 

and Distributed Biological Observatories (National Science Foundation 

[NSF] and Alaska Ocean Observing System [AOOS]) and other large-scale 

ecosystem initiatives (that is, NRPB, BOEM, and NSF). 
Chapter 4 655

Polar Pack Ice

The multiyear pack ice provides habitat for distinctive 

fauna and flora. The extent of the multi-year ice is extremely 

variable inter-annually and is not a static geographic area, but 

rather an ever-changing feature that provides critical habitat 

for many animals. Arctic Cod is a key species in short food 

chains leading to seal and polar bear consumers.

Arctic Climate Change

Non-linear responses of marine ecosystems and 

populations to atmospheric forcing and warming are major 

sources of scientific uncertainty as they pertain to estimating 

probable outcomes of climate scenarios and describing the 

vulnerability of key biotic resources. The major sources 

of variation can be characterized as short-term effects on 

physical (for example, wind, waves, currents, temperature, 

stratification, nutrients, precipitation, and freshwater input) 

and biological (for example, phenology, primary production, 

food availability, and recruitment) environments, and longterm ecological changes (for example, temperature regimes, 

distributional shifts, and trophic cascades). Because warming 

effects are occurring more quickly toward the poles, both 

short- and long-term effects of these changes will be amplified 

in the Arctic. At the Arctic basin scale, marine ecosystems 

are affected by the interactions of physical and biological 

processes occurring at multiple scales of spatial and temporal 

resolution (for example, global—warming, regional—regime 

change, local—predation and survival). At a geographic scale, 

atmospheric forcing—related to the geography, persistence, 

and teleconnections of continental pressure systems—affects 

ecosystem function and structure at different spatial and 

temporal scales (Grebmeier and others, 2006a; 2006b). At 

smaller scales (centimeters to meters), recruitment processes 

can be affected by the dispersal and survival of larval fishes.

Ocean Connections

In association with atmospheric influences, Arctic marine 

ecosystems are also connected through ocean circulation and 

the long distance transport and mixing of north Atlantic and 

northeast Pacific waters. Marine waters over the Chukchi 

Sea shelf tend to be colder and more nutrient rich than water 

in the Beaufort Sea because of upwelling processes in the 

northern Bering Sea and transport of marine nutrients of deep 

northeast Pacific origins. Hydrographic conditions on the shelf 

indicate the presence of three main water masses that move 

to the north along the Alaska coast, across the Bering Sea 

from the northeast Pacific, and to the south along the Russia 

coast (Crawford and others, 2012). Farther north, the offshore 

Beaufort Sea is influenced by a mixing of Chukchi Sea water 

masses and north Atlantic-derived waters (transported to the 

east) and circulation of other Arctic waters in the Beaufort 

Gyre (transported to the west). An ephemeral band of brackish 

water forms along the coast of the North Slope during summer 

months. As summer advances, the frequency of west winds 

along the North Slope increases, resulting in marine intrusions 

and colder, higher salinity water nearshore.

Marine Ecosystem Effects

Extensive and rapid losses of sea ice in the Arctic are 

already affecting marine ecosystems and raising conservation 

concerns for ice-dependent fishes, birds, and mammals 

(Marz, 2010; Jay and others, 2011; Wiese and others, 2015). 

Sea ice melt and break-up during spring strongly drive 

phytoplankton production by enhancing light and stratification 

and stabilization of the water column. The intensity and 

magnitude of the events is therefore determined in large part 

due to temperature and salinity, light conditions, and nutrient 

availability as well as composition and fate of the plankton 

community. Generally, the annual production event, including 

algal growth under sea ice, forms the base of the marine food 

web, which in turn supports higher trophic level consumers 

including zooplankton, fishes, birds, and mammals. The 

consumption of phytoplankton by the herbivorous zooplankton 

in the Chukchi Sea tends to be inefficient (incomplete grazing) 

leading to a tight coupling between pelagic and benthic 

ecosystems and deposition of significant amounts of organic 

debris (decaying phytoplankton) to the benthos. In areas of 

high deposition, sea floor habitats have a diverse and abundant 

benthos that provides important feeding grounds for a rich 

invertebrate community and benthic feeding animals such 

as walrus and bearded seals. Biological hotspots have been 

identified in the Barrow Canyon and Hannah Shoal in the 

northeastern Chukchi Sea near Barrow, Alaska. 

Climate-induced changes in growth conditions for 

primary producers could affect the timing, productivity, and 

spatial extent of biological hot spots—areas of elevated food 

web activity (Leong and others, 2005). As an example, Frey 

and others (2011) reported a 20 percent increase in production 

throughout the Arctic Ocean during 1998–2009; an estimate 

that included a 48 percent increase in primary production 

for the Chukchi Sea. The increases were thought to reflect 

increased bloom durations in open water in time and space. 

Zooplankton communities may be particularly sensitive to 

such changes as their seasonal life cycles are in synchrony 

with the timing of ice-breakup and phytoplankton blooms 

(Smith and Schnack-Schiel, 1990; Deibel and Daly, 2007). 

With warming and changing sea ice conditions, northerly 

shifts in the distribution of marine fish have been projected. 

Although commercial concentrations of marine fish are 

not anticipated in the near term (probably decades due to 

temperature constraints), conditions that are more favorable 

for energy use by pelagic species (gadids, salmonids, and other 

marine fishes) in the Chukchi Sea may result from ecological 
656 Alaska Arctic Marine Fish Ecology Catalog

changes associated with regime shift (Grebmeier and others, 

2006b). In this instance, changes in the food web, such as 

reduced standing stocks of benthic invertebrate foods, could 

potentially affect the distribution and abundance of predators. 

Such changes in distributional patterns already are being 

observed in Pacific walrus (David Douglas, U.S. Geological 

Survey, Juneau, Alaska, oral commun., 2013). 

In a study of possible responses of the Atlantic Cod to 

climate change, Pörtner and others (2008) examined functional 

interactions of ecological and physiological processes 

and effects on this species at various levels of biological 

organization. The potential effects associated to exposures 

beyond optimal temperatures can be broadly generalized 

to other Arctic marine fish species. At the ecosystem level, 

the thermal disturbance (warming) could result in changing 

biogeography, biodiversity, seasonal timing, species 

interactions (mismatch), food web relationships, and overall 

ecosystem (including social interactions) function (Harley and 

others, 2006). At the population level, reduced densities might 

be expected as a result of changes in growth, reproduction, 

recruitment, and mortality. Changes in fish behaviors and 

demographic traits altered the physiological scope for growth 

in individual fish with cumulative effects on population 

processes. The physiological mechanisms operate at cellular 

and molecular levels and can change metabolic processes, 

and alter an organism’s functional integration, condition and 

health, and susceptibility to disease. Within this conceptual 

framework, it is also possible to envision more positive effects 

if warming enhances thermal conditions for a particular 

species life functions and behaviors resulting in possible 

population growth and expansion.

Ocean Acidification

Changes in ocean chemistry, known as ocean 

acidification, are an expected outcome of global warming. 

Ocean acidification (OA) occurs when increases in 

atmospheric carbon dioxide (CO2) cause an increase in the 

oceanic uptake of CO2

. This in turn leads to an increase 

in marine production of carbonic acid, reducing the pH of 

marine waters—ocean acidification. Ocean acidification 

changes ocean chemistry by reducing calcification rates 

and lowering calcium carbonate saturation (Mathis and 

Questel, 2013). Calcium carbonate is a key mineral used in 

calcareous shell production and other biological processes 

(Doney and others, 2009). The changes in ocean chemistry 

from ocean acidification are expected to be amplified in 

the Arctic Ocean17 with potential biological effects to food 

webs (that is, shell forming organisms such euphausiids) and 

marine fish populations (development and survival of early 

life stages of marine fish and foods) could be significant. In 

the U.S. Arctic, the ecological effects from amplification of 

potential acidification in freshwater discharges from the Yukon 

(195 km3

/yr) and Mackenzie (306 km3/yr) Rivers (Milliman 

and Meade, 1983), including nutrients, organic matter, and 

sediments into Chukchi and Beaufort Seas are not known. For 

comparison, the freshwater discharge from the Mississippi 

River into the Gulf of Mexico is 580 km3/yr. 

Shifts in Distribution of Marine Fish Populations

Climate change effects in the Arctic marine environment 

include loss of sea ice in summer, increased stratification, and 

shifts in the timing and intensity of the seasonal production 

cycle (Slagstad and others, 2011; Wassmann and others, 

2011). Several authors have attempted to project how these 

changes will affect marine fish populations (Reist and others, 

2006; Huse and Ellingsen, 2008; Mueter and Litzow, 2008; 

Pörtner and others, 2008; Cheung and others, 2009; Bluhm 

and others, 2011; Mueter and others, 2011; Sigler and others, 

2011; Hunt and others, 2013) and suggest potential effects 

on biogeography (for example, shifts in distributional), 

physiology (reproductive timing) and behavior (diet; 

Hollowed and others, 2013). Cheung and others (2009) 

projected that expanding bioclimatic windows would result 

in increased biodiversity in the Arctic. In contrast, Sigler and 

others (2011), projected that the shallow sill separating the 

northern Bering Sea and the Chukchi Sea, and the persistent 

presence of the cold water over the northern Bering Sea shelf 

(Stabeno and others, 2012a), would serve as a barrier to 

invasions of fish species into the region. Fishery baselines are 

changing rapidly throughout the Arctic; the possibility of new 

fisheries has increased the urgency for scientific attention to 

biodiversity and stock assessments (Christiansen and others, 

2014).

Mueter and Litzow (2008) described the southern edge 

of the cold water pool as defining the ecotone between Arctic 

and subarctic communities in the northern Bering Sea. The 

ecotonal boundary reportedly moved 230 km northward since 

the early 1980s, and Mueter and Litzow (2008) reported that 

community-wide shifts in distributions and related increases in 

biomass, species richness, and average trophic level occurred 

as subarctic species colonized new habitats. The warming 

temperatures were indicated as the primary cause of changing 

community distribution and function although other factors 

were implicated. Importantly, specific responses of individual 

populations to temperature changes were more difficult to 

estimate. The range extensions reported in this report are not 

indicative of significant new additions of subarctic fish species 

to the Arctic fauna. The effects of winter sea ice and thermal 

effects of the Earth’s inclination at high latitudes suggest cold 

water temperatures will persist in the Arctic and act to impede 

successful widespread expansions of many harvestable stocks 

17

Global warming trends have been amplified in the Arctic region relative 

to the global mean, and increased vulnerability of the Arctic, is due to the 

effects of increased freshwater input from melting snow and ice, temperature, 

and from increased CO2 uptake by the sea as a result of ice retreat (Fabry and 

others, 2009).
Chapter 4 657

from the Bering Sea. Given the ecological significance of 

Arctic Cod in Arctic marine food webs and energy cycling, a 

special analysis was done to describe climate change effects 

on Arctic and Saffron Cod in the northern Bering Sea (see 

chapter 5). 

In an Arctic-wide climate-change assessment, 

Hollowed and others (2013) examined the vulnerability of 

17 commercially exploited species in the Bering, Barents, 

and Norwegian Seas to move farther into the Arctic. The 

vulnerability assessment was based on exposure (to variations 

in the environment resulting from climate change), sensitivity 

(degree to which a species responds to variations in the 

marine environment that will be affected by climate change), 

potential affect (probable combined effects of exposure and 

sensitivity), and adaptive capacity (a species physiological 

or behavioral abilities to mitigate the effect). The potential 

for movement of the stocks was evaluated on climate change 

projections and expert opinion relative to life history (for 

example, reproductive ecology, feeding, and migratory 

behavior) and environmental preferences for temperature. 

Several of the Bering Sea species considered (that is, Walleye 

Pollock, Pacific Cod, Arctic Cod, Capelin, Arctic Skate, 

Greenland Halibut, Yellowfin Sole, and Bering Flounder) 

already occur in the Chukchi (all species listed) and Beaufort 

(Walleye Pollock, Arctic Cod, Arctic Skate, Capelin, and 

Greenland Halibut) Seas. Pacific Ocean Perch (Sebastes 

alutus) and Northern Rock Sole (Lepidopsetta polyxystra) 

were considered as having a low potential to move into the 

Arctic. Neither species has been collected from the U.S. 

Beaufort and Chukchi Sea. The potential for expansion of 

Walleye Pollock and Pacific Cod was determined to be low 

due to temperature and spawning site fidelity, respectively. 

Greenland Halibut and Capelin potentially could expand in the 

Arctic if temperature and conditions for larval transport and 

survival, or prey concentrations, respectively, were sufficient 

to sustain increased populations. Three stocks (Arctic Cod, 

Bering Flounder, and Arctic Skate) were determined to have 

a high potential to expand or move into the Arctic. Arctic 

Cod spawning occurs under ice and spawning areas will 

likely change if winter ice cover is lacking in the traditional 

spawning areas.

For many of the species identified by Hollowed and 

others (2013), the issue may not be the potential likelihood 

for northward movements, but climatic and habitat conditions 

(temperature , foods, substrates, and other conditions) for all 

life stages that increase the viability and relative abundance 

and expansion of small populations already present in the 

Arctic. Our analysis of marine fish diversity indicates that 

Pacific Cod and Greenland Halibut are uncommon in the 

Chukchi and Beaufort Seas. Alaska Plaice has been confirmed 

from the Chukchi Sea but not the Beaufort Sea. Yellowfin Sole 

is common in the Chukchi Sea but is rare in the Beaufort Sea. 

Available data support a hypothesis that natural selection 

favors individuals that maximize growth and energy efficiency 

at the expense of ranges of thermal tolerance (Pörtner, and 

others, 2008). Capelin is a cold water, circum-Arctic species 

(temperature preferences from -1 to 6 °C; Rose, 2005). In 

the U.S. Arctic, this species is locally abundant at various 

locations along the coast as spawning occurs in nearshore 

deltaic areas. The vulnerability of the species, or stocks 

adapted to Arctic conditions, to warming and their sensitivity 

to exposures exceeding a narrow range of temperature optima, 

is probably great. Under warming conditions, physiologicallymotivated shifts in distribution and seasonal abundance could 

result in cascading effects on regional food webs and changes 

in foraging behaviors. The disappearance of Capelin in the 

Gulf of Alaska during the temperature regime shift beginning 

in the mid-1970s was hypothesized to be in response to 

elevated temperatures (Anderson and Piatt, 1999; National 

Research Council, 1996). Generally, other marine species 

evaluated by Hollowed and others (2013) that are already 

occurring in the northern Bering Sea or southern Chukchi 

Sea (with the possible exception of Pacific Cod because of 

the species fidelity to spawning areas in the Bering Sea) 

increased abundances of species in the Chukchi Sea and 

further expansions onto the Beaufort Sea shelf and slope are 

considered a highly probable response to climate change.

Effects on Salmon Colonization

Pink and Chum Salmon are the most commonly reported 

species in the Alaska Arctic north of Point Hope (Nielsen 

and others, 2012). Small commercial fisheries operate in 

Kotzebue Sound and subsistence fishing occurs farther 

north. Small numbers of spawning Chum and Pink Salmon 

have been observed in rivers as far east as Prudhoe Bay in 

the southeastern Beaufort Sea (Craig and Haldorson, 1986). 

Whether the spawning observations to the north of Point 

Hope represented putative populations or straying continues 

to be unclear. Lack of suitable spawning habitats and 

extremely cold temperatures have been cited as reasons why 

this salmon straying has not led to sustainable populations 

(Irvine and others, 2009; Nielsen and others, 2012). However, 

each year increasing numbers of Chinook Salmon are being 

captured at Point Barrow and other sites along the coast. 

Over the next 30 years, as thermal barriers to migration 

and successful reproduction break down and, as freshwater, 

coastal, and marine habitats improve for salmon (for example, 

extended growing seasons, increased productivity, and other 

improvements) range expansions for chum, pink, and Chinook 

Salmon in Alaskan Arctic rivers seems certain. The successful 

colonization of these habitats by salmon is more difficult to 

estimate (Nielsen and others, 2012).
658 Alaska Arctic Marine Fish Ecology Catalog

Salmon Disease Ecology

Ichthyophonus in adult Chinook Salmon in the Yukon 

River has been linked to global warming and, because it is 

prevalent in marine fishes elsewhere in the northeast Pacific 

(for example, Pacific Halibut in the Gulf of Alaska), represents 

a potential threat to Arctic marine fishes (James Winton, U.S. 

Geological Survey, oral commun., 2013). Other diseases of 

fish, notably viruses of the Rhaddoviridae family (for example, 

viral hemorrhagic septicemia and infectious hematopoietic 

necrosis virus) would be novel pathogens in the Arctic marine 

fauna and potentially threaten members of the Salmonidae. 

Some ways that global warming can affect the severity and 

distribution of infectious diseases of fish include changes in 

the growth rate of pathogens, the types or strains of pathogens 

present, the distribution or biological carriers and reservoirs, 

the density or distribution of susceptible species, the diets that 

can alter resistance to disease, and the physical habitat that can 

affect disease ecology (Woodson and others, 2011). Although 

disease information is generally lacking, the anticipated 

effects of climate change, especially global warming, on fish 

diseases will include warmer temperatures and, in freshwater, 

lower flows that will exacerbate diseases caused by endemic 

parasites and pathogens, increase growth rates of pathogens, 

favor pathogens or strains that replicate at higher temperatures, 

and alter the strength and speed of host immune response 

disease (Hershberger and others, 2013). Additionally, altered 

freshwater and ocean conditions could change the distribution 

or density of hosts as well as the overlap with vectors, carriers, 

or reservoirs of infection. These altered habitats can produce 

biotic and abiotic stressors that will decrease the resistance to 

disease, and finally, a greater disease burden and associated 

fitness losses will increase the disease component of natural 

mortality among populations of fish.

Acknowledgments

Drs. Jim Winton and Paul Hershberger (USGS, Western 

Fisheries Research Center) provided valuable information 

about the potential effects of climate change on the disease 

ecology of marine fishes. 

Summary

Historical biogeography objectives of this synthesis 

describe the probable origins of distributional patterns as 

determined from systematic studies. The description of 

origins is complicated and largely hypothetical because the 

paleoceanographic record is not well-developed from the U.S. 

Arctic. Despite this, species origins were examined in light 

of possible dispersal and vicariance events as suggested by 

paleontologic and geologic records, climatic histories, and 

known phylogenetic relations. Information is limited, but 

when the known histories are compared with the phylogenetic 

record, preliminary results suggest that most marine fish 

species currently in the U.S. Chukchi and Beaufort Seas 

evolved from a Pacific ancestor. Ecological biogeography 

objectives addressed the environmental relations including 

physiochemical factors (for example, temperature, salinity, 

dissolved oxygen, and turbidity), mechanisms (currents, 

migrations, and movements), and biological processes 

(competition, predation, colonization, and reproduction) that 

limit the distribution, relative abundance, and productivity of 

a species. Global warming effects on this part of the Arctic 

are occurring at twice the rate of change in lower latitudes. 

Changes in regional sea ice coverage, physical and chemical 

oceanography, and hydrology are expected to have profound 

effects on coastal and marine ecosystems. Northerly shifts 

in the distributions of many migratory marine fishes, such 

as Pacific salmon, are expected with physical changes in 

temperature and process changes associated with benthicpelagic coupling over shelf waters.
1

U.S. Geological Survey.

2

U.S. Environmental Protection Agency.

Chapter 5. Arctic Climate Change—A Tale of Two Cods

By Deborah A. Reusser1, Melanie L. Frazier2, Rebecca A. Loiselle1, Henry Lee II2, and Lyman K. Thorsteinson1

Abstract

Climate effects on the ecology of the Arctic marine 

environments are of local, regional, national, and circumpolar 

interest and concern. Habitats are changing at the species level 

and there will be winners and losers with population limiting 

factors and the related temperature effects. In this chapter, 

the potential effects of warming temperatures on Arctic Cod 

and Saffron Cod in the eastern Bering Sea are evaluated. A 

Bering Sea study area was selected because, in contrast to 

the Chukchi and Beaufort Seas, a large volume of long-term 

fishery data was available for 1982–2006. A generalized 

additive modeling (GAM) approach was used to explore the 

effects of depth, bottom temperature, and surface temperature 

on the distribution and abundance of the cod species as well 

each species’ response to warming conditions. The GAMs 

estimate significant contraction and expansion in the marine 

distributions and abundances of Arctic Cod and Saffron Cod, 

respectfully, as simulated bottom temperatures increase in 

0.2 °C increments over shelf and slope habitats to a maximum 

of 2 °C. In the vernacular of “winners and losers,” the GAMs 

suggest that Saffron Cod will be a winner and Arctic Cod a 

loser in warming habitat conditions. The model simulation 

results were examined with respect to their application to 

species interactions and other ecosystem functions in the 

Chukchi and Beaufort Seas where each cod is more abundant 

and of greater ecological importance.

Introduction

Global climate change is an increasingly important 

ecological driver for oceanic and nearshore ecosystems, 

potentially affecting economically important fisheries stocks 

and ecosystem structure and function. Arctic systems are 

estimated to be especially vulnerable because of projected air 

and sea temperature increases (Anisimov and others, 2007). 

Summer sea ice extent in the Arctic has been decreasing at a 

rapid rate with some estimations indicating it will completely 

disappear by 2050 (Wang and Overland, 2009; Overland 

and Wang, 2013). Additionally, Arctic systems are changing 

almost twice as fast as anywhere on Earth from warming 

temperatures and melting ice (National Aeronautics and 

Space Administration, 2012); United Nations Environment 

Programme, 2013). Climate conditions in the eastern Bering 

Sea (EBS) have been highly variable over recent decades 

(Mueter and Litzow, 2008). In the EBS, winter sea-ice cover 

is the dominant factor creating a cold pool where bottom 

temperatures persist through the summer at less than 2 °C 

(Litzow, 2007; Stabeno and others, 2012b). Since 1954, this 

sea-ice cover in the EBS has decreased significantly coincident 

with an increase in summer bottom temperatures (Mueter and 

Litzow, 2008). The southernmost edge of the cold pool in the 

EBS has shifted about 230 km northward since 1982 (Litzow, 

2007). Benthic community shifts in the center of distributions 

northward also have been associated with these changing 

climate conditions in the EBS (Mueter and Litzow, 2008).

A major challenge becomes how to estimate the effects of 

these climate alterations on Arctic biota. Given the complexity 

of projecting future effects, numerous approaches are being 

used in terrestrial and aquatic environments. These approaches 

range from laboratory tolerance experiments (Wernberg and 

others, 2012) to trait-based analyses (for example, Moyle and 

others, 2013; Lee and others, U.S. Environmental Protection 

Agency, written commun., 2014 ) to more mechanistically 

based food webs (for example, Aydin and others, 2007) and 

individual-based models (Chang and others, 2010). Each 

approach has strengths and limitations, as well as drastically 

different data requirements.

This chapter resulted from cooperative research 

between the U.S. Geological Survey and U.S. Environmental 

Protection Agency focusing on potential climate effects on 

Arctic Cod and common interests in the species because of its 

human and ecological values. The research was implemented 

as a demonstration project to feature the application of a 

modeling approach to assess distributional shifts in important 

fish populations in response to potential climate change effects 

in the northern Bering Sea.
660 Alaska Arctic Marine Fish Ecology Catalog

Species Distribution Models

One approach that has gained considerable attention in 

projecting species responses to climate change is the use of 

statistically based species distribution models (SDMs, also 

known as niche or bioclimatic envelope models [Olden and 

others, 2008; Elith and Leathwick, 2009]). These models 

establish statistical relations between a species’ extant 

distribution and (or) abundance and a corresponding suite 

of environmental factors. This relation then can be used to 

project species’ responses to climate-induced environmental 

changes. A major advantage of SDMs is that they can use 

data from existing field surveys. Additionally, as pointed 

out by Heikkinen and others (2006), they are a “particularly 

valuable” tool for providing insights into effects of climate 

warming for species where the range-limiting physiological 

factors are poorly known. However, SDMs have a number 

of limitations (Heikkinen and others, 2006; Robinson and 

others, 2011). They require a moderately large dataset, and 

if effects of warming temperatures in northern latitudes are 

being investigated, it is important that the dataset include the 

southern range of the species. If abundance is being modeled, 

directly comparable sampling techniques are also essential. 

One concern of many ecologists is that most SDMs do not 

incorporate life history attributes or biological interactions 

and, accordingly, the simulated responses may not result from 

the direct effects of the environmental factors in the model, 

but rather result from indirect effects on productivity, predatorprey interactions, or other biotic interactions. Given this final 

limitation, it is perhaps best to view SDMs as statistical “idiot 

savants” and interpret the results through an ecological lens.

Species Selection and Climate Change

With these caveats in mind, SDMs were generated 

for two Arctic fish, Arctic Cod (Boreogadus saida) and 

Saffron Cod (Eleginus gracilis) in the EBS using sea 

bottom temperature, sea surface temperature, and bottom 

depths as key environmental drivers. The Arctic Cod and 

Saffron Cod were selected as target species, partly because 

of their importance to higher trophic levels and because 

they have different biogeographic distributions; therefore, 

they potentially display different responses to climate 

warming. After the models were generated, various climate 

change scenarios were applied to explore how increases 

in temperature are likely to alter their distributions and 

abundances in the EBS. Besides projecting potential changes 

in the EBS, changes that occur in the EBS are likely to be an 

early harbinger of changes in more northerly environments 

especially as they pertain to the potential for expanding 

commercial fisheries. To generate these models, the extensive 

bottom-trawl survey data from the National Oceanic and 

Atmospheric Administration (NOAA) Resource Assessment 

and Conservation Engineering (RACE) Division program 

(National Oceanic and Atmospheric Administration, 2013) 

were used. The RACE surveys in the EBS, Aleutian Islands 

(AI), and Gulf of Alaska (GOA) over the past three decades 

provide a large sample set for modeling and meet the criteria 

of having data from the southern part of the range for Arctic 

Cod and Saffron Cod. The circumpolar distribution of 

Arctic Cod and its intermediate position and importance in 

regional food webs makes this species a strong candidate as 

an indicator of environmental change and, in this regard, its 

interactions with congeners are of special interest.

Life History Considerations

The geographic range of Arctic Cod is circumpolar 

including the Chukchi Sea southward into the EBS. They are 

found from the surface to 1,390 m deep, although they were 

only captured from 30–150 m in the NOAA RACE surveys. 

Adaptations, such as anti-freeze proteins, allow this species 

to live in very cold waters and to use sea ice habitats to hide 

from predators, spawn, and forage for food. Spawning occurs 

in the late autumn–early winter. Eggs and larvae are pelagic, 

and larvae generally hatch in spring. Young Arctic Cod mostly 

feed on plankton in the upper water column, graduating to 

a diet of marine worms, adult copepods, and shrimps. Adult 

Arctic Cod grow to a maximum of 46 cm (Wienerroither and 

others, 2011) with an average length about 25 cm (Cohen and 

others, 1990) and are critical to marine food webs, providing 

food for other fishes such as Dolly Varden (Salvelinus malma), 

seabirds, and marine mammals. Although Arctic Cod are not 

fished commercially in North America, they are harvested in 

Russia and are considered an excellent table fish (Hebert and 

Wearing-Wilde, 2002). 

Saffron Cod also are in Arctic and subarctic seas and, off 

Alaska coasts, have been sampled as far south as the GOA. 

They typically are in nearshore waters less than 50 m deep. 

Physiological adaptations such as the antifreeze proteins in 

Arctic Cod also allow this species to survive in cold waters. 

In winter, they live under the ice in nearshore habitats and 

probably move slightly offshore in summer. Saffron Cod 

grow to a maximum length of 55 cm, live much longer, and 

are more likely to spawn on multiple occasions than Arctic 

Cod. Spawning occurs in winter and eggs are demersal. In 

early spring, eggs hatch and pelagic larvae are transported 

beneath the ice into tidally influenced bays/inlets of 2–10 m 

depths. Saffron Cod are epibenthic feeders and juveniles 

may associate with jellyfish. Like Arctic Cod, Saffron Cod 

are important in nearshore food webs and are consumed by 

other marine fishes, birds, and mammals. They are seasonally 

important in some coastal subsistence fisheries.

Species Interactions

Interactions between adult Arctic Cod and Saffron Cod, 

such as competition and predation, are not well known even 

though each species niche is similar with respect to trophic 

level and position (chapters 2 and 4). Neither species is 
Chapter 5 661

especially abundant in the EBS especially when compared to 

other cod and flatfish species. Under current environmental 

conditions, therefore, the existing information would at least 

tentatively indicate reduced competition between the species 

including limited predation by one species on the other. How 

differences in spawning season affect interspecific competition 

and predation are not known, nor are effects of cannibalism 

(an intraspecific interaction common in gadids) on population 

dynamics. The differences in vertical distribution and the 

apparent absence of biotic interactions are significant with 

respect to climate change modeling. If interactions exist, they 

could reduce the accuracy of model simulations. 

Methods

Data Source

Data were obtained from the NOAA Alaska Fisheries 

Science Center’s RACE Division (http://www.afsc.noaa.

gov/RACE/groundfish/default.php). The RACE resource 

assessment program was implemented in 1971 to monitor the 

condition of demersal fish and crab stocks on shelf regions 

off the coast of Alaska. This historical dataset for the Pacific 

Ocean and EBS includes trawl sampling by NOAA within the 

benthic zone of the United States continental shelf and slope 

areas of the U.S. Exclusive Economic Zone. 

The distribution of NOAA sampling varies by region and 

year, but fish monitoring protocols are consistent across the 

regions (Stauffer, 2004). In the EBS region, monitoring sites 

are located in the center of a regional grid of evenly distributed 

cells (37 × 37 km cells), with occasional samples collected 

at other sites in support of other research and management 

needs (fig. 5.1). In the EBS, monitoring stations were typically 

sampled every year (1982–2006), although Norton Sound in 

the northern Bering Sea was sampled only in 1985 and 1991.

For the AI and GOA regions, sampling stations were 

randomly selected from within grid cells (fig. 5.1) that 

were identified through management and ecological strata 

allocations. Sampling was biennial and took place between 

1984 and 2006 in AI and 1983 and 2007 in GOA. 

For the NOAA monitoring, physical attributes of each 

station were measured at the same time as all biological 

sampling and included bottom depth, bottom-water 

temperature, surface-water temperature, species composition, 

weight (all species), and number (fish only). Latitudes and 

longitudes at the start of the haul were used to describe 

sampling station locations. Only trawls with good performance 

(catch >0) were included in analyses (Stauffer, 2004). 

Figure 5.1. NOAA fishery-resource-assessment survey areas in the 

Eastern Bering Sea, Aleutian Islands, and Gulf of Alaska from NOAA RACE 

data collected between 1981 and 2006. 
662 Alaska Arctic Marine Fish Ecology Catalog

Trawls with haul depths inconsistent with reported station 

depths were excluded from analyses (n=6). Fish abundance is 

expressed in terms of catch (C in total numbers of individuals 

caught per station), relative abundance (numbers per 

kilometer), and density (numbers per kilometer and kilograms 

per hectare) in the RACE database. Initial examination of the 

trawl data revealed that the distributions of Arctic Cod and 

Saffron Cod were almost exclusively in the EBS (table 5.1). 

Therefore, species models were developed using only EBS 

station data (fig. 5.1). 

Statistical and Modeling Approaches

Frequency histograms of bottom depths, bottom 

temperatures, and surface temperatures were created from 

data collected at all stations sampled in the EBS, AI, and 

GOA. The histograms were compared to similar summaries 

from all stations where Arctic Cod and Saffron Cod were 

collected, to describe the niche spaces sampled for each 

environmental variable. The initial histograms including 

data from all survey stations sampled compared with similar 

presentations for each species, allowed comparisons of the 

total marine environmental space sampled and the niche 

space of each species. Generalized additive models (GAMs) 

were used to more rigorously quantify the effects of depth 

and temperature on cod occurrences (presence/absence) and 

abundances. GAMs are smoothing models used to fit nonlinear data using thin plate regression splines. All analyses 

were done in R 2.15.1 (R Core Team, 2012). The mixed GAM 

computation vehicle (mgcv) 1.7-18 package was used for the 

GAM analyses (Wood, 2003; 2006; 2011). A simultaneous 

confidence interval of 95 percent was used to delimit the 

credible regions (Wood, 2006) of each species simulated 

response to depth and temperature variables.

Only data collected from the EBS were used in the 

GAMs because Arctic Cod and Saffron Cod were rarely 

sampled in AI and GOA resource assessment areas. The 

observed effects of bottom temperatures (°C), surface 

temperatures (°C), bottom depths (m), and their interactions 

on abundance (C) as [ln (abundance + 1)] and distribution 

(presence/absence frequency of occurrence) were evaluated 

in the GAMs. Restricted maximum likelihood techniques 

were used to determine optimal smoothing functions in the 

regression analyses. The binomial family was used to estimate 

occurrence by depth and temperature relations. To select the 

best predictive model, we compared candidate GAM models 

with combinations of the three environmental variables 

including model runs with only one or two variables. Different 

smoothing functions also were evaluated. The best model for 

each fish species (occurrence and abundance) was selected 

from the candidate models using Akaike Information Criterion 

(AIC; Akaike, 1973). In addition to AIC, environmentalresponse curves were developed to select among models with 

similar AIC scores. Environmental response curves are a way 

of displaying model simulations. To generate these plots, the 

model was used to simulate a species probability of occurrence 

or abundance in relation to a single environmental variable 

while holding the other environmental variables constant. 

Spatial Interpolation of Environmental Data

Data from the World Ocean Database (National Oceanic 

and Atmospheric Administration, 2009) were downloaded 

and analyzed to examine the content and structure of bottom 

and surface temperatures to potentially increase the spatial 

extent of model simulations. Various interpolation schemes 

were applied to these data and tested for compatibility with the 

environmental values in the NOAA RACE data. Paired t-tests 

indicated that these data were not sufficiently compatible with 

the NOAA RACE data to expand the spatial extent of model 

simulations with any reliability; therefore, for this research, 

model simulations were limited to coverage within a smaller 

spatial extent that could be reliably interpolated with the 

NOAA RACE environmental data.

The three environmental variables (bottom depth, bottom 

temperature, and surface temperature) from the NOAA RACE 

program were spatially interpolated to create continuous 

coverage raster grids describing these variables across the 

EBS. An Inverse Distance Weighted (IDW) interpolation 

was completed in ArcGIS 10. The physical data for spatial 

interpolation were collected from a network of circular sample 

areas (37 km radius) in accordance with a random sampling 

plan that included greater than or equal to 12 sample points 

per area. The size of the circular area for spatial data collection 

was selected to approximate the grid size used in the NOAA 

RACE fish monitoring design. The resolution of all layers was 

set to 4.7 km. The data were then clipped to a buffer around 

the edge of the regular set of sampling stations to reduce 

extraneous information. The clipped region does not include 

Norton Sound because this area was sampled only during 

2 years and the temperature varied significantly between 1985 

and 1991. Consequently, although all EBS trawls were used in 

model development and analysis, data in Norton Sound were 

determined to be too sparse to create an interpolated average 

for each of the environmental surfaces. The clipped region is 

the spatial extent of estimated distributions and abundances of 

the cod species. 
Chapter 5 663

Simulated Effects of Climate Warming

To compare the accuracy of the model simulations to the 

measured data, the best GAMs for each fish species were used 

to simulate cod distributions and abundances based on the 

spatial interpolations of the environmental data. To evaluate 

the potential effects of climate change on each species, 

+0.2 °C was incrementally added to every cell of the spatially 

interpolated bottom-water temperature layer until the total 

change measured a +2 °C increase. Bottom temperature was 

varied because this variable had the greatest influence on the 

distributions and abundances of each species in GAMs. At 

each increment, GAMs were applied to established frequency 

baselines to predict changes in distributions (probability of 

occurrence) and abundances [ln (abundance + 1)] of the Arctic 

Cod and Saffron Cod. The predicted changes described are for 

total habitat and estimated numbers of fish. Total probability 

is calculated as the cumulative sum of individual probabilities 

of occurrence estimated for each species in all the grid cells 

in the EBS. Because the GAMs underestimate abundance, the 

predicted effects of bottom temperatures are best considered as 

indicators of relative change. The predicted changes also are 

shown as habitat maps for each species.

Results and Discussion 

Catch Characteristics of the RACE Surveys, 

1982–2006

Physical and biological data from 18,470 trawl samples 

from NOAA surveys off Alaska were used to evaluate climatewarming effects on the distribution and abundance of Arctic 

Cod and Saffron Cod in the EBS. Trawl depths ranged from 

0 to 1,000 m with most fish sampling occurring at stations 

located between 0 and 200 m. Bottom temperatures in the 

Bering Sea surveys ranged from 2 °C to about 14 °C with 

most measurements between 0 and 8 °C. Surface temperatures 

ranged from 0 to 16 °C with a few measurements indicating 

warmer temperatures.

Of the 9,423 trawls completed in the EBS between 1982 

and 2006, about 10.5 percent or 901 trawls contained Arctic 

Cod (table 5.1). Arctic Cod were widely distributed across the 

shelf and were sampled in Norton Sound and southeast of St. 

Lawrence Island (fig. 5.2). Highest abundance was associated 

with the coldest bottom temperatures in northern parts of the 

study area. 

Figure 5.2. Locations where Arctic Cod (Boreogadus saida) samples were collected by NOAA in the eastern 

Bering Sea (source: NOAA RACE database, 1981–2006) (National Oceanic and Atmospheric Administration, 

2013). The colors of the dots indicate the percentage of trawls in which Arctic Cod were captured. This 

analysis of presence-absence is possible because the same station locations are consistently sampled in the 

NOAA RACE resource assessment surveys. The NOAA RACE database includes samples collected in Norton 

Sound in 1985 (n=74) and 1991 (n= 47).
664 Alaska Arctic Marine Fish Ecology Catalog

In contrast, the highest abundance of Saffron Cod outside 

of Norton Sound was in Kuskokwim Bay and to the north 

and west of Nunivak Island (fig. 5.3). Saffron Cod were 

not collected in samples farther offshore (fig. 5.3). Of the 

9,423 trawls completed by NOAA in the EBS, 5.7 percent or 

531 trawls contained Saffron Cod (table 5.1). Both cod species 

were captured together in only about 1 percent or 109 trawls. 

About 81 percent or 89 of the 109 trawls that captured both 

cod species was in Norton Sound.

Table 5.1. Number of trawls completed by National Oceanic and 

Atmospheric Administration’s marine fishery resource assessment 

completed where Arctic Cod and Saffron Cod were collected in 

the eastern Bering Sea, Aleutian Islands, and Gulf of Alaska.

Trawls Eastern Bering Sea Aleutian Islands Gulf of Alaska

Total 9,423 3,202 5,845

Arctic Cod 901 0 0

Saffron Cod 531 0 7

Figure 5.3. Locations where Saffron Cod (Eleginus gracilis) samples were collected by 

NOAA in the eastern Bering Sea (source: NOAA RACE database, 1981–2006) (National 

Oceanic and Atmospheric Administration, 2013). The colors of the dots indicate the 

percentage of trawls in which Saffron Cod were captured. This analysis of presenceabsence is possible because the same station locations are consistently sampled in the 

NOAA RACE resource assessment surveys. The NOAA RACE database includes samples 

collected in Norton Sound in 1985 (n= 74) and 1991 (n =47).
Chapter 5 665

Environmental Variables and Models

The physical and biological variables reported in the 

NOAA surveys and used in this study are summarized in 

figure 5.4. The histograms present frequency data for selected 

measurements made throughout the three Alaska regions 

sampled (figs. 5.4a, d, g) and for capture sites of Arctic Cod 

(figs. 5.4b, e, h) and Saffron Cod (figs. 5.4c, f, i). 

The catch data indicate some overlap in species distributions 

in shallow waters (figs. 5.4b and c), but distinct differences 

in their response to bottom temperatures (figs. 5.4e and f). 

Arctic Cod were most frequently found at trawl stations 

having bottom temperatures of less than about 3 °C (fig. 5.4e) 

and Saffron Cod were more frequently detected in warmer 

waters (fig. 5.4f). These distributional relations were further 

supported in the GAM analyses (appendix C). Overall, 

Figure 5.4. Histograms showing bottom depth, bottom temperature, and surface temperature for all trawl samples 

collected in eastern Bering Sea, Aleutian Islands, and Gulf of Alaska. Trawl samples include 901 Arctic Cod and 538 Saffron 

Cod. Total samples, 18,470.
666 Alaska Arctic Marine Fish Ecology Catalog

the models that performed best (lowest AIC scores) for 

distribution and abundance of both species contained all three 

environmental variables. These models are highlighted in 

the tables included in appendix C. When individual physical 

variables were modeled as a predictor of abundance in the 

EBS, bottom temperature was the strongest single predictor 

with an r2 = 0.46 for Arctic Cod, and r2 = 0.35 for Saffron Cod. 

In comparison, surface temperature was the weakest single 

predictor with an r2 = 0.006 and r2 = 0.11 for Arctic Cod and 

Saffron Cod, respectively.

The southernmost edge of Arctic Cod distribution is 

coincident with the southern boundary of the cold pool and 

does not extend beyond the 2 °C isotherm in the EBS. The 

location of this isotherm and geography of the cold pool varies 

from year to year (fig. 5.5). The Arctic Cod distributional 

pattern relative to the cold pool is consistent with the results 

of our predictive distribution (occurrence by presence/

absence) and abundance modeling (appendix tables C1, 

C4, and fig. 5.6), which indicates the species is not likely 

to occur at locations where bottom temperatures are much 

warmer than 2 °C (fig. 5.6c). Catch and model simulations 

indicate that Arctic Cod are most common on the inner 

shelf and that their abundance generally decreases from 0 

to 200 m, with few fish predicted to occur at depths greater 

than 200 m (figs. 5.6a and b). A similar pattern of decreasing 

abundance with increasing bottom temperatures is evident 

in figure 5.6d. An interesting exception to this regional 

pattern occurred in the northern Bering Sea where the model 

predicted a slight increase in Arctic Cod abundance at 

bottom temperatures ranging between 2 and 4 °C, declining 

abundance as temperatures warmed to 7–8 °C, and a return 

to an increasing abundance pattern at temperatures higher 

than this. However, the uncertainty (dashed lines in fig. 5.6d) 

associated with estimated abundance at bottom temperatures 

greater than 4 °C is great and these results are questionable. 

The underlying statistical anomalies supporting abundance 

patterns in the northern Bering Sea may relate to interannual 

variations in hydrographic conditions across the northern 

Bering Sea shelf (Stabeno and others, 2012a; 2012b) and the 

associated interannual changes in the geographic extent of 

the cold pool. During warm years, the distribution of Arctic 

Cod would be geographically constricted in southern parts of 

its range providing at least a partial explanation for simulated 

abundance patterns in the northern Bering Sea. Arctic Cod 

responded to surface temperature with an optimal temperature 

of about 8 °C; however, this variable was less important than 

bottom temperature (figs. 5.6e and f). 

The presence/absence model selected to develop 

environmental response curves for Saffron Cod was not 

determined to be the best fit as indicated by AIC score, 

but it seemed to perform better than others based on the 

environmental response curves (appendix table C5). The 

model performed well enough to predict the probability of 

occurrence (r2

 = 0.42), and overall, demonstrated that depth, 

surface temperature, and bottom temperature are major 

determinants of this species patterns of distribution and 

abundance (fig. 5.7). The model simulation results indicated 

that Saffron Cod prefer a warm, shallow habitat and that 

although bottom temperature is not a strong predictor of 

distribution (fig. 5.7), it is for abundance (fig. 5.7d).

The selected models were used to predict the probability 

of species’ distribution and abundance in response to depth 

and temperature using the inverse distance weighted methods 

in ArcGIS 10 (fig. 5.8). The darkest area in figure 5.8b 

indicates the location of the cold pool.

Figure 5.5. Average bottom temperatures from the bottom trawl survey comparing the 

extent of the cold pool (<2 °C) during years warmer and colder than the 1982–2008 grand 

mean, eastern Bering Sea shelf (National Oceanic and Atmospheric Administration, 

2014).
Chapter 5 667

Figure 5.6. Environmental response curves for the selected presence/absence and abundance models for Arctic Cod 

in the eastern Bering Sea. Solid curves are the function estimates and dashed curves delimit the 95 percent confidence 

regions for each function.
668 Alaska Arctic Marine Fish Ecology Catalog

Figure 5.7. Environmental response curves for the selected presence/absence and abundance models for Saffron Cod 

in the eastern Bering Sea. Solid curves are the function estimates and dashed curves delimit the 95 percent confidence 

regions for each function.
Chapter 5 669

Figure 5.8. Physical description of the marine environment by (a) depth, and mean distributions of (b) bottom temperature, 

and (c) surface temperatures of the eastern Bering Sea (source: National Oceanic and Atmospheric Administration, 2012). 

Data from 1982–2006 interpolated using inverse distance weighted methods in ArcGIS 10. 

Model performance was evaluated by making 

comparisons of the observed and simulated patterns of 

Arctic Cod distribution and abundance over shelf and slope 

habitats as determined by depth and temperature attributes 

(fig. 5.9). Generally, the models performed reasonably well 

with respect to regional spatial patterns although they tended 

to underestimate the observed distribution and abundance 

associated with NOAA’s survey data. For example, the 

distribution of Arctic Cod presented in figure 5.9a closely 

approximates the model-predicted occurrence in figure 5.8b. 

The same relation holds true for abundance (figs. 5.9c and d). 

The analysis of temperature effects on the distribution 

(fig. 5.10) and abundance (fig. 5.11, table 5.2) of Arctic Cod 

shows a gradual reduction in the species range in the EBS 

with warming bottom temperatures. As bottom temperatures 

increase, the model also showed how the cold pool contracts 

northward and predicted that Arctic Cod distributions will 

similarly shift northward in association with the disappearance 

of cold water habitats. The main concentration of Arctic Cod 

to the southwest of St. Lawrence Island almost completely 

disappears with an overall increase of 2 °C. There is great 

uncertainty associated with predictions of increasing abundance 

in the more southern areas of the species range, which reflects 

the difficulties of quantitative analysis with sparse datasets due 

to low detection rates of Arctic Cod in warmer environments.

The GAM models underestimated reported distribution 

and abundance patterns of Saffron Cod developed from NOAA 

catch data (fig. 5.12, table 5.3). For instance, the difference 

between observed and modeled patterns is especially evident 

in this species distribution in figures 5.12a and b. The reason 

for the difference may relate to the Saffron Cod’s apparent 

restricted distribution near the Yukon Kuskokwim Delta 

and the limited sampling in the RACE surveys in nearshore 

environments of the inner shelf compared to offshore areas 

where commercial fisheries occur. The low frequency of Saffron 

Cod occurrence in trawl catches in deeper and colder parts of 

the EBS is probably responsible for the model’s performance. 

It is important to remember that the value of the models is as 

a tool for simulating relative changes in response to changing 

environmental conditions (figs. 5.13 and 5.14).
670 Alaska Arctic Marine Fish Ecology Catalog

Table 5.2. Changes in Arctic Cod habitat and relative abundance simulated with incremental warming of bottom 

temperature (+ 0.2 °C increments) and where surface temperature and depth are constant in the eastern Bering Sea.

[°C, degrees Celsius]

Simulated change in total Arctic Cod habitat Simulated change in Arctic Cod abundance

Temperature

increase

(°C)

Total

probability 

Total area

(percent)

Total abundance

(number of fish)

Relative abundance 

compared to

time zero

(percent)

Time Zero 3,274 12.12 39,416 100

0.2 2,825 10.45 27,884 71

0.4 2,397 8.87 19,950 51

0.6 2,002 7.41 14,525 37

0.8 1,654 6.12 10,857 28

1.0 1,363 5.05 8,434 21

1.2 1,137 4.21 6,922 18

1.4 974 3.61 6,083 15

1.6 869 3.22 5,757 15

1.8 811 3.00 5,833 15

2.0 790 2.92 6,212 16

Figure 5.9. Observed and simulated distribution and abundance of Arctic Cod in the eastern Bering Sea (source: National 

Oceanic and Atmospheric Administration, 2012) . Simulations were generated using best performing Generalized Additive 

Models and fishery resource data from 1982 to 2006 (see appendix C).
Chapter 5 671

Figure 5.10. Changes in the distribution of Arctic Cod (probability of occurrence) in the eastern Bering Sea as simulated 

bottom temperatures increase in 0.2 °C increments. 
672 Alaska Arctic Marine Fish Ecology Catalog

Figure 5.11. Changes in the abundance of Arctic Cod in the eastern Bering Sea as simulated bottom temperatures increase 

in 0.2 °C increments.
Chapter 5 673

Figure 5.12. Observed and simulated distribution and abundance of Saffron Cod in the eastern Bering Sea (source: National 

Oceanic and Atmospheric Administration, 2012; fishery resource data from 1982–2006). Simulations were generated using 

best performing GAM models (see appendix C).

The models for Saffron Cod and Arctic Cod predict 

increases in the range and abundance in and around 

Kuskokwim Bay as bottom temperatures increase. This is 

probably a modeling artifact for Arctic Cod but not for Saffron 

Cod. The model simulation results shown in figures 5.12

and 5.13 suggest that habitats more suitable for Saffron Cod 

than for Arctic Cod would be created with warming bottom 

temperatures. The models predicted an increase in Saffron 

Cod distribution to the north, south, and offshore (fig. 5.12) to 

and potentially at depths greater than 50 m beyond the outer 

boundary of the coastal domain described for EBS (Kinder 

and Schumacher, 1981). Similarly, as bottom temperatures 

increase, Saffron Cod abundance shows a gradual increase in 

Kuskokwim Bay northward beyond Nunivak Island, south to 

Bristol Bay and farther offshore (fig. 5.13). 

Table 5.3. Changes in Saffron Cod habitat and relative abundance simulated with incremental warming of bottom 

temperature (+0.2 °C increments) and where surface temperature and depth are constant in the eastern Bering Sea.

[°C, degrees Celsius]

Simulated change in total Saffron Cod habitat Simulated change in Saffron Cod abundance 

Temperature

increase

(°C)

Total

probability 

Total area

(percent)

Total abundance

(number of fish)

Relative abundance 

compared to

time zero

(percent)

Time Zero 1,487 2.83 14,139 100

0.2 1,499 3.08 18,493 131

0.4 1,516 3.29 24,362 172

0.6 1,537 3.49 32,216 228

0.8 1,562 3.69 42,649 302

1.0 1,593 3.89 56,400 399

1.2 1,628 4.08 74,371 526

1.4 1,667 4.26 97,638 691

1.6 1,710 4.48 127,454 901

1.8 1,757 4.74 165,274 1,169

2.0 1,806 3.92 212,793 1,505
674 Alaska Arctic Marine Fish Ecology Catalog

Figure 5.13. Changes in the distribution of Arctic Cod (probability of occurrence) in the eastern Bering Sea simulated as 

bottom temperatures increase in 0.2 °C increments.
Chapter 5 675

Figure 5.14. Changes in the abundance of Saffron Cod in the eastern Bering Sea simulated as bottom temperatures increase 

in 0.2 °C increments.
676 Alaska Arctic Marine Fish Ecology Catalog

Norton Sound

The NOAA RACE database accessed herein included 

2 years (1985 and 1991) of data collected from Norton 

Sound. Although Arctic Cod were frequently caught in the 

trawl catches, the thermal environmental conditions seem to 

be different to the preferred ranges suggested in figure 5.4. 

Mean bottom temperatures in the Norton Sound trawls were 

relatively warm in both years (7 °C in 1985 and 6.1 °C in 

1991) and, on first inspection, seem to approach or exceed the 

upper limits of thermal preferentia for this species. Perhaps 

this result reflects greater thermal tolerance for this species 

than described by the RACE data set. The occurrence of 

Arctic Cod also could be indicative of bathymetric migrations, 

habitat differences by life cycle stages, or other kinds of 

ecological benefits conferred by warmer waters. An example 

of the latter could be physiological (for example, growth) or 

related to increased prey abundance that overrides optimal 

temperature preferences.

The model outputs would also suggest that, because 

the mean bottom temperature decreased from 7 °C in 1985 

to 6.1 °C in 1991, the distribution and abundance of Arctic 

Cod should have increased. However, although mean bottom 

temperatures showed a 0.85 °C decrease between 1985 

and 1991, the presence and abundance of Arctic Cod also 

decreased from 65 of 74 trawls in 1985 to 24 of 47 trawls in 

1991. Likewise, the average number of Arctic Cod per trawl 

in Norton Sound in 1985 was 82.7 compared to 6.9 in 1991. 

Corresponding changes in surface temperatures, however, 

showed an opposite trend with a mean temperature of 6.6 °C 

in 1985 and 11.7 °C in 1991. 

Although Arctic Cod in Norton Sound generally respond

negatively to increases in bottom temperatures, other factor(s), 

such as food abundance, may be involved. The large volumes 

of freshwater discharged by the Yukon and Kuskokwim Rivers 

creates hydrographic conditions in Norton Sound that may be 

similar to conditions in the nearshore Alaska Beaufort Sea in 

summer. The regional comparison is significant in that Arctic 

Cod are a common, if not dominant species, in nearshore 

waters farther north. Their occurrence in marine and brackish 

habitats of the Beaufort Sea has been related to invertebratedriven food webs that are also important in Norton Sound 

(Thorsteinson and others, 1989). In comparison, the mean 

abundance and distribution of Saffron Cod remained about the 

same in 1985 and 1991 despite the observed environmental 

changes. A potential future project could include improving 

the models as more data become available in this area and 

farther north. 

Arctic Applications

The overlap in habitats of Arctic Cod and Saffron Cod 

is much greater in the Chukchi and Beaufort Seas (chapter 3) 

than has been described for the EBS. However, because of 

the incomplete nature of fishery data from the high Arctic, 

it is difficult to describe how changing distribution patterns 

might affect interactions between these species. It is known 

that Arctic Cod play a keystone role in the bioenergetics of 

Arctic food webs as a consumer of primary and higher levels 

of biological production and are a significant prey resource 

for many marine fishes, birds, and mammals. The potential 

for large-scale changes in the forage base of the Chukchi and 

Beaufort Seas suggests the possibility for cascading ecosystem 

effects and corresponding needs for long-term and multivariate 

approaches to data collection and greater attention to 

scientific goals for population and functional understanding. A 

broadening of the ecological lens in climate change modeling 

is needed to understand the adaptive capacity and resilience 

of these species and their interaction with others. Although 

Arctic and Saffron cods are not currently of commercial 

importance in North America, their ecological and subsistence 

values are considerable and worthy of marine indicator status. 

Environmental changes that affect their distribution and 

abundance could alter the structure and function of the marine 

ecosystem and change the traditional and economic uses of 

these and other organisms in Arctic food webs.

The SDM results suggest that based on warming 

temperatures, without viewing climate effects more broadly, 

similar distributional changes as predicted for the EBS 

might be projected for the Chukchi and Beaufort Seas. These 

changes include a constriction in range to the north in Arctic 

Cod and a similar expansion in range to the north by Saffron 

Cod. Warming of bottom temperatures in the northern Bering 

Sea can be expected to result in other large-scale shifts in 

distributions of other marine fishes including Walleye Pollock, 

a species that, although presently known from the Chukchi 

and western Beaufort Seas, seems to be limited in numbers 

and biological condition in cold-water environments. Pollock 

are extremely abundant in the EBS and the comparatively 

low relative abundances of Arctic Cod and Saffron Cod 

relates to the zoogeographic boundary in the northern Bering 

Sea discussed in chapters 2 and 4. The observed distribution 

and abundance data from the EBS suggests that deeper shelf 

and upper slope habitats are not occupied by Arctic Cod, in 

contrast to distribution and abundance in the Chukchi and 

Beaufort Seas, which suggests predation and other competitive 

processes with Pollock and other species, in addition to 

temperature, may be occurring. 
Chapter 5 677

Favorable environmental temperatures associated with 

possible large-scale changes toward more pelagic regimes 

over the Chukchi Sea shelf could result in difficult to estimate 

changes in predator-prey dynamics and species composition 

of regional fish assemblages (Grebmeier and others, 2006a). 

Questions about the stock composition in Arctic Cod and 

Saffron Cod populations and areas of life history importance 

(for example, spawning locations) are currently outstanding 

and thus expansions to the north would not necessarily 

translate to the Arctic Cod’s extirpation from the Chukchi 

Sea. Movements to deeper, colder and more stable marine 

environments might remove these species from easily 

accessible coastal fishing areas, or at least make harvest more 

difficult, for subsistence users.

Acknowledgments

The authors wish to thank the U.S. Department of the 

Interior, Bureau of Ocean Energy Management (BOEM), 

Environmental Studies Program, the U.S. Environmental 

Protection Agency (EPA), the U.S. Geological Survey (USGS) 

through the Outer Continental Shelf (OCS) Studies Program, 

the USGS Western Fisheries Research Center, and the USGS 

Alaska Region for supporting this study. The EPA provided 

support through Interagency Agreement Number DW-14-

92231501-1. The editors and chapter authors appreciate the 

support of this study through the EPA’s National Health and 

Environmental Effects Research Laboratory (NHEERL). 

Technical reviews were managed by the BOEM Alaska 

OCS Region, the USGS, and the NHEERL Western Ecology 

Division. The authors wish to thank the following EPA 

employees for their contributions to this report: Patrick 

Clinton, for providing input and guidance into interpolation 

methods within ArcGIS and Lee McCoy for collaboration in 

developing the R scripts to generate the maps of predicted 

distribution and abundance of the two cod species.

Summary

The GAMs generated using NOAA’s RACE survey data 

appear to be sufficiently dynamic to simulate the general 

response of Arctic Cod and Saffron Cod to climate-related 

changes in deeper shelf waters of the EBS. The modeling 

results suggest that both species likely will be affected by 

global warming trends, but in different ways. Analysis of fish 

catch and bottom water temperature data from the NOAA 

surveys further suggest that the southern boundaries of the 

cold pool in EBS vary annually in spatial extent and generally 

correspond to the southern range limit of Arctic Cod, an 

endemic Arctic species. It is hypothesized that Arctic Cod 

distribution in the northern Bering Sea will shift (contract) to 

the north with warming bottom temperatures. The presence 

of sea ice in the Bering Sea during winter and persistent cold 

bottom temperatures to the south of St. Lawrence Island will 

likely dampen this warming effect for the foreseeable future. 

The model simulations presented in this chapter 

support this premise and indicate that Arctic Cod are 

sensitive to changes in bottom temperature and depth. 

For example, the model simulations indicate that small 

increases in bottom temperature will dramatically reduce 

their distribution and abundance. Significantly, a 2 °C 

increase in bottom temperatures is expected to reduce Arctic 

Cod distribution in the EBS by approximately 80 percent 

and its relative abundance by 90 percent. By comparison, 

the model simulations indicate a nearly opposite effect 

on Saffron Cod. Their distribution, with a 2 °C increase 

in bottom temperatures, would be expected to expand by 

approximately 28 percent with a corresponding increase 

in abundance of 92 percent. As data become available, the 

application of GAMs in the Chukchi and Beaufort Seas would 

enhance assessments of effects of climate change on marine 

ecosystems and subsistence in regions where Arctic Cod, and 

to a lesser extent Saffron Cod, are more important in food 

web dynamics.
678 Alaska Arctic Marine Fish Ecology Catalog

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1

U.S. Geological Survey.

Chapter 6. Conservation of Arctic Alaska’s Marine Fish 

Resources

By Lyman K. Thorsteinson1

Abstract

Limitations in data and scientific uncertainties in our 

understanding of Arctic marine fish ecology are pronounced 

and indicative of the science that will be needed for 

effective resource management, environmental regulation, 

and conservation. The existing datasets are not conducive 

to quantitative evaluations of population dynamics or 

determinations of species abundance, and in most instances, 

it is not possible to estimate vital demographic parameters 

such as age and size structure, instantaneous growth rates, or 

mortality and survival rates. Cost and logistical constraints 

associated with Arctic research will likely focus future 

efforts on monitoring of trends in these key population 

parameters rather than actual changes in fish population sizes. 

Physiological data are lacking especially with respect to our 

understanding of key environmental effects on rates of growth, 

reproduction, and survival. Field and laboratory experiments 

will be needed to develop bioenergetics models to assess 

climate change effects or disturbances, such as possible Arctic 

oil spills. For now, in the absence of population understanding, 

information about species life history and ecological traits 

can be used to classify Arctic marine fish species into 

strategic groups based on the r-K theory to assist resource 

management and environmental decision-making including 

two previously undescribed groups we call Amphidromic 

and Cryophilic Strategists. Knowledge about the seasonal 

diets and food habits of dominant Arctic fishes are primarily 

known from stomach samples collected during ice-free 

months. Similarly, descriptions of predator-prey relationships 

and biological interactions tend to be limited to prominent 

species in food webs leading to apex predators. Integrated 

and interdisciplinary science approaches are priorities for 

long-term data collection that will be needed for the Arctic 

fishery-based ecosystem management recommended by 

the U.S. Department of Commerce. The information needs 

for 21 priority marine species identified in chapter 3 are 

considered in light of integrated research and monitoring in 

the Chukchi and Beaufort Seas.

General Understanding of Marine Fish 

Life History

Much new biological information about marine Arctic 

fishes and their environmental relationships was acquired, 

reviewed, and synthesized in this study. It is an important 

synthesis because it expands the nature of information 

presented in the Fishes of Alaska (Mecklenburg and others, 

2002) and along with biodiversity baseline presented in 

Mecklenburg and Steinke (2015) and Mecklenburg and others 

(2016) significantly updates the inventory represented for the 

Arctic in that monograph. The update of this information was 

timely because the user community is increasingly large and 

it requires accurate resource inventories and access to data 

and information. The need extends beyond a listing of species 

(Cook and Bundy, 2012) and, given the wide variety of uses 

for resource information, requires a consistent and reliable 

source of scientific information for each species (geographic 

distribution, abundance, and habitats). Although life history 

information is lacking for most species, it is generally true 

that, within a life cycle context, the most information is 

available for adult stages and much less is known about 

juvenile marine fish and younger stages (chapter 3). The 

ichthyoplankton sampling that has been done is an important 

source of information about the biodiversity of the region. 

This synthesis is therefore timely for purposes of 

assessing potential climate effects and informing decision 

making about regional fisheries, offshore energy development, 

and other human uses (for example, increased marine shipping 

and tourism, and port developments). It is an encyclopedic 

effort with respect to content (biodiversity, life history, 

population ecology, and regional ecosystems), visualization 

of data, and scientific information presented and reviewed. 

The narratives, maps, and literature references provided in 

individual species accounts are meant to serve as trailheads or 

guides to more detailed information about the zoogeography, 

taxonomy, life histories, species niches, and life requirements 

in sectors of the Arctic marine ecosystem of the United States. 

We have attempted to present scientific information and 

concepts in language that will be understood and useful not 

only to professional scientists and resource managers but the 

broader swath of user communities involved in environmental 

research, community planning and development, and Arctic 

policy making.
680 Alaska Arctic Marine Fish Ecology Catalog

Applications for Arctic Outer Continental Shelf 

Energy Development

The Chukchi and Beaufort Seas differ with respect to 

geographic setting, expanse of continental shelves, regional 

oceanography, and freshwater-marine interactions (rivers and 

streams). The differences (for example, topography of the 

shelf and slope habitats) affect patterns of species occurrence 

in Arctic marine habitats. The Chukchi Sea shelf is broad 

and generally less than 200 m deep. In contrast, the Beaufort 

Sea shelf is narrow and slope waters are deep (>1,000 m). 

The presence of sea ice is an important feature in both seas 

as it relates to environmental temperatures, light penetration, 

habitat for species such as the Arctic Cod (food, shelter, and 

potential nursery for early life stages), and processes affecting 

production cycles.

This report documents the confirmed occurrence of 

109 species of known marine fishes in 24 families and 

63 genera in the U.S. Chukchi and Beaufort Seas. The 

occurrences were based on published literature, specimens 

examined in museum collections, and species confirmed from 

expert identifications in ongoing field research. Of these, our 

review of Alaskan records indicates 97 species are found in 

the Chukchi Sea and 83 in the Beaufort Sea. Sixty-eight fishes 

are common to both seas. Since the latest publication covering 

all known Alaskan fishes (Mecklenburg and others, 2002), 

18 new species to the region have been confirmed. In time, 

other species will be confirmed from this region. For example, 

at least three other species whose taxonomic identification is 

in progress are known, as well as several others marine fishes 

that are likely, but have not been observed in sampling. Given 

the intensity of sampling in nearshore areas, new species and 

range extensions are expected from these waters.

The NEPA process requires that BOEM analysts use the 

best resource information available in their assessments of 

environmental effects associated with offshore oil and gas 

development in Arctic OCS Planning Areas. Through review 

and synthesis, our summaries provide current information 

about which fishes are present, when and where they are 

present, and the physical and biological processes that affect 

their distribution and abundance. This information is valuable 

because in reporting what is known about the Arctic marine 

fish fauna and its traditional, cultural, and economic values, 

this study also indicates where further study is needed. 

The immediate applications of this work extend beyond 

the BOEM’s purposes and are of immediate value to other 

Federal and state research and management agencies to Alaska 

Native and other Arctic residents interested in Arctic marine 

ecosystems, potential Arctic fisheries, and the conservation of 

Arctic marine fishery resources. At the same time, the format 

and information presented, especially in the species accounts, 

are intended to be useful to a public audience. 

Theoretical Applications to Resource 

Management

Information about species-specific and interspecific 

life history and ecological traits informs not only theoretical 

ecology (such as r-K adaptations), but has implications 

for resource management (Hardie, 2003). The relationship 

between multiple traits is addressed in the fast-slow continuum 

hypothesis, which “explains life history traits as reflecting 

the causal influence of mortality patterns in interaction with 

trade-offs among traits, particularly more reproductive effort 

at a cost of shorter lives” (Paemelaere and Dobson, 2011). 

Bjørkvoll and others (2012) recently examined this hypothesis 

with respect to the population dynamics of nine species of 

marine fishes from the Barents Sea. A major finding was 

that mean natural mortality rates, annual recruitment, and 

population growth rates were lower in long-lived species 

(slow end of the continuum) than in short-lived species (fast 

end). Interspecific characteristics also were associated with 

ecological traits where it was determined that species at the 

fast end were mainly pelagic with short generation times and 

high natural mortality, annual recruitment, population growth 

rates, and high temporal variability in these demographic 

traits. In contrast, species at the slow end of the continuum 

were long-lived, demersal species with low rates and reduced 

temporal variability in the same demographic traits. 

Life History and Ecological Traits

The relations between basic life history characteristics 

and population dynamics have implications for managing 

marine fish resources where population data are limited or 

non-existent, like the Arctic. To illustrate, in a conceptually 

similar analysis, Winemiller (2005) used a triangular model 

(Kawasaki, 1980: Winemiller and Rose, 1992) to classify 

the life history traits of mature individuals of marine fishes 

into life history strategist groups (based on position within 

the fast-slow continuum) to investigate their predictive 

capacity and management applications. King and McFarlane 

(2003) completed a much larger analysis (n = 42 species) 

and identified 5 strategist groups represented by species 

common to pelagic, benthic, and nearshore habitats in the 

western Gulf of Alaska. Two of the strategic groups had been 

previously described by Winemiller and Rose (1992) including 

opportunistic and periodic strategists. Three additional distinct 

groupings, equilibrium strategists, salmonic strategists, 

and intermediate strategists were described by King and 

McFarland (2003). King and McFarlane (2003) described each 

group as:
Chapter 6 681

Opportunistic Strategists.—Opportunistic strategists 

are short-lived with a small body size at maturation, low 

fecundity, high growth rates, and small eggs. They are surface 

and midwater pelagic species that exhibit little if any parental 

investment and are planktivores or lower-order carnivores. 

They occupy habitats that have a high degree of environmental 

variability but potentially large resources of energy. As such, 

their population responses tend to be large in amplitude and 

species in this group display highly variable, fluctuating 

population patterns. Species include clupeids (for example, 

Pacific Sardine, Pacific Herring), smelts (for example, 

Eulachons), and other forage fishes (for example, Northern 

Lampfish and Arctic Sand Lance). Because abundance is 

dynamic and survival rates are variable, the opportunistic 

strategists are susceptible to rapid depletion augmented 

by fisheries.

Periodic Strategists.—Periodic strategists are long-lived 

and slow growing with a high fecundity, but are medium in 

size, have a midrange for size at maturity, and have medium 

sized-eggs. These fishes are higher-order carnivores that 

inhabit shelf or slope benthic habitat and exhibit some parental 

investment. In the eastern Pacific, rockfishes and flatfishes 

are good examples of periodic strategists. These species were 

classified as having a steady-state population pattern. The 

period between strong year classes can be relatively long 

(as much as several decades), and these species can exhibit 

decadal scale patterns in recruitment coincident with climate–

ocean regimes. Annual recruitment is only a fraction of the 

spawning stock biomass, and maintaining an appropriate 

age-structure in the spawning stock biomass should be a 

paramount management goal for these relatively long-lived, 

late maturing species. 

Equilibrium Strategists.—Equilibrium strategists are 

dominated by elasmobranchs (skates and sharks), which 

are slow growing, have low fecundity, are large in size, and 

have large eggs (K selection traits). These species exhibit 

modest to great degrees of parental investment, are higher 

order carnivores and piscivores, and inhabit a range of 

habitats. Equilibrium strategists have a low fecundity and late 

maturation, and therefore, not able to recover as quickly as 

other fishes after population reduction.

Salmonic Strategists.—Salmonic strategists, 

Oncorhynchus spp., are relatively short-lived, but fast growing 

and large sized. Compared with other marine species, they are 

not extremely fecund, but have large eggs. The life history 

traits of the Pacific salmon differ from opportunistic strategists 

with their semelparous nature and the higher degree of 

parental investment. Large-scale changes in ocean condition 

affect salmon productivity. Ocean survival, notably during 

the first marine summer conditions, may be especially critical 

in the salmon strategist population dynamics. Improved 

estimation of freshwater (egg-smolt production) and marine 

(smolt-adult production) survival in population models will 

aid management forecasts.

Intermediate Strategists.—Intermediate strategists, 

cods and scombrids, have life history traits that are mid-range 

when compared to opportunistic and periodic strategists. They 

have a longer life span than the opportunistic strategists, with 

maximum ages typically 10–20 years, but exhibit the same 

population dynamics as this group. Gadids are considered 

typical groundfish species (that is, benthic or bathypelagic), 

but are different from the other groundfish species (for 

example, rockfish, flatfish, and sablefish) grouped with 

periodic strategists. For example, gadids are not as long-lived 

and scombrids are highly migratory surface pelagics, but 

differ from the other surface pelagics because of their larger 

size and longevity. Populations of intermediate strategists can 

withstand periods of unfavorable environmental conditions 

for recruitment better than the opportunistic strategists, but 

they do not exhibit the more stable populations as in periodic 

strategists. Their shorter generation time makes them more 

vulnerable to fluctuations in biomass through fluctuations in 

recruitment. Intermediate strategists are large-sized, highly 

migratory pelagic species that are able to move from areas of 

poor conditions to areas of better conditions as reflected in 

large distributional changes.

The study by King and McFarlane (2003) included 

species from the Gulf of Alaska where fish life histories and 

population dynamics and trends are relatively well known 

from long-term research. The species accounts demonstrate 

that, although in many instances, life history and ecological 

trait information is not available from United States waters, 

the general information that is available is of sufficient quality 

to identify the relative positions of certain species on the 

fast-slow continuum. Additionally, the similarities between 

Arctic and subarctic traits allow an initial classification of 

Arctic marine fishes using the strategist grouping of King and 

McFarlane (2003) as an initial screen. 

The assignments are a first-order projection and, as such, 

are preliminary in nature; however, as noted by King and 

McFarlane (2003), in the absence of information on absolute 

or relative biomass (Arctic case) this conceptual framework 

can guide management options. Although a comprehensive 

analysis of the entire fauna is not feasible, initial decisions 

about (1) what species to include were based on abundance 

(that is, selected species were common in both Chukchi and 

Beaufort Seas), and (2) assignment to strategic groupings were 

based on life history knowledge including resiliency (that is, 

estimates of population doubling time). The Arctic marine 

fish strategic groupings for selected dominant species are 

shown in table 6.1 following criteria described by King and 

McFarland (2003).
682 Alaska Arctic Marine Fish Ecology Catalog

Table 6.1. Life history strategy groupings of common Arctic marine fishes in the Chukchi and Beaufort Seas.

[Resiliency: Based on estimates of population doubling time: low (4.5–15 years); medium (1.4–4.4 years); and high (<15 months) turnover. Life history:

Information is based on King and McFarland (2003). <, less than]

Taxonomic

family Resiliency

Life history strategic grouping

Opportunistic Periodic Equilibrium Salmonic Intermediate

Common fish species

Petromyzontidae Low Arctic Lamprey

Squalidae Low Spotted Spiny 

Dogfish

Rajidae Low Arctic Skate 

Clupeidae Medium Pacific Herring 

Osmeridae Medium Arctic Smelt, 

Capelin

Salmonidae High Chum Salmon

Myctophidae1

Gadidae Medium Walleye Pollock, 

Saffron Cod 

Gasterosteidae Medium Ninespine 

Stickleback

Cottidae Low to high Hamecon, 

Fourhorn Sculpin, 

Shorthorn Sculpin

Agonidae Medium Alligatorfish, 

Arctic Alligatorfish, 

Veteran Poacher

Liparidae Medium Gelatinous Snailfish,

Kelp Snailfish

Zoarcidae Low to medium Fish Doctor, 

Marbled Eelpout, 

Polar Eelpout

Stichaeidae Low to medium Fourline 

Snakeblenny, 

Slender Eelbenny, 

Arctic Shanny

Ammodytidae Medium Arctic Sand Lance 

Pleuronectidae Low to medium Bering Flounder, 

Arctic Flounder 

1

Not commonly found in Chukchi or Beaufort Seas.
Chapter 6 683

Two additional strategic groupings are described to 

capture other adaptation strategies known to the Arctic marine 

fish fauna:

Amphidromic Strategists.—The amphidromous 

life history strategy is significantly different from the 

strategies of other Arctic and subarctic marine fishes. For 

this reason, we created a new life history strategy grouping, 

amphidromic strategists. Amphidromous strategists (that is, 

Dolly Varden char; Least Cisco, and Bering Cisco; Broad 

and Humpback Whitefish; and Inconnu) strongly exhibit 

K selection traits (Craig, 1984) and are similar to salmonic 

strategists with respect to migratory behavior, occupation of 

multiple habitats, and parental investment in young. Unlike 

the salmonic strategists they are iterperous and long-lived 

(>20 years). Density-dependent factors operating in freshwater 

overwintering sites result in a relatively steady state. The 

Arctic Cisco life cycle in the Alaska Beaufort Sea involves a 

long-distance migration of young-of-the-year (YOY) juveniles 

between the McKenzie River, Canada and Colville River, 

United States. Overwintering of immature ciscoes (generally 

<7–8 year old fish) in Alaskan waters occurs in salinityinfluenced areas of the delta and thus the species shares some 

anadromous characteristics with the salmonic strategists. The 

abundance of migrant YOY ciscoes is variable and survival 

rates, at least during early life phases, are similar to the 

opportunistic strategists.

The amphidromic strategists include key, iconic species 

of Arctic marine fishes. Long-term harvest records for Arctic 

Cisco from a small fishery in the Colville River provide 

information about trends in population abundance for the 

species in Alaskan habitats. Upon reaching age 7–8 (average 

age of maturity), the ciscoes return to Canada to spawn. 

Information about Arctic Cisco from the McKenzie River is 

extremely limited. Dolly Varden stocks spawn and overwinter 

in rivers and streams originating in the Brooks Range, Alaska 

(with perennial springs) in the eastern sector of the Alaska 

Beaufort Sea. Arctic Cisco, Least Cisco, and Broad Whitefish 

freshwater habitats are farther west in the Sagavanirktok 

River and westward of Prudhoe Bay. For the latter two 

species, habitats are located in low-lying rivers that are 

usually connected to lakes. Maintaining the connectivity and 

quality of freshwater and coastal habitats is critical for these 

species long-term sustainability. The combination of delayed 

reproduction, low reproductive effort, and increased longevity 

adapts the organism to fluctuations in recruitment and 

management concepts described for periodic strategists apply. 

Cryophilic Strategists.—The Arctic Cod and Ice Cod 

have strong ice affiliations in their life histories although 

the latter species is rarely encountered in U.S. waters. The 

Arctic Cod is a keystone species and its association with 

cold temperatures and sea ice presents a special case referred 

to herein as the cryophilic strategist. King and McFarlane 

(2003) classified the gadids with the intermediate strategists. 

Like its congeners in the south, Arctic Cod are widespread 

and abundant; however, unlike the other cods, the Arctic 

Cod is a small and short–lived species. They are the most 

important forage species in the Chukchi and Beaufort Seas 

and share many of the life traits of opportunistic strategists 

including a medium resiliency with respect to its population 

dynamics (table 6.1). The relative importance of the ice 

edge ecosystem (Alexander, 1992) in the life cycle of this 

species is ecologically different enough that we classified 

them in a unique Arctic grouping. The life cycle association 

beneath the sea surface ice may provide refuge habitat and 

special food web dynamics that can dampen the effects of 

density-independent factors on population and possibly stock 

abundance. Genetic stock information is not yet available 

and, although ongoing research suggests the presence of large 

schools over slope waters (Crawford and others, 2012), the 

species is also a coastal dominant nearer to shore. The role of 

ice in the reproductive ecology of Arctic Cod is not known.

Most of our understanding about Arctic Cod in United 

States waters is from data and information collected from 

nearshore environments. The abundance of Arctic Cod in 

coastal waters of the Alaska Beaufort Sea varies greatly from 

year to year and even between adjacent sites within a year 

(Craig and others, 1982; Palmer and Dugan 1990; Wiswar and 

Fruge 2006) although in the Canadian Arctic, large schools 

move into some discrete areas with some predictability (Welch 

and others, 1993). This often-large influx makes Arctic Cod 

by far the most abundant fish in nearshore waters (Crawford 

and Jorgenson, 1996; Fechhelm and others, 1996; Gillispie 

and others, 1997). In a recent acoustic survey, Crawford and 

others (2012) observed large schools of unconfirmed Arctic 

Cod over shelf break and slope waters of the Chukchi and 

Beaufort Seas. Estimates of summer cod abundances have 

been as high as 12–27 million fish in Simpson Lagoon, Alaska 

(Craig and Haldorson, 1981), and 900 million fish in a small 

area off Cornwallis Island in the Canadian Arctic (Crawford 

and Jorgenson, 1996). Interannual variations in patterns of 

abundance of Arctic Cod in coastal waters may be related to 

the timing, frequency, and magnitude of westerly storm events 

relative to the location of water masses and currents relative 

and spatial distributions and sizes of schooling cod over shelf 

and slope waters (for example, passive transport inshore). 

In contrast, some segment of the Arctic Cod population may 

actively migrate inshore in response to the abundance of 

invertebrate prey in coastal waters.

Although there has been considerable speculation 

regarding the environmental parameters that drive inshore 

migrations, the data often are contradictory. For instance, in 

the Chukchi Sea, Fechhelm and others (1984) determined 

that catches increased when water temperatures increased 

and salinities decreased, but the opposite was noted by 

Griffiths and others (1998) in the Sagavanirktok River 

Delta. In Prudhoe Bay, Moulton and Tarbox (1987) noted 

highest densities in frontal areas bordering low salinity 

and high temperature surface waters and high salinity 

and low temperature bottom waters, perhaps an area of 

high productivity. In another Chukchi Sea study, Gillispie 
684 Alaska Arctic Marine Fish Ecology Catalog

and others (1997) determined there was no association 

between abundance and any environmental parameter and 

they hypothesized that food availability may underlay fish 

movements. Thorsteinson (1996) reported the presence of high 

numbers of YOY Arctic Cod in surface waters of Camden 

Bay, Alaska, and hypothesized their vertical distribution and 

separation from older cod as an avoidance mechanism from 

cannibalism noted in other gadids. 

The autumn and winter behavior of Arctic Cod 

throughout the Arctic, but particularly in the U.S. Chukchi 

and western Beaufort Seas, is poorly understood. In the 

Beaufort and Chukchi Seas, at least some fish spend winters 

under nearshore ice (presumably spawning) (Lowry and 

others, 1980; Craig and Haldorson 1981; Fechhelm and 

others, 1984; Schmidt and others, 1987; Craig 1989b; and 

Thorsteinson and others, 1990). However, whether the bulk 

of the population overwinters and spawns in shallow waters 

is not known (Craig and others, 1982). For instance, Craig 

and others (1982) reported spawned-out cod both near the 

coast and 175 km (109 mi) off Prudhoe Bay. Arctic Cod have 

been reported to spawn near the bottom along the ice edge 

(Ponomarenko, 1968) and underneath ice (Andriyashev, 

1954). Ponomarenko (1968) reported that in the autumn and 

winter, large, spawning-oriented migrations occurred in the 

Russian Arctic and that spawning in the Barents Sea may have 

occurred from nearshore to hundreds of kilometers off the 

coast. Thus far, the most complete study of winter behavior 

off North America was completed by Benoit and others (2008; 

2010) in Franklin Bay, eastern Beaufort Sea. They determined 

that after spawning during the early winter (perhaps over deep 

waters in the Amundsen Gulf [D. Benoit, Université Laval, 

Quebec, Canada, oral. commun., 2013]), large numbers of 

fish either migrated to, or were passively carried into, waters 

primarily deeper than 180 m in Franklin Bay. Migration out of 

these waters coincided with spring phytoplankton blooms and 

the beginning of feeding. Whether cod in the western Beaufort 

Sea or Chukchi Sea perform similar migrations is not known, 

although fish that apparently spawned in winter in Simpson 

Lagoon had departed by February (Craig and others, 1982; 

Craig and Haldorson, 1986). 

In other research in the Northeast Water Polynya 

(Greenland Sea), Fortier and others (2006) tested a hypothesis 

that the survival of Arctic Cod larvae is limited at sea ice 

cover greater than 50 percent and sea-surface temperature 

less than 0 °C. The authors described variable recruitment 

rates in a spring and summer cohort of cods. Although only a 

low percentage (12 percent) of the spring cohort survived to 

winter, their larger size was described as having evolutionary 

significance with respect to the survival and persistence of 

this cohort. The existence of multiple cohorts in Arctic Cod 

from the Chukchi and Beaufort Seas is suggested in the length 

frequency distributions of many coastal surveys.

Resiliency

Many environmental factors affect the population 

dynamics of any particular species. Stock assessments for 

Arctic marine fish have not been done and the population 

dynamics for the species listed in this report are not known. 

For the present, the qualitative evaluation of population 

resilience based on life history traits (table 6.1) suggests the 

time requirements for recovery of dominant species from a 

large-scale mortality event such as an oil spill. Quantitative 

approaches to evaluate the magnitude of population effects 

and recoveries of hypothetical oil spills on well-studied fishery 

species in the southeastern Bering Sea have considered the 

cumulative effects of natural and anthropogenic changes in 

mortality rates over the species life cycle in space and time 

(Laevastu and others, 1985). The many information gaps 

(chapter 3) in understanding of distribution and abundance 

of life history stages, population processes, and effects 

(positive and negative) of climate change, sea ice, and ocean 

acidification preclude a defensible quantitative approach. 

Long-term data and integrative science approaches are needed 

and may be an area where traditional ecological knowledge 

and Bayesian analysis could guide resilience thinking and 

science planning with respect to managing and protecting this 

component of marine ecosystem goods and services.

Marine Fishery Science in Support of 

Ecosystem-Based Management

Fishery research in the Arctic is challenging given the 

rough weather conditions, presence of sea ice, short openwater season, irregular seafloor topography, and great depths 

off the continental shelf. Despite these challenges, important 

Arctic fish studies have been accomplished since 2002. 

Notable is an earlier Arctic expedition in 2004 by the Russian 

American Long-Term Census of the Arctic, which was 

sponsored by NOAA and the Russian Academy of Sciences. 

BOEM, in cooperation with other agencies and several 

universities (including University of Alaska, University of 

Washington, University of Maryland, and University of 

Texas), has been investigating fish use and ecological process 

in the Chukchi and Beaufort Seas. The results of recent 

(2008–12) and ongoing marine fish studies (2013), when 

available, are expected to make important new contributions to 

our understanding of Arctic marine fish diversity, population 

dynamics, and community interactions. These contributions 

will be significant in their role in ushering a new era of fishery 

ecosystem-based management by the United States and its 

circum-arctic partners.
Chapter 6 685

An important step in this synthesis process was the 

identification of general research needs and their priority in 

near- and long-term science activities. In many instances these 

complement or address similar needs that were previously 

identified for systematics of Arctic fishes (for example, 

Collette and Vecchione, 1995; Mecklenburg and others, 2011); 

life history and environmental relations (for example, Reist 

and others, 2006; Mueter and others, 2009; DeGange and 

Thorsteinson, 2011; von Biela and others, 2011; Hollowed and 

others, 2013) or quantitative population ecology (for example, 

Monterio, 2002; Katsanevakis, 2006; Wilson and Orsmeth, 

2009). A Structured Information Management process 

(Bayesian analysis) involving expert opinions is recommended 

for priority setting and addressing the most relevant Arctic 

issues in an ecosystem-based structure (for example, HollandBartels and Pierce, 2011; Jay and others, 2011). Although 

a broad list of science themes for further consideration is 

provided, it should be noted that, if implemented, many of 

the specific needs identified in the species accounts would be 

addressed over the long-term.

The Chukchi and Beaufort Seas differ with respect to 

geographic setting, expanse of continental shelves, regional 

oceanography, and freshwater-marine interactions. The 

differences affect ecological processes and the patterns of 

species occurrence in shelf and slope habitats. The presence 

of sea ice is an important feature in both seas as it relates 

to environmental temperatures, light penetration, habitat 

for species, and more than 100 marine fish, including the 

Pacific salmon that are known from the region. Life history 

information is lacking for most, is best for adult stages, and is 

best known for species occurring nearest to the shore. Major 

limitations of existing information relate to the absence of 

large-scale fisheries and lack of related resource assessment 

surveys. This lack is beginning to be addressed in light of 

changing Arctic conditions, but needs much greater scientific 

attention to abundance patterns and dispersal processes, 

population dynamics, physiological requirements, and 

community relationships.

A commitment to long-term data collection within 

an integrated science framework is needed to develop 

quantitative population understanding (similar to current 

fisheries research and assessment surveys in the Bering Sea). 

Logistical, technological, and cost considerations have limited 

the practicality of early spring and winter surveys. As a result, 

under-ice resource information is inadequate for evaluation 

of effects such as those that might be related to an oil spill in 

winter months.

A combination of laboratory, field, and modeling 

approaches is needed to estimate the effects of climate effects 

and ocean variability on production cycles and the distribution, 

abundance, and movement behaviors of Arctic marine fishes. 

As appropriate, these approaches should incorporate local and 

traditional ecological knowledge. Research and monitoring 

should focus on key species in Chukchi and Beaufort Seas fish 

assemblages in strategic locations (for example, Distributed 

Biological Observatories (long-term monitoring sites and 

biological hotspots) and include studies of human interactions. 

Human interactions extend beyond subsistence activities and 

may include changes associated with increases in marine 

transportation and OCS oil and gas activities on important 

biological habitats and ecosystems. 

Crosscutting technology-analytical themes for integrated 

research are needed to make best use of historical and 

new data collection. Modern geospatial tools are needed 

to effectively and efficiently investigate distribution and 

abundance patterns of marine fishes in time and space. Greater 

reliance on modern scientific technologies and their fishery 

applications, such as gliders, remote sensing, telemetry, 

genetics, cellular and molecular biology, and quantitative 

ecology (for example, predictive models) is needed to 

establish species environmental relationships, address existing 

gaps about relative importance of habitats, understand 

natural variation in fluctuating stocks, and accurately assess 

anthropogenic effects.

Chukchi and Beaufort Seas Marine 

Ecosystem Studies

The Interagency Arctic Research Policy Commission 

(IARPC) published a conceptual model for the U.S. Arctic 

marine ecosystem as part of its planning for integrated, 

process-oriented research in the Chukchi and Beaufort Seas 

(Wiese and others, 2013). An IARPC team is using this 

framework to develop priority needs for future ecosystem 

research. The team has highlighted the importance of winds, 

currents, and advection on nutrient dynamics and consequent 

plankton distribution and production (Wiese and others, 2015). 

Additionally, the effects of changes in sea ice and other drivers 

in the ecosystem on energy pathways (benthic and pelagic 

systems), ecosystem structure and function, and the phenology 

and location of key elements of the food web (hot-spots, hottimes, biodiversity), including access for subsistence activities 

are seen as research priorities. In coastal waters, nearshore 

changes caused by ice, winds, currents, and freshwater runoff 

and their implications for biota and communities, especially 

changes in the habitats of fish, seabirds, and marine mammals 

and subsequent implications for subsistence use and culture 

are included in the planning process. The role of humans 

within the marine ecosystem as predators, as a source of 

perturbation, and as receivers of ecosystem services, will be an 

objective of future Arctic marine ecosystem research.
686 Alaska Arctic Marine Fish Ecology Catalog

In Arctic Alaska important subsistence fisheries are 

located in seas projected to experience rapid transitions in 

temperature, pH, and other chemical parameters caused 

by global change, especially ocean acidification (Mathis 

and Questel, 2013). Many of the marine organisms that are 

most intensely affected by OA contribute substantially to 

Alaska’s local traditional economies and subsistence way 

of life. Management concerns about OA effects on marine 

organisms and ecosystems relate to food web and community 

interactions, are interdisciplinary in nature, and are far 

reaching with respect to their potential consequences: reduced 

calcification rates; significant shifts in key nutrient and 

trace element speciation; shifts in phytoplankton diversity; 

reduced growth, production and life span of adults, juveniles, 

and larvae; reduced tolerance to other environmental 

fluctuations; changes to fitness and survival; changes to 

species biogeography; changes to key biogeochemical 

cycles; changes to food webs; reduced sound absorption; 

reduced homing ability; reduced recruitment and settlement; 

changes to ecosystem goods and services; and changes to 

behavior responses. The rate and extent of change in pH in 

the northeastern Chukchi Sea suggests the importance of 

the region as a bellwether for other coastal seas (Mathis and 

Questel, 2013).

Information Gaps: Priority Marine Fishes

Twenty-one or about 20 percent of marine fish species 

are identified as high priority [A] in chapter 3 (table 6.2). 

These species are important in Arctic food webs and human 

economies or are of potential commercial interest or indicator 

status in long-term monitoring. The species comprise a mix of 

marine ecosystem goods (7 being of subsistence importance) 

and services (8 having food web values). Six species may 

support viable commercial fisheries someday. The most 

conspicuous members of the nearshore fish assemblage 

(charr, ciscoes, and whitefish) are the best known of the Arctic 

fish fauna, but fishery information is dated and population 

understanding is non-existent. Additionally, Pacific salmon 

already are becoming important in recreational fisheries 

and in some villages (for example, Nuiqsut) the increasing 

abundance and potential interaction with traditional food 

species is of concern. Most species identified are common 

to both seas (13 species) or to the Chukchi Sea only. As new 

data become available in BOEM studies, it is possible that the 

true relative abundance of species such as Greenland Halibut 

especially on shelf and slope habitats across the Beaufort Sea 

will become better known. Its potential as a marine dominant 

in these habitats is an intriguing gap in our understanding of 

this ecosystem.

Walleye Pollock has the greatest potential to be a target 

of large-scale industrial fisheries in the Chukchi Sea. In 

chapters 4 and 5, potential changing ecosystem conditions 

and distributional shifts are discussed in light of warming 

effects. Currently, cold water temperatures in the Chukchi 

and Beaufort Seas are limiting to Pollock and it has been 

hypothesized that the persistence of low temperatures in the 

northern Bering Sea will for the foreseeable future, inhibit 

large-scale shifts in fishable biomass to the north. If, in the 

future, warming favors a northward shift, this species effects 

on the Arctic marine fish assemblage would be significant. 

The effects on other congeners, especially Arctic and Saffron 

cods, is hypothesized here to result in a rearrangement of 

dominance structure through competition and predation in a 

warmer, pelagic ecosystem through changes in top-down and 

bottom-up processes, respectively.

Arctic Cod, Capelin, and Dolly Varden have circumpolar 

distributions and for reasons related to ecological significance 

and food security make them of interest in monitoring 

programs like the Arctic Council’s Arctic Monitoring and 

Assessment Programme. The colonizing potential of many 

other priority species from the Bering Sea lends similar 

support to the candidacy as indicators of climate change. 

Such selections must be based on a fuller suite of ecological 

indicators in an interdisciplinary monitoring design. That 

design should include pelagic and benthic marine ecosystem 

components, nearshore and offshore components of the survey 

region, and species whose values are representative of local 

concerns, information needed by resource managers, and 

species such as Arctic Cod, that are key to ecosystem function. 

The 21 species and collective fishery component of the Arctic 

marine ecosystem identified herein, would strategically 

contribute to the matrix of habitats, food webs, life history 

adaptations, Arctic and other zoogeographic patterns, and 

human uses that should be considered in a comprehensive 

experimental design. The composition of species is varied 

enough that processes important to the distribution and 

abundance of all marine fish and higher level consumers 

will be highlighted and strengthened by their inclusion in an 

integrated approach.

The information gaps identified for each of these species 

are similar to those of all other species. The needs were 

categorized in the species accounts (chapter 3) into eight 

life history, habitat, population, and ecological areas that 

are of importance for baseline development, environmental 

assessment, and fisheries management. The priority species 

identified inhabit all of the marine habitats and represent the 

major adaptive strategies discussed in this report (table 6.2). 

The significance of these features in future research and 

monitoring is, that lower priority species ([B] in chapter 3) 

may not be emphasized in field collections; they will be 

sampled and important information about them will more 

slowly accrue. In this way, new information about their life 

histories and ecological roles will emerge.
Chapter 6 687

Table 6.2. Characteristics of high priority [A] species in the Chukchi and Beaufort Seas.

[Priority determinations and high-priority species are described in chapters 2 and 3, respectively. Status: Fisheries reflects commercial potential as 

hypothesized by colonization potential, or for Pacific Salmon, existing fisheries in the southeastern Chukchi Sea, as well as potential expansions. For Pacific 

Herring, there is longstanding interest in Port Clarence, Kotzebue Sound, Alaska. Abundance: X, indicates “common” occurrence in Chukchi and Beaufort 

Seas. If common in one sea but not the other, the sea is indicated. Pollock is “uncommon,” but listed for the Chukchi Sea because of its colonizing potential. 

Adaptive strategy: Incorporates information about major habitats in life-cycle context]

Common name

Species characteristics

Status Abundance Adaptive

strategy

Food web 

position

Habitat 

orientation

Pacific Herring Fisheries Chukchi Linked Marine-Estuarine Intermediate Pelagic

Capelin Ecological X Linked Marine-Estuarine Intermediate Pelagic

Arctic Smelt Ecological X Nearshore Marine High Pelagic

Arctic Cisco Subsistence Beaufort Amphidromy Intermediate Pelagic

Broad Whitefish Subsistence X Amphidromy Intermediate Pelagic

Humpback Whitefish Subsistence X Amphidromy Intermediate Pelagic

Least Cisco Subsistence X Amphidromy Intermediate Pelagic

Pink Salmon Fisheries Chukchi Anadromy High Pelagic

Chum Salmon Fisheries Chukchi Anadromy Intermediate Pelagic 

Dolly Varden Subsistence X Amphidromy High Pelagic

Inconnu Subsistence Chukchi Amphidromy High Pelagic

Arctic Cod Ecological X Marine Intermediate Demersal

Saffron Cod Subsistence X Nearshore Marine Intermediate Demersal

Walleye Pollock Fisheries Chukchi Marine High Demersal

Arctic Staghorn Sculpin Ecological X Marine Intermediate Benthic

Fourhorn Sculpin Ecological X Marine Intermediate Benthic

Arctic Sand Lance Ecological X Marine Intermediate Pelagic-Demersal

Bering Flounder Fisheries X Marine Intermediate Benthic

Yellowfin Sole Fisheries Chukchi Marine Intermediate Benthic

Arctic Flounder Ecological X Nearshore Marine Intermediate Benthic

Greenland Halibut Ecological Uncommon Marine High Benthic

Fishery Objectives for Ecosystem-Based 

Management

It is within an ecosystem context and interdisciplinary 

science approach that the long-term and most outstanding 

fishery research and resource assessment needs for the 

Chukchi and Beaufort Seas will be addressed most 

appropriately. This is particularly true because, with respect 

to continued offshore oil and gas development in the Chukchi 

and Beaufort Sea Planning Areas, the science needs for 

NEPA requirements, oil-spill damage assessment, ecological 

restoration, and assessment of climate change effects, are not 

the same (Holland-Bartels and Pierce, 2011). Thus, within 

the context of current national policy for the United States 

Chukchi and Beaufort Seas (for example, White House, 2013) 

and planning for interagency marine ecosystem framework 

(for example, North Pacific Research Board [Chukchi Sea], 

Bureau of Ocean Energy Management [Beaufort Sea], and 

Interagency Arctic Research Policy Committee [IARPC]

[Chukchi and Beaufort Seas]), five science areas are identified 

having fishery objectives for possible address:

• Evaluate biological responses of populations and 

communities to natural and anthropogenic stressors 

with improved seasonal and geographic information on 

the distribution and abundance, life histories, habitats, 

community structure, and demographics for Arctic 

marine fishes.

• Determine how variability in environmental conditions 

(for example, temperature, salinity, light penetration, 

pH, water masses, and currents) influences ecological 

processes (advection of nutrients, zooplankton prey, 

and early life stages; recruitment; competition; feeding; 

reproduction; population growth and survival) and the 

abundance, and distribution of fish species, including 

the potential for Bering Sea species to move into high 

Arctic waters. 

• Describe onshore-offshore linkages (physical and 

biological) for key species in a life history context 

focusing on seasonal habitats, food webs, and 

biological interactions.
688 Alaska Arctic Marine Fish Ecology Catalog

• Determine physiological requirements of Arctic fish 

condition and health with emphasis on (1) effects 

of environmental factors responsible for changes 

in demographic rates, (2) environmental tolerances 

and preferences, and (3) effects of hydrocarbon 

contaminants and dispersants.

• Investigate the diversity and biogeography of Arctic 

marine fish through improved understanding of 

systematic, taxonomic, and phylogenetic relationships. 

Fishery Research Priorities for United 

States Chukchi and Beaufort Seas 

These broad fishery objectives are interconnected and 

would best be addressed in an integrated science approach 

as envisioned by the IARPC. Regular synthesis efforts are 

part of the planned process and, with the amount of ongoing 

BOEM research on Alaska Arctic fishes nearing completion, 

such activity should be supported. Within the objectives 

described in section, “Fishery Objectives for EcosystemBased Management,” several recommendations that stepdown for priority consideration are offered here. These 

recommendations specifically relate to the major information 

gaps identified in the marine fish species accounts and 

synthesis goals of this study. 

Marine Fish Systematics, Taxonomy, and 

Phylogenetics

Systematists study the diverse forms of life and 

determine the evolutionary relations among them. There is 

a continuing need to support the collection, analysis, and 

archive of specimen vouchers in fisheries investigations. 

Concurrently, continued, or expanded, museum studies 

are needed to resolve systematic problems and update fish 

diversity evaluations. New fishery surveys collect large 

amounts of materials; voucher specimens should be retained 

to confirm field identifications. Field data and existing 

historical records should continue to be evaluated in light of 

new taxonomic and phylogenetic understanding to update 

regional resource inventories. Reliable comparisons of fauna 

across international boundaries are difficult due to widespread 

problems with taxonomic identifications, nomenclatural 

issues, and lack of attention to standard systematic 

conventions. For the harmonization and congruency goals 

of the Arctic Council to be effectively realized, considerable 

attention to the evaluation of historical ichthyological data 

(through education and coordination) by many countries 

(for example, Russia) will be needed for effective marine 

conservation across the Arctic Basin. Finally, it has been more 

than a decade since the first edition of the Fishes of Alaska

(Mecklenburg and others, 2002) was published. There have 

been many scientific advances in taxonomic knowledge and 

understanding since 2002. The recently published baseline 

assessment (Mecklenburg and Steinke, 2015) and atlas and 

guide to Pacific Arctic Marine Fishes (Mecklenburg and 

others, 2016) fill the need for an update of the taxonomy, 

diagnostic characteristics, geographic distribution, and basic 

habitat of the marine fishes of the Alaskan Arctic region. The 

same authors, with additional collaborators from around the 

Arctic region and with funding primarily from the Norwegian 

Ministry of Foreign Affairs, are building on the Pacific Arctic 

atlas and guide to produce a reference covering the entire 

Arctic region which also includes summary information on 

life history and diet. The primary objective of the Pacific 

Arctic and pan-Arctic works is to provide baseline references 

for identifying marine fish species of the Arctic region and 

evaluating changes in diversity and distribution. A critical 

component of the ongoing research is completion of a panArctic DNA barcode reference library. These works are critical 

to help prevent errors and inform future research.

Enhance Species Accounts

A more complete understanding of the multi-dimensional 

temporal and spatial aspects of population maintenance 

is achievable by incorporating age-specific food habits 

information (included in the species accounts) in the life zone 

schemata developed for vertical distribution. This application 

would allow a novel visualization of life history stage, habitat, 

and trophic linkages and would especially be valuable for 

environmental assessments including those associated with 

National Oceanic and Atmospheric Administration needs for 

evaluating Essential Fish Habitat (for example, Rosenberg 

and others, 2000; National Oceanic and Atmospheric 

Administration, 2013). 

Advanced Geospatial Technology for 

Biodiversity Assessments

Modern geospatial tools are urgently needed to most 

effectively and efficiently investigate distribution and 

abundance patterns of marine fishes in time and space. The 

application of existing fishery data within a Geographic 

Information System environment would allow users 

to (1) explore fishery environmental relationships, and 

(2) determine population responses to changing ocean at 

multiple scales of resolution. Advanced geospatial analysis 

tools are needed for NEPA and climate change assessments, 

marine spatial planning, fisheries management applications, 

and emergency response in the unforeseen event of an Arctic 

oil spill. Research applications of modern technologies would 

focus attention on (1) geographic coverage of sampling, and 

(2) reporting of data (standards and automations) such that 

more dynamic, quantitative geospatial analyses are possible.
Chapter 6 689

Advance Scientific Technologies in Fisheries 

Applications

Greater reliance on modern scientific technologies and 

their applications, such as remote sensing, telemetry, genetics, 

cellular and molecular biology, and quantitative ecology (for 

example, predictive models) is needed to establish species 

environmental relationships, address existing gaps about 

relative importance of habitats, understand natural variation 

in fluctuating stocks, and to more accurately assess effects of 

proposed offshore oil and gas activities. As an example, the 

validation and use of environmental DNA (eDNA) approaches 

to rapid biodiversity assessments should be explored for 

through-the-ice sampling applications.

Life History and Ecological Traits

Information about status and trends, habitat requirements, 

relative distribution and abundance, and knowledge of life 

history stages of marine fish is incomplete and unavailable 

for large expanses of Arctic nearshore, shelf, and slope waters 

and should be developed for indicator species (that is, species 

that are broadly distributed, of subsistence or ecological 

significance, readily available for vulnerability assessments, 

and deemed sensitive to offshore oil and gas development 

and climate changes, see, for example, Parrish and others, 

2003; Roessig and others, 2004; and Logerwell and others, 

2015). Onshore-offshore linkages (physical and biological) 

associated with life history requirements (for example, 

seasonal movements and migrations and ontogenetic shifts in 

prey preference) have not been described. Many well studied 

marine fishes show shifts in diet over time with increasing size 

moving from low to higher trophic levels (and from smaller to 

larger prey sizes).

Quantitative Ecology

A commitment to long-term data collections is needed to 

develop population understanding. The new research that is 

currently underway will improve scientific understanding of 

abundance patterns, habitat relations, life history parameters, 

trophic relations and bioenergetics (including predators, for 

example, Brown and others, 2002), and genetic diversity and 

structure, for some species. Logistical, technological, and cost 

considerations have limited the practicality of early spring and 

winter surveys. As a result, under-ice resource information is 

limited and inadequate for evaluation of effects such as those 

that might be related to an oil spill in winter months. As of 

2016, Arctic fieldwork is expected to continue in late summer 

sampling periods and significant information gaps will remain 

with respect to spatial and temporal coverage and life stage 

coverage. The data requirements for estimating population 

parameters are substantial and an initial focus on fewer 

indicator species (common species in Chukchi and Beaufort 

Seas) may be a useful short-term approach to understanding 

change in representative marine habitats. The inclusion of 

bioenergetics components in coupled population models 

should be explored as part of efforts to evaluate population 

effects of climate change.

Empirical Data in Support of Field Work and 

Population Models

Laboratory studies are needed to investigate the causal 

mechanisms responsible for shifts in distribution or changes 

in population rates (growth, recruitment, and survival) and 

to determine the environmental preferences (for example, 

temperature and temperature-salinity relationships) of 

key Arctic species. Inferences drawn from correlations 

established in the field do not directly address mechanisms 

and their effects with respect to single or multiple stressors. 

Mueter and others (2009, p. 108) recognized the need for 

an analysis of biodiversity shifts of marine fishes in the 

northern Bering Sea: “Although biological responses to past 

temperature changes provide some basis for predicting future 

changes, such predictions are fraught with danger because 

extrapolating observed relationships beyond the historical 

range of temperatures cannot account for potential thresholds 

or non-linearities. To be able to predict where the fish is going 

one has to gather biological and ecological field data and 

the results of experiments that provide an estimate of the fish 

environmental preferences.” 

Laboratory experiments are needed to understand the 

effects of variable environmental conditions on physiological 

processes and animal health. Processes of special concern 

include feeding and digestion, assimilation and growth, 

fish behavior (responses to stimuli such as orientation and 

swimming speed), and reproduction. Collectively, these 

processes are integral to an overall assessment of fish 

condition and health. They are dependent on key water 

properties, including temperature, salinity, light penetration, 

and oxygen concentration. Animal health also is affected by 

the presence of toxic substances, infectious pathogens, and 

parasites. Experiments are needed to examine the effects 

of ocean acidification on the development, behaviors, 

and productivity of key species, their food supplies, and 

natural predators.

The relation between field (including genetics) 

and laboratory data are critical to improved population 

dynamics and modeling. How marine fish populations (and 

metapopulations) respond to changing ocean conditions, 

which are variable in space and time, creates spatial 

distributions and abundance patterns (Mueter and others, 

2013). Fish movements in these systems impose gradients 

in growth and survival through the effects of temperature, 

food concentration, sensory capabilities, predator density, 

and detection risk (Monterio, 2002; Carey and others, 2012). 
690 Alaska Arctic Marine Fish Ecology Catalog

Many marine fish stocks undertake seasonal horizontal 

migrations and the extent of these varies with age, size, and 

environmental conditions. Although field studies may reveal 

patterns at a given time and place, laboratory investigations 

may isolate causal effects. Using field and laboratory results, 

quantitative models can be used to study the effects of 

multiple environmental influences on population dynamics in 

continuous space and time (Fordham and others, 2013).

A combination of laboratory, field, and modeling 

approaches is needed to explore the potential effects of ocean 

variability on production cycles and the distribution behavior, 

movement, and abundance of Arctic marine fishes. Research 

and monitoring should focus on selected fishery resources in 

strategic locations to include effects of human interactions, 

which extend beyond subsistence use and may include 

effects associated with increased tankering, vessel support, 

and offshore construction activities on important biological 

habitats and ecosystems. The effects of invasive species also 

are an area of concern.

Participate in Regional Research and 

Monitoring Networks

Reference sites in biological hotspots should be estimated 

to support and contribute to existing long-term research and 

monitoring of coastal and marine ecosystems, including 

human interactions. Potential sites and ecological topics 

include: Bering Strait (marine ecosystem processes and fish 

distribution); Kasegaluk, Simpson, and Beaufort Lagoons 

(population dynamics of nearshore fish assemblages); Barrow 

Canyon-Hannah Shoal (benthic productivity and marine 

fish interactions); Capes Lisburne and Thompson (seabird 

colony and fishery oceanography dynamics); Point Barrow 

(dynamics of this transitional biogeographic zone); Boulder 

Patch (kelp bottom ecosystem processes); Stefansson SoundCamden Bay (Arctic Cod ecology); Mackenzie, Colville, and 

Canning River deltas (onshore-offshore linkages); ice edge 

and polynyas (biological significance to marine fish and higher 

level consumers). Local residents are often the first to notice 

changes in fish and wildlife populations. Mechanisms should 

be developed to better solicit and integrate local and traditional 

ecological knowledge as a basic source of information.

Participate in Additional Investigations of 

Iñupiaq Taxonomy

Research in Kotzebue Sound (Georgette and Shiedt, 

2005) demonstrated the complexities and subtleties of the 

Iñupiag classification system. Additional investigations are 

needed in concert with subsistence resource surveys to fully 

incorporate traditional ecological knowledge into biological 

research on fishery populations and their habitats.

Summary

In many respects, present day understanding of Arctic 

Alaska’s marine fishes is similar to what was known about 

the Bering Sea in the 1970s. Quantitative data are lacking 

or dated, but new information is slowly developing. The 

relationships between basic life history characteristics and 

population dynamics have implications for managing marine 

fish resources where population data are limited or nonexistent. These relationships previously were used to describe 

strategic groups in the Northeast Pacific as an aid to fishery 

management. Taking a qualitative approach, we applied the 

criterion for these groupings to the Arctic marine fish fauna 

and added two new groups. One group, the amphidromic 

strategists, includes those Arctic fishes displaying an 

amphidromous life strategy. Arctic Cod, unlike other gadids, 

are similar to the opportunistic strategists, but because of their 

dependence on sea ice habitats, life history, and central role in 

Arctic marine ecosystems, they are considered independently 

as a cryophilic strategist. Long-term fishery research 

objectives for the Chukchi and Beaufort Seas are described 

in relation to information gaps and they are addressed in 

planned marine ecosystem research in the U.S. Arctic. Some 

suggestions for more immediately needed studies relate to 

access of existing information, description of environmental 

preferences for key marine species, participation in regional 

monitoring networks and cooperative research, and continued 

biodiversity assessments through field, laboratory, and 

museum studies. 
1

The von Bertalanffy Growth Model is also expressed as the von 

Bertalanffy Growth Function (VBGF) and von Bertalanffy Growth Equation 

in the scientific literature.

Chapter 7. Glossary of Ecological Terms

Information Sources

The terminology used in chapters 1 through 6 is 

common in fishery science and quantitative ecology. The list 

of science terms identified in this glossary focuses not only 

on the common vocabulary used but also the uncommon 

and specialized terminology used in this report and its 

appendices. Each is accompanied by a definition that best 

reflects the meaning conveyed by the authors. A number of 

online resources and scientific references were consulted 

in preparation of the glossary. In many instances, the terms 

relate to fish population dynamics and quantitative concepts 

and, in these cases, the primary scientific references have 

been cited. The web sites used to develop definitions for more 

descriptive biological terminology include (1) A Glossary of 

Ecological Terms (http://www.terrapsych.com/ecology.html); 

(2) The Nature Education Knowledge Project (http://www.

nature.com/scitable/knowledge); (3) Encyclopedia Britannica 

(https://www.britannica.com/); (4) FishBase (http://www.

fishbase.org/); (5) Merriam-Webster Online (http://www.

merriam-webster.com/); and (6) The Free Dictionary (http://

www.thefreedictionary.com/). An extensive glossary of fishery 

science terms is presented in Mecklenburg and others (2002). 

Where appropriate, the definitions are presented with special 

attention to their application to meaning in the Chukchi and 

Beaufort Seas.

Science Terms 

Abundance Generally, there are three types of abundance 

measures: total number of animals in a population (absolute 

abundance), number of animals per unit area (density), and 

abundance and density of one population to another (relative 

abundance). Because temporal variation in a population 

is often described in density and biomass parameters, the 

distinction between biomass and other abundance measures is 

important. Biomass, expressed as mass per unit area, or mass 

per volume, reflects the quantity of resources incorporated 

by the population (that is, bioenergetics) and as such, is an 

indicator of its relative role in the ecosystem (Odum, 1985). 

Age The length of time during which a being or thing 

has existed; the number of years of life completed (fishery 

statistics). Age estimation is a fundamental part of studies 

of the life history of fish and a key area in the monitoring, 

assessment, and management of fish stocks because age 

determined parameters underlie the population-dynamics 

models used for fish stock assessments. The age-determined 

parameters include age profiles and age at first maturity, and, 

when considered with length and weight measurements, 

provide valuable information about stock composition, 

age at maturity, life span, mortality, and stock production 

(Ricker, 1975). 

Age at first maturity (t

m) Mean or median age at first maturity 

(that is, age at which 50 percent of a cohort spawn for the first 

time). Values are estimated from population parameters in the 

von Bertalanffy Growth Model (VBGM)1 (Ricker, 1975). 

Age and growth Basic information about the age and 

growth of a species and the environmental area inhabited by 

that species is foundational to effective fishery management. 

Habitat is the natural environment that influences and is 

used by a species population. The relations between length 

and weight and size and age form the basis for population 

dynamics (that is, population growth and production 

models) and quantitative analysis of environmental effects. 

Importantly, the relation between total length (L) and total 

weight (W) for nearly all species of fish is expressed by the 

equation (Ricker, 1975): 

W = aLb or ln W = ln a + b ln L (1)

where 

W is weight, in kilograms;

L is length, in centimeters; 

a is y-intercept; and 

b is slope.
692 Alaska Arctic Marine Fish Ecology Catalog

A well-known growth model is the VBGM, which is based 

on a bioenergetics expression of fish growth converts lengthfrequency data into age composition (see, for example, 

Essington and others, 2001). The model expresses length (L) 

as a function of the age of the fish (t):

( ) ( )

( ) ( )

0 (–

0

) 1 or 

* 1 *

kt t L L e Lt t

L exp K t t

−

∞

∞

= −

= −−−    

(2)

where

Lt is the length at time t; t

0

 the hypothetical age 

of the fish at zero length; 

K is the growth coefficient, the rate at which 

L1 approached expressed as the rate (per 

year) at which L∞ is approached (also see 

“Carrying capacity”); and

L∞ is the asymptotic length, the mean length 

a given stock would reach if it were to 

growth indefinitely.

Estimation of growth parameters (K, L∞ , t0) of the VBGM are 

good indicators of stock condition and health. The right hand 

side of the equation contains the age, t, and some parameters. 

They are: “L∞” (read “L-infinity”), “K” and “t

0

” (read “t-zero”). 

Different growth curves will be created for each different set 

of parameters; therefore, it is possible to use the same basic 

model to describe the growth of different species simply by 

using a special set of parameters for each species. 

Age structure The distribution of ages, or classes of ages, of 

individuals of a population. The presence of separate cohorts 

in fish populations may reflect adaptations in life strategies 

to cope against year to year variability in environmental 

conditions (Ricker, 1975). 

Ammocoetes The larval stage of the primitive lamprey, 

known as an ammocoetes larva. Resembling a small eel, the 

larval lamprey can remain concealed in the sediments of rivers 

and estuaries from 7 to 17 years. Burrows are detectable as 

funnel shaped depressions, and in shallow, clear, flowing 

springs and streams, the ammocoetes larva will often expose 

their heads orienting their buccal cavities and gills into 

the current. Throughout the larval stage, there is a passive 

downstream migration of ammocoetes leading to maturity, and 

in some species, parasitism. 

Amphidromy (of a migratory fish) that travels between 

freshwater and salt water, but does not travel to breed. 

Anadromy (of a migratory fish) that lives in the sea and 

breeds in fresh water. 

Arctic realm One of 12 marine realms as designated by 

the World Wildlife Federation and The Nature Conservancy. 

It includes the coastal regions and continental shelves of 

the Arctic Ocean and adjacent seas, including the Arctic 

Archipelago, Hudson Bay, and the Labrador Sea of northern 

Canada, the seas surrounding Greenland, the northern and 

eastern coasts of Iceland, and the eastern Bering Sea. 

Axon An axon (from Greek, axis) also known as a nerve 

fiber, is a long, slender projection of a nerve cell, or neuron, 

that typically conducts electrical impulses away from the 

neuron’s cell body. The function of the axon is to transmit 

information to different neurons, muscles, and glands. 

Axoplasm Axoplasm is the cytoplasm within the axon of a 

neuron (nerve cell). Neural processes (axons and dendrites) 

contain about 99.6 percent of the cell’s cytoplasm, and 

99.7 percent of that is in the axons. 

Baseline Initial collection of data, which serves as a basis 

for comparison with the subsequently acquired data. 

Behavior Traditional fishing methods, scientific surveys, 

population modeling, and fishery management strategies 

are dependent upon a fundamental understanding of fish 

behavior. Behavioral research develops information about 

the relationships between fish behavior and environmental 

variables (physical and biological), and how this influences 

distribution, survival, and recruitment processes. Species 

have evolved particular behaviors to cope with environmental 

temperature and light, foods and predators, and reproduction.

Biodiversity Biological variety of the kind that preserves 

species and their DNA. Other definitions are (1) the number of 

species in an ecosystem, (2) the diversity between ecosystems, 

and (3) the diversity of entire regions. Depleted biodiversity 

leads to population crashes, declines in genetic variability, 

and extinctions. 

Biogeography Biogeography is the study of the distribution 

of species and ecosystems in geographic space and through 

geological time. Organisms and biological communities vary 

in a highly regular fashion along geographic gradients of 

latitude, elevation, isolation, and habitat area. 

Biogeographic province Provinces, as defined in 

biogeographic classification schemes, are nested within 

the realms. 

Provinces are large areas defined by the presence 

of distinct biotas that have at least some cohesion 

over evolutionary time frames. Provinces will hold 

some level of endemism, principally at the level 

of species. Although historical isolation will play 

a role, many of these distinct biotas have arisen 

as a result of distinctive abiotic features that 

circumscribe their boundaries. These may include 

geomorphological features (isolated island and 

shelf systems, semienclosed seas); hydrographic 

features (currents, upwellings, ice dynamics); or 

geochemical influences (broadest-scale elements of 

nutrient supply and salinity) (Spalding and others, 

2007, p. 575). 

The BOEM Arctic Planning Areas are nested within the Arctic 

realm and the boundaries that include Chukchi and Beaufort 

Sea provinces. The Beaufort Sea province includes continental 

coast and shelf waters of the United States and Canada. 
Chapter 7 693

Brackish In the Arctic, brackish typically refers to salinity 

conditions of 10–25 practical salinity units (psu). Warm 

(5–10 °C) brackish waters have estuarine-like qualities 

(Craig, 1989a). 

Carrying capacity (K) In nature, population growth must 

eventually slow, and population size ceases to increase. As 

resources are depleted, population growth rate slows and 

eventually stops, known as logistic growth. The population 

size at which growth stops is generally called the carrying 

capacity (K), which is the number of individuals of a particular 

population that the environment can support. At carrying 

capacity, because population size is approximately constant, 

birthrates must equal death rates, and population growth is 

zero. Growth rate K (1 – L (t)/L∞) decreases with time as the 

length of the fish reaches asymptotic length (Ricker, 1975). 

Chukchi borderlands The gateway to the Arctic Ocean, 

located 600 miles north of the Bering Strait and 800 miles 

south of the North Pole. From Jakobsson and others (2008, 

p. 527): 

The Chukchi Borderland is comprised of a group 

of generally less than 1000 m deep, north-trending 

ridges that surround the extensional Northwind 

Basin (Figure 2) (Hall, 1990). The easternmost of 

these ridges is the Northwind Ridge, which is deeper 

than its western neighbours and is characterized by 

an exceptionally steep slope towards the Canada 

Basin and a gently rounded to flat topped ridge 

crest. The Chukchi Spur–Chukchi Plateau composite 

ridge lies on the western side of the Northwind 

Basin and has a wide (>140 km at 78° N) flat topped 

crest mainly shallower than 600 m (Figure 2(a)). In 

addition to the large Northwind Ridge and Chukchi 

composite ridge, several much smaller ridges rise 

above the floor of the Northwind Basin.

Location of existing fish sampling and related catch 

characteristics is available in Mecklenburg and others (2014).

Colonization The spreading of a species into a new habitat. 

Successful colonization, as used in this report, implies the 

success of a reproductive event in the newly occupied habitat. 

Compensation (density dependent processes) Population 

processes such as survival, growth, reproduction, and 

movement are considered density dependent if their rates 

change as a function of population abundance (Ricker, 1975). 

Processes that limit population growth at high abundances 

(for example, slower growth, increased emigration, and 

lower survival) or increase numerical growth at low 

population abundances (for example, faster growth, increased 

immigration, and higher survival) are examples of direct 

density dependence or compensation. Direct density dependent 

processes operate as a negative feedback and tend to stabilize 

population abundance. 

Crepuscular feeder Crepuscular feeding occurs primarily 

at twilight (that is, during dawn and dusk). This is opposed 

to diurnal and nocturnal feeding behaviors, which occur 

respectively during hours of daylight and dark. 

Dendrite Dendrites are the branched projections of a neuron 

that act to propagate the electrochemical stimulation received 

from other neural cells to the cell body, or soma, of the neuron 

from which the dendrites project. Electrical stimulation is 

transmitted onto dendrites by upstream neurons (usually their 

axons) through synapses located at various points throughout 

the dendritic tree. Dendrites play a critical role in integrating 

these synaptic inputs and in determining the extent to which 

action potentials are produced by the neuron. 

Depensation (density independent processes) If population 

processes, such as survival and growth, decrease at low 

population abundances or increase at high abundances, they 

are referred to as inverse density dependence or depensation 

(Ricker, 1975). Inverse density dependent processes operate as 

a positive feedback and tend to destabilize populations. 

Diadromy Migration of fish in either direction, from fresh to 

seawater or the reverse. 

Ecosystem A biotic community and its surroundings, 

part inorganic (abiotic) and part organic (biotic), the latter 

including producers, consumers, and decomposers. 

Ecological niche Describes the relational position of a 

species or population in an ecosystem. More formally, the 

niche includes how a population responds to the abundance 

of its resources and enemies (for example, by growing 

when resources are abundant and predators, parasites and 

pathogens are scarce) and how it affects those same factors 

(for example, by reducing the abundance of resources through 

consumption and contributing to the population growth of 

enemies by falling prey to them). The abiotic or physical 

environment also is part of the niche because it influences how 

populations affect, and are affected by, resources and enemies. 

The description of a niche may include descriptions of the 

organism’s life history, habitat, and place in the food chain. 

Endemism The ecological state of a species being unique 

to a defined geographic location, such as an island, nation, 

country or other defined zone, or habitat type; organisms that 

are indigenous to a place are not endemic to it if they are also 

found elsewhere. 

Epipelagic Of, relating to, or constituting the part of 

the oceanic zone in which enough light penetrates for 

photosynthesis. 

Extralimital (pertaining to a species) Not found within a 

given geographical area. 

Eurythermic The animal is adaptable to a wide range of 

temperatures in the environment. 
694 Alaska Arctic Marine Fish Ecology Catalog

Food and feeding The numerical assignment of an organism 

to a trophic level has evolved to include ecological insight 

about a species position and interactions within a food web 

or ecosystem (Odum and Heald, 1975). In classic food web 

studies (Lindeman, 1942)2, five (T1-5) trophic levels were 

recognized by ecologists. Lower trophic levels classified 

as producers (T1) and herbivores (T2), and higher trophic 

levels and primary and secondary carnivores (T3 and T4, 

respectively) and apex consumers (T5). Analysis of stomach 

contents often is used to associate a species with a specific 

level, intermediate standing between levels, or multiple 

levels. Recent models of trophic interactions (for example, 

Christensen and Pauly, 1992; Pauly and Watson, 2005) include 

a food-based fractional component in trophic level analysis 

to quantify the relative importance of prey in the consumers 

diet as well as the role of primary production. Model results 

present trophic level designations as continuous numbers 

ranging from 1 to 5. Age-related variations in foods habits 

are known in marine fish and thus, in reality, trophic level 

status will change over time. In most instances, deficiencies 

in life-history information have limited existing analyses 

to trophic level determinations based on combined data 

sets for juvenile and adult life stages. Although imperfect, 

the fractional analysis is meant to improve ecosystem 

understanding and energy flows because it more realistically 

addresses the complexities of consumer feeding behaviors 

(omnivory and feeding across multiple trophic levels) and 

predator-prey relationships. To illustrate, the mean trophic 

level for Blackline Prickleback (Acantholumpenus mackayi) 

is 3.1 (±0.31). This mid food web value is indicative of a 

primary carnivore that feeds across trophic levels, in this case 

on lower level herbivores. The trophic level values reported 

herein are as reported in FishBase (Froese and Pauly, 2012). 

Generation time The average age (t

g

) of parents at the time 

their young are born. In most fishes, Lopt (derived as a growth 

parameter in the VBGM) is the size class with the maximum 

egg production. The corresponding age (t

opt) is a good 

approximation of generation time in fishes. It is calculated 

using the parameters of the VBGM as t

g = topt = t0 - ln(1 - Lopt / 

Linf) / K. Note that in small fishes (<10 cm) maturity is often 

reached at a size larger than Lopt and closer to Linf. In these 

cases, the length class where about 100 percent (instead 

of 50 percent) first reach maturity will contain the highest 

biomass of spawning fishes, usually resulting in the highest 

egg production. As an approximation for that length class, 

most fish most fish are assumed to have reached maturity at 

a length that is slightly longer than Lm, namely: Lm100 = Lm + 

(Linf - Lm) / 4, and generation time is calculated as the age at 

Lm100. This is applied whenever Lm >= Lopt. 

Gonochoristic The state (gonochorism) of having just one of 

at least two distinct sexes in any one individual organism. 

Growth The study of growth means the determination of 

the body size as a function of age (Ricker, 1975). Therefore, 

all stock assessment methods work essentially with age 

composition data. This measures the growth of individuals 

in size and length. This is important in fisheries where the 

population is often measured in terms of biomass. The VBGM 

is widely to estimate of productivity parameters. 

Specific growth rate (K) is given by:

K = (ln wt – ln wo) / t (3)

where 

wt is the final weight and w0 the initial weight, 

t is equal to the number of days considered, and

K is assumed to be constant. 

Hence, the individual growth is described by an exponential 

expression, relating the weight of the fish to time. 

Growth model A primary interest of fisheries biologists is 

to estimate rates of fish population growth and understand the 

processes and factors that influence growth. Age and growth 

information is critical for research addressing questions about 

basic ecological relations and for managing fisheries. In the 

latter case, growth information is frequently used to populate 

assessment models with vital rates and age-specific length, 

weight, fecundity, and vulnerability to exploitation. 

Additionally, information on growth may be used to 

estimate the age of fish based on size. Given the importance 

of understanding growth, much effort has been expended to 

understand factors that influence growth, to develop models 

to describe observed growth patterns, and to estimate the 

parameters of those models. 

The decision about which growth model to use is 

important because most ecological models deal with transfer 

of energy or matter along the trophic chain. Growth can be 

constant or change during development periods. Although 

initial and final weights estimated with different models may 

be similar, growth curves as well as the total amount of energy 

or matter consumed over time may vary considerably during 

the growth period. It is suggested that the Gompertz or the 

parabolic growth models seem to be more appropriate for the 

description of young fish growth. For older fish, the VBGM 

equation or some modified form adjusted to seasonal change is 

preferable (Gamito, 1998). 

Intrinsic rate of increase (r

m) The rate of increase in 

populations that reproduce within discrete time intervals and 

possess generations that do not overlap. The rm is calculated 

as number of births minus number of deaths per generation 

time (reproduction rate less the death rate). Values greater 

than zero indicate that the population is increasing, the higher 

the value, the faster the growth rate. A population that has 

intrinsic rate of increase of zero is considered to have a stable 

age distribution (neither growing nor decreasing in numbers). 2

Some scientific literature refers to trophic levels T1-5 as I-V.
Chapter 7 695

A growing population has more individuals in the lower 

age classes than does a stable population, and a decreasing 

population has more individuals in the older age classes than a 

stable population (Musick, 1999). 

Iteroporous A reproductive strategy (iteroparity) in which 

the species is characterized by multiple reproductive cycles 

over the course of a lifetime. 

K-selection Species in stable environments tend to live 

longer and produce fewer, and sometimes larger, offspring. 

(K is the constant for carrying capacity in terms of 

population growth.) 

Lateral line A system of sense organs in aquatic vertebrates, 

mainly fish, used to detect movement and vibration in 

the surrounding water. The sensory ability is achieved by 

modified epithelial cells, known as hair cells, which respond to 

displacement caused by motion and movement and transduce 

these signals into electrical impulses through excitatory 

synapses. Lateral lines serve an important role in schooling 

behavior, predation, and orientation. 

Length (L) The length-frequency of a fish population is a 

key determinant in analyses of status and trends (Ricker, 

1975). The length distributions of fish in samples give the 

simplest index of the composition of the stock from which the 

catch was taken. (Weight measurements are also required for 

determination of length-weight relationships and condition 

analysis.) Length measurements are reported as standard 

length, total length, and fork length by fishery investigators.

• Standard length (SL) refers to the length of a fish 

measured from the tip of the snout to the posterior 

end of the last vertebra or to the posterior end of the 

mid-lateral part of the hypural plate. This measurement 

excludes the length of the caudal fin.

• Total length (TL) refers to the length from the tip of 

the snout to the tip of the longer lobe of the caudal fin, 

usually measured with the lobes compressed along the 

midline. It is a straight-line measure, not measured 

over the curve of the body. 

• Fork length (FL) refers to the length from the tip of 

the snout to the end of the middle caudal fin rays and 

is used in fishes in which it is difficult to tell where the 

vertebral column ends. 

Life span (t

max) Life span is a central aspect of life history 

diversification. The parameter t

max is the approximate 

maximum age that fish of a given population would reach. It is 

calculated as the age at 95 percent of L∞ using the parameters 

of the VBGM. Life span is frequently estimated from 

maximum age and size data (Ricker, 1975). 

Life span depends on the organism’s survival schedule 

and is often associated with fecundity. The combination of 

survival and fecundity fitness components constitutes the basis 

for understanding the evolution of life histories. The theory of 

r- and K-selection expects that life histories can evolve toward 

short or long life spans because of variation in ecological 

factors such as resource availability. Resource limitations 

impose trade-offs between different fitness components (for 

example, reproductive effort compared with individual growth 

rate) that are expected to translate into different demographic 

patterns. For example, as life span increases, the importance 

of fecundity for overall population dynamics is progressively 

replaced by that of survival. 

Limiting factors All living things need food, water, shelter, 

and space to survive. As long as organisms have all of these 

things available to them their population will continue to grow. 

However, populations cannot grow forever. Some form of 

environmental resistance will stop the growth of a population 

(Ricker, 1975). The form of environmental resistance is called 

a limiting factor because it limits the population. However, 

limiting factors also may increase a population. In nature, 

population growth must eventually slow, and population size 

ceases to increase. As resources are depleted, population 

growth rate slows and eventually stops (known as logistic 

growth). The population size at which growth stops is 

generally called the carrying capacity (K), which is the number 

of individuals of a particular population that the environment 

can support. At carrying capacity, because population size is 

approximately constant, birthrates must equal death rates, and 

population growth is zero.

Limitations to population growth are either densitydependent or density-independent. Density-dependent 

factors include disease, competition, and predation. Densitydependent factors can have either a positive or a negative 

correlation to population size. With a positive relationship, 

these limiting factors increase with the size of the population 

and limit growth as population size increases. With a negative 

relationship, population growth is limited at low densities and 

becomes less limited as it grows.

• Density-dependent factors may influence the size of the 

population by changes in reproduction or survival (for 

example, food availability—effects on fecundity and 

[or] habitat condition)

• Density dependent factors also may affect population 

mortality and migration.

Factors that decrease population growth can be defined as 

environmental stress including limitations in food, predation, 

and other density-dependent factors. However, many sources 

of environmental stress affect population growth, irrespective 

of the density of the population. Density-independent 

factors, such as environmental stressors and catastrophe, are 

not influenced by population density change. Although the 

density-dependent factors are often biotic, density-independent 

factors are often abiotic. These density-independent 

factors include food or nutrient limitation, pollutants in the 

environment, and climate extremes, including seasonal cycles 

such as monsoons. Catastrophic factors such as fires and 

hurricanes also can affect population growth (Monterio, 2002).
696 Alaska Arctic Marine Fish Ecology Catalog

Some important density-independent factors include:

• The quality of nutrients (for example, food quality, and 

amount of particular plant nutrients) in an environment 

affects the ability of an organism to survive, grow, and 

reproduce. 

• Pollutants also contribute to environmental stress, 

limiting the growth rates of populations. 

• Environmental catastrophes such as oil spills, fires, 

earthquakes, volcanoes, and floods can strongly 

affect population growth rates by direct mortality and 

habitat destruction. 

Logistic population growth The geometric or exponential 

growth of all populations is eventually limited by food 

availability, competition for other resources, predation, 

disease, or some other ecological factor. If growth is limited 

by resources such as food, the exponential growth of the 

population begins to slow as competition for those resources 

increases. The growth of the population eventually slows 

nearly to zero as the population reaches the carrying capacity 

(K) for the environment. The result is an S-shaped curve of 

population growth known as the logistic curve (Ricker, 1975). 

Myomeres The blocks of skeletal muscle tissue found 

commonly in chordates. They are commonly zig-zag, W- or 

V-shaped muscle fibers. The myomeres are separated from 

adjacent myomere by connective tissues and most easily seen 

in larval fishes. Myomere counts are sometimes used for 

identifying specimens because their number corresponds to the 

number of vertebrae in the adults. 

Microphthalmia A developmental disorder of the eye that 

literally means small eye. One (unilateral microphthalmia) or 

both (bilateral microphthalmia) eyes may be involved.

Myosin structure Myosins are a family of Actin Motor 

Protein that are Adenosine triphosphate (ATP)-dependent 

motor proteins and are best known for their role in muscle 

contraction and their involvement in a wide range of other 

eukaryotic motility processes. They are responsible for actinbased motility. All myosins have head, neck, and tail domains 

with distinct functions. 

Natural mortality (M) Natural mortality is a parameter in 

most fish stock assessment models (Ricker, 1975). Natural 

mortality can occur through predation or non-predation events 

such as senescence and disease. It is generally accepted that 

the natural morality is high during larval stages and decreases 

as the age of fish increases, approaching a steady state. 

The rate then increases exponentially when the fish nears 

maximum age. Natural mortality also may vary with size, sex, 

parasite load, density, food availability, and predator numbers. 

However, in most cases, a single value—usually 0.2—for M is 

assumed for stock assessments. 

Neritic Shallow marine waters extending from mean low 

water to 200-meter depths. 

Neuromast The major unit of functionality of the lateral line 

is the neuromast. The neuromast is a mechanoreceptive organ 

that allows the sensing of mechanical changes in water. There 

are two main varieties of neuromasts in animals, superficial 

or freestanding neuromasts and canal neuromasts. Superficial 

neuromasts are located externally on the surface of the body, 

whereas canal neuromasts are located along the lateral lines 

in subdermal, fluid filled canals. Each neuromast consists of 

receptive hair cells whose tips are covered by a flexible and 

jellylike cupula. See “lateral line.” 

Otophysic A connection between the swimbladder and the 

inner ear that enhances the hearing capability in various types of 

fish. 

Oviparous Fishes producing eggs that hatch outside the body 

of the mother.

Poikilotherm An organism whose internal temperature varies 

with the temperature of its surroundings. 

Paedmorphic Of, relating to, or resulting from the retention of 

juvenile characteristics by an adult. 

Phylogenetic relationships In biology, phylogenetics is the 

study of evolutionary relationships among groups of organisms 

(for example, species and populations), which are discovered 

through molecular sequencing data and morphological data 

matrices. The result of phylogenetic studies is a hypothesis 

about the evolutionary history of taxonomic groups: their 

phylogeny. 

Population distribution Variation of population density over a 

particular geographic area. 

Population dynamics Major abiotic and biotic factors that 

tend to increase or decrease the population size, age, and sex 

composition of a species. 

Populations or stocks The subdivision of species into local 

populations and the adaptive nature of genetic differences 

between these populations are interlinked by the ecological and 

genetic processes that subdivide and determine the discreteness 

of these stocks. Genetic discreteness implies some restriction 

of gene flow and spatial and temporal mechanisms of isolation. 

Differences in genetic stock structure reflect behavioral 

processes and adaptation within the species particular life 

history strategy.

Management at the stock-level is a cornerstone of 

conservation biology. In fisheries, the stock refers to the part 

of the fish population that is under actual or potential use. 

Population dynamics describes the ways in which a given 

population grows and shrinks over time as controlled by birth, 

death, and emigration or immigration. The population levels 

of exploited marine fish stocks are regulated through many 

underlying processes. Biological and environmental conditions 

as well as exploitation rate and pattern determine the balance 

between the increase in stock size due to recruitment and growth 

and losses caused by fishing and predation mortality. Critical 

parameters for population dynamics evaluations include stock 

size, age structure, recruitment, and growth (Ricker, 1975). 
Chapter 7 697

Relative abundance The contribution a species makes to 

the total abundance of the fishery community (Miller and Lea, 

1976). It is a measure that provides an index of the number 

of individuals present, but not the actual numbers. Relative 

abundance terms, such as “abundant,” “common,” or “rare” 

often are used to express the general population status of 

a given species, but are most useful when they are defined 

by something that is measured or estimated in a manner 

that makes comparison meaningful. The definitions used in 

individual studies conducted in Chukchi Sea vary greatly and 

often reflect the differences in sampling gears, geographical 

localities and habitats sampled, time of sampling, and 

statistical approach. Thus, attempts to tie reported indices of 

abundance from catch summaries of individual studies (that is, 

species composition or catch-per-unit effort data) proved too 

challenging initially for this analysis, especially for the more 

poorly known fauna. 

Recruitment The reaching of a certain size or reproductive 

state. For fisheries, recruitment usually means addition of new 

fish to the vulnerable population by growth from smaller size 

categories (Ricker, 1975). 

Reproduction (Reproduction mode, chapter 3) The 

reproductive strategies of marine, amphidromous, and 

anadromous fishes demonstrate sharp contrasts related to 

their adaptation to cold Arctic conditions. Marine species 

generally have shorter lives, grow more quickly, and achieve 

earlier maturity than the anadromous counterparts. The low 

abundance of many species suggests tendencies for niche 

generalists, rather than specialists, animals that feed at lower 

trophic levels, and whose population may be more affected by 

density independent factors. For most Arctic species, marine 

and anadromous-like, population or stock information about 

the population characteristics (for example, sex ratios, age-atmaturity, and fecundity) that affect reproductive potential does 

not exist. Little is known about spawning times and locations 

or mating behaviors of marine species, and what is known has 

come from data collection outside the study area (chapter 3). 

Resilience In ecology, resilience traditionally refers to the 

ability of a biotic community to sustain or return to its former 

state after a disturbance. The rate of recovery is a measure of 

resilience determined by the population processes involved in 

restoring abundance to healthy, sustainable, or pre-disturbance 

levels (Musick, 1999). Resilience also is thought of as a 

measure of the amount of change or disruption that is required 

to transform a system from being maintained by one set of 

mutually reinforcing processes and structures to a different set 

of processes and structures. 

The competing definitions emphasize different aspects 

of stability. The practical application of resilience concepts 

requires substantial information about population status and 

trends (for example, intrinsic rate of population growth), 

biological interactions, genetic relationships, and effects of 

natural and anthropogenic stressors.

The American Fisheries Society proposed a method to 

classify a population’s resilience (vulnerability to extinction) 

using VBGM parameters and other life history traits 

(Musick, 1999). In the two-tiered approach a population in 

is initially assigned to one of four productivity categories. 

The second part of the vulnerability assessment examines 

recent trends in population size and further classifies the 

productivity categories according to observed rates of decline. 

This approach to studying population resilience is a first 

approximation of extinction risk. This means that a population 

that is determined to be vulnerable merits further study or 

status review.

In the first step, the five most important productivity 

parameters evaluated are (in order of importance): (1) intrinsic 

rate of growth (rm, expressed as an instantaneous or annual 

percentage), (2) growth coefficient (K), (3) fecundity (number 

of eggs per year), (4) age at maturity (t

m), and (5) maximum 

age (t

max) (table 7.1). The intrinsic rate of increase is 

considered to be most important because its calculation 

incorporates all of the other parameters. Musick (1999) and 

others note that this parameter is the most difficult to estimate 

and is usually not available. As such, when information 

on rm is available, it takes precedence in assignments to 

a productivity category. Age at maturity is the next most 

important parameter because it often is correlated with the 

growth coefficient (K) and maximum age. Musick (1999) 

noted that although fecundity also is a key indicator, it needs 

to be properly assessed in light of demographics features 

of the population, especially maximum age. Many fish 

populations exhibit a direct correlation between age or size 

and fecundity. Because of this, Musick (1999) recommended 

using fecundity-at-first-maturity to assess resilience. 

As a rule, in the absence of an estimate of rm, the fish 

population should be classified according to the lowest 

productivity category for which data are available (table 7.2). 

Table 7.1. Productivity parameters proposed to the American 

Fisheries Society for classifying resilience of marine fish 

populations.

[Adapted from Musick (1999). Life history parameter: rmax, intrinsic rate of 

increase; K, growth coefficient; t

m, age at maturity; tmax, maximum age. <, less 

than; >, greater than]

Life history

parameter

Productivity category

High Medium Low Very low

rmax

 (1/year) >0.5 0.16–0.50 0.05–0.15 <0.05

K (1/year) >0.3 0.16–0.30 0.05–0.15 <0.05

Fecundity 

(1/year)

>10,000 100–1,000 10–100 <10

t

m (years) <1 2–4 5–10 >10

t

max (years) 1–3 4–10 11–30 >30
698 Alaska Arctic Marine Fish Ecology Catalog

Table 7.2. Decline thresholds as a function of resilience/

productivity and population decline.

[Adapted from Musick (1999). Observed population decline: Threshold is 

percentage of rate of decline in numbers or biomass of mature individuals in a 

marine fish population over the longer of 10 years or 3 generations]

Decline thresholds 

Productivity category Observed population decline

High 0.99

Medium 0.95

Low 0.85

Very Low 0.70

As an example, a fish with high fecundity (>104), but late 

maturity (5–10 years), and long life span (>30 years), would 

be classified under the Very Low Productivity category. 

Generally, populations having VBGM growth coefficients (K) 

less than 0.10 and (or) intrinsic rates of increase (rmax) less than 

10 percent per year may be especially vulnerable.

Once a population’s productivity category is established, 

the second tier of the assessment considers rates of population 

decline (table 7.2). Population trends are analyzed over the 

longer of two periods: 1 decade or 3 generations. Information 

about decline (from Musick, 1999) is based on (1) population 

decline observed, estimated, inferred, or suspected in the past, 

or (2) population decline projected or suspected in the future 

based on (a) direct observation; (b) an index of abundance 

appropriate for the taxon; (c) a decrease in area of occupancy, 

extent of occurrence, and (or) quality of habitat; (d) actual or 

potential levels of exploitation; or (e) the effects of introduced 

taxa, hybridization, pathogens, pollutants, competitors, 

or parasites.

Richness Species richness is the number of different species 

represented in an ecological community, landscape, or region. 

Species richness is simply a count of species, and it does 

not take into account the abundances of the species or their 

relative abundance distributions. [3]

R-Selection Rapid growth and occupation through 

early reproduction, short life spans, low biomass, and 

many offspring. 

Salinity The concentration of salts dissolved in water, 

measured in parts per thousand (‰) or practical salinity units 

(psu). Seawater averages 34 psu. Arctic salinities typically 

range from 27 to 32 psu. Ocean salinity is generally defined 

as the salt concentration (for example, sodium and chloride) 

in sea water. Salinity is most frequently measured in practical 

salinity units, a unit based on the properties of seawater 

conductivity, which is equivalent to parts per thousand or to 

grams per kilogram. 

Semelparous Refers to a strategy whereas reproduction or 

breeding occurs only once in a lifetime. 

Stock size (in weight) The weight of a fish stock or of some 

defined portion of a fish is known as biomass. Biomass is 

indicative of energy flows and the relative importance of a 

particular species to an assemblage or community level of 

organization. The established relations between fish length and 

weight are critical to the development of other key population 

parameters (Ricker, 1975). 

Temperature Temperature is arguably the most important 

environmental variable to fish; Reynolds (1977, p. 734) 

describes this importance thusly: 

Temperature serves as a proximate factor (cue, 

guidepost, sign stimulus, or directive factor) 

affecting locomotor responses of fishes. Although 

temperature can also serve as an ultimate ecological 

factor, as in behavioral thermoregulation, 

nonthermal factors may in some cases provide 

the ultimate adaptive or ecological value of a 

temperature response; some examples are habitat 

selection, intraspecific size segregation, interspecific 

niche differentiation, isolating mechanisms, 

predator avoidance, prey location, escape reactions, 

and migrations (thermoperiodic, diel, seasonal, 

spawning). Conversely, nonthermal variables 

such as light intensity or water depth may act as 

accessory proximate factors in thermoregulation. 

In spawning migrations, thermal requirements 

of eggs and larvae may take precedence over the 

(often different) preferenda or optima of adults. 

Although thermal responses of fishes are largely 

innate and species specific, ontogenetic and other 

changes can occur. Since temperature can serve as 

an unconditioned reinforcer in operant conditioning, 

thermal responses are not limited to simple kineses 

or taxes. Nonthermal factors such as photoperiod, 

circadian rhythms, currents, social and biotic 

interactions, stresses, infections, or chemicals can 

affect thermal responses, and may account for some 

lack of conformity between laboratory preferenda 

and field distributions and behaviors.

Trophic level The step in a nutritive series, or food chain, 

of an ecosystem. The organisms of a chain are classified into 

these levels on the basis of their feeding behavior. The first 

and lowest level contains the producers, green plants. The 

plants or their products are consumed by the second-level 

organisms—the herbivores, or plant eaters. At the third level, 

primary carnivores, or meat eaters, eat the herbivores; and 

at the fourth level, secondary carnivores, eat the primary 

carnivores. These categories are not strictly defined, as many 

organisms feed on several trophic levels; for example, some 

carnivores also consume plant materials or carrion and are 

called omnivores, and some herbivores occasionally consume 

animal matter. A separate trophic level, the decomposers, or 

transformers, consists of organisms such as bacteria and fungi 

that break down dead organisms and waste materials into 

nutrients usable by the producers. 
Chapter 8. References Cited

Able, K.W., and McAllister, D.E., 1980, Revision of the 

snailfish genus Liparis from Arctic Canada: Ottawa, 

Department of Fisheries and Oceans, Canadian Bulletin of 

Fisheries and Aquatic Sciences 208.

Abookire, A.A., Piatt, J.F., and Norcross, B.L., 2001, Juvenile 

groundfish habitat in Kachemak Bay, Alaska, during late 

summer: Alaska Fishery Research Bulletin, v. 8, no. 1, 

p. 45–56.

Abookire, A.A., Piatt, J.F., and Robards, M.D., 2000, 

Nearshore fish distributions in an Alaska estuary in relation 

to stratification, temperature and salinity: Estuarine, Coastal 

and Shelf Science, v. 51, no. 1, p. 45–59.

Acuna, E.I., Goddard, P., and Kotwicki, S., 2003, 2002 bottom 

trawl survey of the eastern Bering Sea continental shelf: 

National Marine Fisheries Service, Alaska Fisheries Science 

Center, Processed Report 2003-01.

Ahlstrom, E.H., Amaoka, K., Hensley, D.A., Moser, H.G., and 

Sumida, B.Y., 1984, Pleuronectiformes—Development, 

Moser, H.G., and others, eds., in Ontogeny and systematics 

of fishes: American Society of Ichthyologists and 

Herpetologists.

Akaike, H., 1973, Information theory and an extension of 

the maximum likelihood principle, in Petrov, B.N., and 

Caski, F., eds., Proceedings of the Second International 

Symposium on Information Theory: Budapest, Akademiai 

Kiado, p. 267–281.

Alaska Fisheries Science Center, 2013, Ichthyoplankton 

Information System: Alaska Fisheries Science Center, 

accessed November 29, 2013, at http://access.afsc.noaa.gov/

ichthyo/index.cfm.

Albers, W.D., and Anderson, P.J., 1985, Diet of Pacific cod, 

Gadus macrocephalus, and predation on the northern pink 

shrimp, Pandalus borealis, in Pavlof Bay, Alaska: Fishery 

Bulletin, v. 83, no. 4, p. 601–610.

Albert, O.T., Harbitz, A., and Høines, Å.S., 2003, Greenland 

halibut observed by video in front of survey trawl—

Behaviour, escapement, and spatial pattern: Journal of Sea 

Research, v. 50, no. 2–3, p. 117–127.

Alexander, V., 1992, Arctic marine ecosystems, chap. 17 of, 

Peters, R.L., and Lovejoy, T.L., eds., Global warming and 

biological diversity: New Haven, Conn., Yale University 

Press, p. 221–232.

Allen, M.J., and Smith, G.B., 1988, Atlas and zoogeography 

of common fishes in the Bering Sea and northeast Pacific: 

U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration Technical Report NMFS 66, 

151 p.

Allen, M.S., and Hightower, J.E., 2010, Fish population 

dynamics—Mortality, growth, and recruitment, in Hubert, 

W.A., and Quist, M.C., eds., Inland fisheries management in 

North America: Bethesda, Md., American Fisheries Society, 

p. 43–79.

Alt, K.T., 1969, Taxonomy and ecology of the inconnu, 

Stenodus leucichthys nelma, in Alaska: Anchorage, 

University of Alaska, Biological Papers of the University of 

Alaska, no. 12, 63 p.

Alt, K.T., 1973b, Age and growth of the inconnu (Stenodus 

leucichthys) in Alaska: Journal of the Fisheries Research 

Board of Canada, v. 30, no. 1, p. 457–459.

Alt, K.T., 1973a, Contributions to the biology of the Bering 

cisco (Coregonus laurettae) in Alaska: Journal of the 

Fisheries Research Board of Canada, v. 30, no. 12, 

p. 1,885–1,888.

Alt, K.T., 1987, Review of sheefish (Stenodus leucichthys) 

studies in Alaska: Juneau, Alaska Department of Fish and 

Game, Division of Sport Fish, Fishery Manuscript no. 3, 

69 p.

Alt, K.T., 1988, Biology and management of inconnu 

(Stenodus leucichthys) in Alaska: Finnish Fisheries 

Research, v. 9, p. 127–132.

Alt, K.T., 2008, Alaska wildlife notebook series—Sheefish: 

Alaska Department of Fish and Game.

Alton, M.S., Bakkala, R.G., Walters, G.E., and Munro, P.T., 

1988, Greenland turbot Reinhardtius hippoglossoides of 

the eastern Bering Sea and Aleutian Island region: National 

Oceanic and Atmospheric Administration, Technical Report 

NMFS 71, 31 p.

Alverson, D.L., 1960, A study of annual and seasonal 

bathymetric catch patterns for commercially important 

groundfishes of the Pacific Northwest coast of North 

America: Pacific Marine Fisheries Commission Bulletin 4, 

66 p.
700 Alaska Arctic Marine Fish Ecology Catalog

Anderson, P.J., and Piatt, J.F., 1999, Community 

reorganization in the Gulf of Alaska following ocean 

climate regime shift: Marine Ecology Progress Series, 

v. 189, p. 117–123.

Anderson, R.O., and Gutreuter, S.J., 1983, Length, weight, and 

associated structural indices, in Nielsen, L.A., and Johnson, 

D.L., eds., Fisheries Techniques (2d ed.):Bethesda, Md., 

American Fisheries Society, p. 283–300.

Andriyashev, A.P., 1954, Fishes of the northern seas of the 

U.S.S.R,—Keys to the fauna of the U.S.S.R.: Academy of 

Sciences of the U.S.S.R., Zoological Institute, no. 53, 566 

p. [In Russian, translation by Israel Program for Scientific 

Translation, Jerusalem, 1964, 617 p., available from U.S. 

Department of Commerce, Springfield, Virginia].

Andriyashev, A.P., 1961, Forms of the hook-horn sculpin 

genus Artediellus Jordan (Pisces: Cottidae) in the Bering 

Sea: Vopr Ikhtiol, v. 1, p. 231–242. [In Russian.]

Andriyashev, A.P., 1970, Cryopelagic fishes of Arctic and 

Antarctic and their significance in polar ecosystems,

in Holdgate, M.W., ed., Antarctic ecology: New York, 

Academic Press, p. 297–304.

Andriashev, A.P., 1986, Agonidae, in Whitehead, P.J.P., ed., 

Fishes of the north-eastern Atlantic and the Mediterranean: 

Paris, United Nations Educational, Scientific, and Cultural 

Organization, p. 1,265–1,268.

Andriyashev A.P., and Chernova, N.V., 1994, Annotated list of 

fishlike vertebrates and fish of the Arctic seas and adjacent 

waters: Journal of Ichthyology, v. 35, no. 1, p. 81–123.

Anisimov, O.A., Vaughan, D.G., Callaghan, T.V., Furgal, C., 

Marchant, H., Prowse, T.D., Vilhjálmsson, H., and Walsh, 

J.E., 2007, Polar regions (Arctic and Antarctic), in Parry, 

M.L., Canziani, O.F., Palutikof, J.P., van der Linden, P.J., 

and Hanson, C.E., eds., Climate change 2007—Impacts, 

adaptation and vulnerability, contribution of working group 

II to the fourth assessment report of the Intergovernmental 

Panel on Climate Change: Cambridge, U.K., Cambridge 

University Press, p. 653–685.

Ansaldo, M., Luquet, C.M., Evelson, P.A., Polo, J.M., and 

Llesuy, S., 2000, Antioxidant levels from different Antarctic 

fish caught around South Georgia Island and Shag Rocks: 

Polar Biology, v. 23, no. 3, p. 160–165.

Antonenko, D.V., and Vdovin, A.V., 2001, Seasonal 

distribution of the common greenling Hexagrammos stelleri

(Hexagrammidae) in Peter the Great Bay (the Sea of Japan): 

Journal of Ichthyology, v. 41, p. 524–528.

Archer, S.D., and Johnston, I.A., 1989, Kinematics of 

labriform and subcarangiform swimming in the Antarctic 

fish Notothenia neglecta: The Journal of Experimental 

Biology, v. 143, p. 195–210.

Armstrong, R.H., Carstensen, R.L., and Wilson, M.F., 2004, 

Hotspots—Bird survey of Mendenhall Wetlands April 

2002 to May 2003: Juneau Audubon Society and Taku 

Conservation Society, 9 p.

Armstrong, R.H., and Morrow, J.E., 1980, The Dolly Varden 

charr, Salvelinus malma, in Balon, E.K., ed., Charrs, 

salmonid fishes of the genus Salvelinus: The Hague, 

Netherlands, Dr. W. Junk B.V. Publishers, p. 99–140.

Aronovich, T.M., Doroshev, S.I., Spectorova, L.V., and 

Makhotin, V.M., 1975, Egg incubation and larval rearing 

of navaga (Eleginus navaga Pall.), Arctic Cod (Boreogadus 

saida lepechin) and Arctic flounder (Liopsetta glacialis

Pall.) in the laboratory: Aquaculture, v. 6, no. 3, p. 233–242.

Arvey, W.D., 1991, Stock status of anadromous Dolly Varden 

in waters of Alaska’s North Slope: Alaska Department of 

Fish and Game, Fishery Manuscript no. 91-3, 20 p.

Aschan, M., Karamushko, O.V., Byrkjedal, I., Wienerroither, 

R., Borkin, I.V., and Christiansen, J.S., 2009, Records of 

the gadoid fish Arctogadus glacialis (Peters, 1874) in the 

European Arctic: Polar Biology, v. 32, no. 7, p. 963–970.

Atkinson, E.G., and Percy, J.A., 1992, Diet comparison among 

demersal marine fish from the Canadian Arctic: Polar 

Biology, v. 11, no. 8, p. 567–573.

Axelsson, M., 2005, The circulatory system and its control, in

Farrell, A.P., and Steffensen, J.F., eds., The physiology of 

polar fishes: Amsterdam, Elsevier, p. 239–280.

Aydin, K., Gaichas, S., Ortiz, I., Kinzey, D., and Friday, N., 

2007, A comparison of the Bering Sea, Gulf of Alaska, and 

Aleutian Islands large marine ecosystems through food 

web modeling: U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration, National Marine 

Fisheries Service, Alaska Fisheries Science Center, NOAA 

Technical Memorandum NMFS-AFSC-178, 309 p.

Bailey, K.M., Abookire, A.A., and Duffy-Anderson, J.T., 

2008, Ocean transport paths for the early life history stages 

of offshore-spawning flatfishes—A case study in the Gulf of 

Alaska: Fish and Fisheries, v. 9, no. 1, p. 44–66.

Bailey, K.M., Brown, E.S., and Duffy-Anderson, J.T., 2003, 

Aspects of distribution, transport and recruitment of Alaska 

plaice (Pleuronectes quadrituberculatus) in the Gulf of 

Alaska and eastern Bering Sea: comparison of marginal and 

central populations: Journal of Sea Research, v. 50, no. 2–3, 

p. 87–95.

Bailey, K.M., and Picquelle, S.J., 2002, Larval distribution of 

offshore spawning flatfish in the Gulf of Alaska—Potential 

transport pathways and enhanced onshore transport during 

ENSO events: Marine Ecology Progress Series, v. 236, 

p. 205–217.
Chapter 8 701

Bailey, K.M., Powers, D.M., Quattro, J.M., Villa, G., 

Nishimura, A., Traynor, J.J., and Walters, G., 1999, 

Population ecology and structural dynamics of Walleye 

Pollock (Theragra chalcogramma), in Loughlin, T.R., and 

Ohtani, K., eds., Dynamics of the Bering Sea: Fairbanks, 

University of Alaska Sea Grant, AK-SG-99-03, p. 581–613.

Baird, P.A., and Gould, P., 1983, The breeding biology and 

feeding ecology of marine birds in the Gulf of Alaska: U.S. 

Fish and Wildlife Service, Denver Wildlife Research Center, 

Final Report, Outer Continental Shelf Environmental 

Assessment Program, Research Unit 341, Final Report 45 

(1986), p. 1–120.

Bakkala, R.G., 1970, Synopsis of biological data on the chum 

salmon, Oncorhynchus keta (Walbaum, 1792): U.S. Fish 

and Wildlife Service, FAO Fisheries Synopsis no. 41, 89 p.

Bakkala, R.G., 1981, Population characteristics and ecology 

of yellowfin sole, in Hood, D.W., and Calder, J.A., eds.: The 

eastern Bering Sea shelf—Oceanography and resources: 

National Oceanic and Atmospheric Administration, 

p. 553–574.

Balanov, A.A., Badaev, O.Z., Napazakov, V.V., and 

Chuchukalo, V.I., 2006, Distribution and some biological 

features of Lycodes raridens (Zoarcidae) in the western part 

of the Bering Sea: Journal of Ichthyology, v. 46, no. 2, 

p. 148–155.

Barber, W.E., Smith, R.L., Vallarino, M., and Meyer, R.M., 

1997, Demersal fish assemblages of the northeastern 

Chukchi Sea, Alaska: Fishery Bulletin, v. 95, no. 2, 

p. 195–209.

Barber, W.E., Smith, R.L., and Weingartner, T.J., 1994, 

Fisheries oceanography of the northeast Chukchi Sea: 

Final Report to U.S. Department of the Interior, Mineral 

Management Service, Anchorage, Alaska, OCS Study 

MMS-93-0051 [variously paged].

Beamish, R.J., 1980, Adult biology of the river lamprey 

(Lampetra ayresi) and the Pacific lamprey (Lampetra 

tridentata) from the Pacific Coast of Canada: Canadian 

Journal of Fisheries and Aquatic Sciences, v. 37, no. 11, 

p. 1,906–1,923.

Beamish, R.J., McFarlane, G.A., and King, J.R., 2005, 

Migratory patterns of pelagic fishes and possible linkages 

between open ocean and coastal ecosystems off the Pacific 

coast of North America: Deep-Sea Research II, v. 52, 

p. 739–755.

Bean, T.H., 1887, The fishery resources and fishing-grounds 

of Alaska, in Goode, G.B., ed., The fisheries and fishery 

industries of the United States, Section III: United States 

Commission of Fish and Fisheries, p. 81–115.

Beauchamp, D.A., Shepard, M.F., and Pauley, G.B., 1983, 

Species profiles—Life histories and environmental 

requirements of coastal fishes and invertebrates (Pacific 

Northwest)—Chinook Salmon: U.S. Fish and Wildlife 

Service, Biological Services Program FWS/OBS 82 (11.6), 

15 p.

Becker, K., Wöhrmann, A.P.A., and Rahmann, H., 1995, Brain 

gangliosides and cold-adaptation in high-Antarctic fish: 

Biochemical Systematics and Ecology, v. 23, nos. 7–8, 

p. 695–707.

Belyanina, T.N., 1968, Dynamics of smelt populations in 

sub-arctic waters: Rapports et procès-verbaux des réunions/ 

Conseil permanent international pour l’exploration de la 

mer, v. 158, p. 74–79.

Bendock, T.N., 1977, Beaufort Sea estuarine fishery study: 

Alaska Department of Fish and Game Annual Report, 

Contract #03-5-022-69, p. 670–729.

Benoit, D., Simard, Y., and Fortier, L., 2008, Hydroacoustic 

detection of large winter aggregations or Arctic cod 

(Boreogadus saida) at depth in ice-covered Franklin Bay 

(Beaufort Sea): Journal of Geophysical Research, v. 113, 

no. CO6S90, 9 p.

Benoit, D., Simard, Y., Gagné, J., Geoffroy, M., and Fortier, 

L., 2010, From polar night to midnight sun—Photoperiod, 

sea predation, and the diel vertical migrations of polar cod 

(Boreogadus saida) under landfast ice in the Arctic Ocean: 

Polar Biology, v. 33, no. 11, p. 1,505–1,520.

Benton, T.G., Plaistow, S.J., and Coulson, T.N., 2006, 

Complex population dynamics and complex causation—

Devils, details, and demography: Proceedings of the Royal 

Society B, Biological Sciences, v. 273, no. 1591, 

p. 1,173–1,181. 

Benz, G.W., Hocking, R., Kowunna, A., Bullard, S.A., and 

George, J.C., 2004, A second species of Arctic shark—

Pacific sleeper shark (Somniosus pacificus) from Point 

Hope, Alaska: Polar Biology, v. 27, no. 4, p. 250–252.

Benziger, D., and Umminger, B.L., 1973, Role of hepatic 

glycogenolytic enzymes in the cold-induced hyperglycemia 

of the killifish, Fundulus heteroclitus: Comparative 

Biochemistry and Physiology, v. 45, no. 3, p. 767–772.

Best, E.A., 1981, Halibut ecology, in Hood, D.W., and Calder, 

J.A., eds., The eastern Bering Sea shelf—Oceanography and 

resources: Seattle, University of Washington Press, 

p. 495–508.

Bjørkvoll, E., Grøtan, V., Aanes, S., Sæther, B.E., Engen, S., 

and Aanes, R., 2012, Stochastic population dynamics and 

life-history variation in marine fish species: American 

Naturalist, v. 180, no. 3, p. 372–387. 
702 Alaska Arctic Marine Fish Ecology Catalog

Blackburn, J.E., and Jackson, P.B., 1982, Seasonal 

composition and abundance of juvenile and adult marine 

finfish and crab species in the nearshore zone of Kodiak 

Island’s eastside during April 1978 through March 1979: 

Outer Continental Shelf Environmental Assessment 

Program, Alaska Department of Fish and Game, Final 

Report, Research Unit 552, p. 377–570.

Blackburn, J.E., Jackson, P.B., Warner, I.M., and Dick, M.H., 

1981, A survey for spawning forage fish on the east side 

of the Kodiak Archipelago by air and boat during spring 

and summer 1979: Outer Continental Shelf Environmental 

Assessment Program, Final Reports of Principal 

Investigators, RU 0552, p. 309–376.

Blood, D.M., Duffy-Anderson, J.T., Matarese, A.C., Sohn, 

D., Stabeno, P.J., and Cheng, W., 2009, Early life history 

of Greenland halibut (Reinhardtius hippoglossoides) in the 

eastern Bering Sea based on recent field studies (2007–

2009)—Spawning, distribution, and physical processes 

affecting drift and dispersal: Portland, Oregon, 33rd 

Annual Larval Fish Conference and American Society of 

Ichthyologists and Herpetologists.

Bluhm, B., Dunton, K., Grebmeier, J., and Sirenko, B., 2008, 

Benthos, in Hopcroft, R., Bluhm, B., and Gradinger, R., 

eds., Arctic Ocean synthesis—Analysis of climate change 

impacts in the Chukchi and Beaufort Seas with strategies 

for future research: Institute of Marine Sciences, University 

of Alaska Fairbanks, p. 56–65.

Bluhm, B.A., Gebruk, A.V., Gradinger, R., Hopcroft, R.R., 

Huettmann, F., Kosobokova, K.N., Sirenko, B.I., and 

Weslawski, J.M., 2011, Arctic marine biodiversity—An 

update of species richness and examples of biodiversity 

change: Oceanography, v. 24, no. 3, p. 232–248.

Bluhm, B.A., and Gradinger, R., 2008, Regional variability 

in food availability for Arctic marine mammals: Ecological 

Applications, v. 18, no. 2 Supplement, p. S77-S96.

Bond, W.A., 1982, A study of the fishery resources of 

Tuktoyaktuk Harbour, southern Beaufort Sea coast, with 

special reference to life histories of anadromous coregonids: 

Canadian Technical Report of Fisheries and Aquatic 

Sciences, no. 1119, 90 p.

Bond, W.A., and Erickson, R.N., 1987, Fishery data from 

Phillips Bay, Yukon, 1985: Winnipeg, Manitoba, Canadian 

Data Report of Fisheries and Aquatic Sciences, Central and 

Artcic Region, Department of Fisheries and Oceans, no. 

635, 47 p.

Bond, W.A., and Erickson, R.N., 1989, Summer studies of 

the nearshore fish community at Phillips Bay, Beaufort Sea 

coast, Yukon: Winnipeg, Manitoba, Canadian Technical 

Report of Fisheries and Aquatic Sciences, Central and 

Arctic Region, Department of Fisheries and Oceans, 

no. 1676, 102 p.

Bond, W.A., and Erickson, R.N., 1993, Fisheries 

investigations in coastal waters of Liverpool Bay, Northwest 

Territories: Winnipeg, Manitoba, Canada Department of 

Fisheries and Oceans, Central and Arctic Region, Canadian 

Manuscript Report of Fisheries and Aquatic Sciences 

No. 2204, 59 p.

Bonham, K., 1954, Food of the dogfish Squalus acanthias: 

Washington Department of Fisheries, v. 1, no. 2, p. 25–36.

Booth, S., Zeller, D., and Pauly, D., 2008, Baseline study of 

marine fisheries catches from Arctic Alaska 1950–2006: 

Report Fisheries Center, University of British Columbia, 

Canada, 16 p.

Bradstreet, M.S.W., 1982, Occurrence, habitat use, and 

behavior of seabirds, marine mammals, and Arctic cod at 

the Pond Inlet ice edge: Arctic, v. 35, no. 1, p. 28–40.

Bradstreet, M.S.W., Sekerak, A.D., Finley, K.J., Griffiths, 

W.B., Evans, C.R., Fabijan, M.F., and Stallard, H.E., 1986, 

Aspects of the biology of Arctic cod (Boreogadus saida) 

and its importance in Arctic marine food chains: Canada 

Department of Fisheries and Oceans, Canadian Technical 

Report of Fisheries and Aquatic Sciences, iss. 1491, 

ISSN 0706-6457, 193 p.

Brand, M.D., Steverding, D., Kadenbach, B., Stevenson, P.M., 

and Hafner, R.P., 1992, The mechanism of the increase 

in mitochondrial proton permeability induced by thyroid 

hormones: European Journal of Biochemistry, v. 206, no. 3, 

p. 775–781.

Braund, S.R., and Burnham, D.C., 1984, Subsistence 

economics and marine resource use patterns, in 

Truett, J.C., ed., The Barrow Arch environment and 

possible consequences of planned offshore oil and gas 

development: Outer Continental Shelf Environmental 

Assessment Program, National Oceanic and Atmospheric 

Administration/Ocean Assessment Division, p. 167–215.

Braund, S.R., and Kruse, J., 2009, Synthesis—Three decades 

of research on socioeconomic effects related to offshore 

petroleum development in coastal Alaska: U.S. Department 

of the Interior, Minerals Management Service, MMS Study 

Number 2009-006, 405 p., plus 3 appendixes.

Briggs, J.C., 1974, Marine zoogeography: McGraw-Hill, New 

York, 475 p.

Briggs, J.C., 1995, Global biogeography: Elsevier, 

Amsterdam, 452 p.

Briggs, J.C., 2003, Marine centres of origin as evolutionary 

engines: Journal of Biogeography, v. 30, p. 1–18. doi: 

10.1046/j.1365-2699.2003.00810.x

Briggs, J.C., 2004, Using marine invertebrate fossils for 

biogeography: Journal of Biogeography, v. 31, p. 1,711. doi: 

10.1111/j.1365-2699.2004.01157.x
Chapter 8 703

Briggs, J.C., and Bowan, B.W., 2012, A realignment of marine 

geographic provinces with particular references to fish 

distributions: Journal of Biogeography, v. 39, no. 1, 

p. 12–30.

Brix, O., Thorkildsen, S., and Colosimo, A., 2004, 

Temperature acclimation modulates the oxygen binding 

properties of the Atlantic cod (Gadus morhua L.) 

genotypes-HbI*1/1, HbI*1/2, and HbI*2/2- by changing 

the concentrations of their major hemoglobin components 

(results from growth studies at different temperatures): 

Comparative Biochemistry and Physiology Part A—

Molecular and Integrative Physiology, v. 138, no. 2, 

p. 241–251.

Brodeur, R.D., 1998, In situ observations of the association 

between juvenile fishes and scyphomedusae in the Bering 

Sea: Marine Ecology Progress Series, v. 163, p. 11–20.

Brodeur, R.D., and Livingston, P.A., 1988, Food habits and 

diet overlap of various Eastern Bering Sea fishes: National 

Oceanic and Atmospheric Administration Technical 

Memorandum NMFS F/NWC-127, 76 p.

Brodeur, R.D., and Rugen, W.C., 1994, Diel vertical 

distribution of ichthyoplankton in the northern Gulf of 

Alaska: Fishery Bulletin, v. 92, no. 2, p. 223–235.

Brown, E.D., 2002, Life history, distribution, and size 

structure of Pacific capelin in Prince William Sound and the 

northern Gulf of Alaska: ICES Journal of Marine Science, 

Journal du Conseil, v. 59, no. 5, p. 983–996.

Brown, J., and Green, J.M., 1976, Territoriality, habitat 

selection, and prior residency in underyearling Stichaeus 

punctatus (Pisces: Stichaeidae): Canadian Journal of 

Zoology, v. 54, no. 11, p. 1,904–1,907.

Brown, R.J., 2004, A biological assessment of whitefish 

species harvested during the spring and fall in the Selawik 

River Delta Selawik National Wildlife Refuge, Alaska: 

Alaska Fisheries Technical Report no. 77.

Brown, R.J., Brown, C., Braem, N.M., Carter, W.K., III, 

Legere, N., and Slayton, L., 2012, Whitefish biology, 

distribution, and fisheries in the Yukon and Kuskokwim 

river drainages in Alaska—A synthesis of available 

information: U.S. Fish and Wildlife Service, Alaska 

Fisheries Data Series Number 2012-4, 316 p.

 Browne, P.J., Laake, L., and DeLong, R.L., 2002, Improving 

pinnipeds diet analyses through identification of multiple 

skeletal structures in fecal samples: Fishery Bulletin, v. 100, 

no. 3, p. 423–433.

Bulatov, O.A., 1983, Distribution of eggs and larvae 

of Greenland halibut, Reinhardtius hippoglossoides

(Pleuronectidae), in the eastern Bering Sea: Journal of 

Ichthyology, v. 23 no. 1, p. 157–159.

Bureau of Ocean Energy Management, 2012, Proposed Final 

Outer Continental Shelf Oil and Gas Leasing Program 

2012-2017, U.S. Department of the Interior, 211 p., website 

accessed November 13, 2013, at http://www.boem.gov/

uploadedFiles/BOEM/Oil_and_Gas_Energy_Program/

Leasing/Five_Year_Program/2012-2017/Program_Area_

Maps/Alaska%20Program%20Areas.pdf.

Bureau of Ocean Energy Management Regulation and 

Enforcement, 2011, Chukchi Sea Planning Area: U.S. 

Department of the Interior, Bureau of Ocean Energy 

Management Regulation and Enforcement, Alaska OCS 

Region, OCS EIS/EA BOEMHRE 2011-041, Final 

Supplemental Environmental Impact Statement, v. 1, 

chaps. I–IV, 590 p.

Burgner, R.L., 1991, Life history of sockeye salmon 

(Oncorhynchus nerka), in Groot, C., and Margolis, L., eds., 

Pacific salmon life histories: University of British Columbia 

Press, p. 3–117.

 Burns, J.J., 1990, Proceedings of a technical workshop on 

fishes utilized in subsistence fisheries in National Petroleum 

Reserve-Alaska—Barrow, Alaska, October 26–28, 1988: 

Prepared for North Slope Borough, Department of Wildlife 

Management, Barrow, Alaska, 94 p.

Burton, D.W., 1988, Melanophore comparisons in different 

forms of ambicoloration in the flatfish Pseudopleuronectes 

americanus and Reinhardtius hippoglossoides: Journal of 

Zoology, v. 214, no. 2, p. 353–360.

Busby, M.S., Matarese, A.C., and Mier, K.L., 2000, Annual, 

seasonal, and diel composition of larval and juvenile fishes 

collected by dip-net in Clam Bay, Puget Sound, Washington, 

from 1985 to 1995: U.S. Department of Commerce, 

National Oceanic and Atmospheric Administration, NOAA 

Technical Memorandum NMFS-AFSC-111, 48 p.

Busby, M.S., Mier, K.L., and Brodeur, R.D., 2005, Habitat 

associations of demersal fishes and crabs in the Pribilof 

Islands region of the Bering Sea: Fisheries Research, v. 75, 

no. 1–3, p. 15–28.

Byers, T., and Prach, R.W., 1988, Diet of the kelp snailfish, 

Liparis tunicatus, in Jones Sound, Canadian High Arctic: 

The Canadian Field-Naturalist, v. 102, no. 2, p. 242–245.

Byrkjedal, I., Brattegard, T., and Møller, P.R., 2009, Lycodes 

adolfi Nielsen and Fosså, 1993 (Teleostei: Zoarcidae) 

recorded near Jan Mayen and in the eastern part of the 

Norwegian Sea: Fauna Norvegica, v. 28, p. 1–3.

Byrkjedal, I., Hadler-Jacobsen, S., Rees, D., and Orlov, A., 

2014, Molecular and morphological variation in ‘saddled’ 

fathead sculpins (Psychrolutidae—Cottunculus) of the 

northeast Atlantic, with a synonymy of three species: 

Journal of Fish Biology, v. 85, no. 3, p. 821–837.
704 Alaska Arctic Marine Fish Ecology Catalog

Byrkjedal, I., Langhelle, G., Wenneck, T.d.L., and 

Wienerroither, R., 2011, Lycodes adolfi Nielsen and Fosså, 

1993 (Teleostei: Zoarcidae) found in the Arctic Ocean: 

Polar Biology, v. 34, no. 3, p. 465–467.

Campbell, H.A., Fraser, K.P.P., Bishop, C.M., Peck, L.S., and 

Egginton, S., 2008, Hibernation in an Antarctic fish—On ice 

for winter: PLoS ONE: v. 3, iss. 3, e1743, 9 p., doi:10.1371/

journal.pone.0001743.

Carey, A.G., Jr,. 1987, Particle flux beneath fast ice in the 

shallow southwestern Beaufort Sea, Arctic Ocean: Marine 

Ecology Progress Series, v. 40, p. 247–257.

Carey, A.G., Jr., and Ruff, R.E., 1977, Ecological studies 

of the benthos in the western Beaufort Sea with special 

reference to bivalve molluscs, in Dunbar, M.J., ed., Polar 

oceans: Calgary, Alberta, Arctic Institute North America, 

p. 505–530.

Carey, M.P., Levin, P.S., Townsend, H., Minello, T.J., 

Sutton, G.R., Francis, T., Harvey, C.J., Toft, J.E., Arkema, 

K.K., Burke, J.L., Kim, C-K., Guerry, A., Plummer, M., 

Spiridonov, G., and Ruckelshaus, M., 2013, Characterizing 

coastal food webs with qualitative links to bridge the 

gaps between theory and the practice of ecosystem-based 

management: ICES Journal of Marine Science, v. 71, no. 3, 

p. 713–724, doi:10.1093/icesjms/fst012.

Carey, M.P., Sanderson, B.L., Barnis, K.A., and Olden, 

J.L., 2012, Native invaders—Challenges for science, 

management, policy, and society: Frontiers in Ecology and 

the Environment, v. 10, no. 7, p. 373–381.

Carlson, H.R., 1995, Consistent yearly appearance of age-0 

Walleye Pollock, Theragra chalcogramma, at a coastal site 

in southeastern Alaska, 1973–1994: Fishery Bulletin, v. 93, 

no. 2, p. 386–390.

Carmack, E., and Kulikov,Y., 1998, Wind-forced upwelling 

and internal Kelvin wave generation in Mackenzie Canyon, 

Beaufort Sea: Journal of Geophysical Research, v. 103, 

no. 18, p. 447–458.

Carmack, E., and Wassmann, P., 2006, Food webs and 

physical-biological coupling on pan Arctic shelves—

Unifying concepts and comprehensive perspectives: 

Progress in Oceanography, v. 71, nos. 2–4, p. 446–477.

Carmack, Eddy, Winsor, Peter, and Williams, William, 2015, 

The contiguous panarctic Riverine Coastal Domain—A 

unifying concept: Progress in Oceanography, v. 139, 

p. 13–23, accessed December 5, 2015, at http://dx.doi.

org/10.1016/j.pocean.2015.07.014.

Carothers, C., Cotton, S., and Moerlein, K., 2013, Subsistence 

use and knowledge of salmon in Barrow and Nuiqsut, 

Alaska: U.S. Department of the Interior, Bureau of Ocean 

Energy Management, OCS Study BOEM 2013-0015, 52 p.

Carscadden, E., and Vilhjálmsson, H., 2002, Capelin—What 

are they good for? Introduction: ICES Journal of Marine 

Science, v. 59, p. 863–869.

Chambers, R.C., 1997, Environmental influences on egg and 

propagule sizes in marine fishes, in Chambers, R.C., and 

Trippel, E.A., eds., Early life history and recruitment in fish 

populations: London, Chapman and Hall, p. 63–102.

Chang, Y.J., Sun, C.L., Zhang, Y., and Yeh, S.Z., 2010, 

Incorporation climate changes into population dynamic 

modeling—An individual-based modeling approach for 

lobster: Canadian Journal of Fisheries Aquatic Science, 

v. 68, no. 1, p. 122–136.

Chen, A.-L., and Sakurai, Y., 1993, Age and growth of saffron 

cod (Eleginus gracilis): Scientific Reports of Hokkaido 

Fisheries Experimental Station, v. 42, p. 251–264.

Cheng, C.-H.C., Cziko, P.A., and Evans, C.W., 2006, Nonhepatic origin of notothenioid antifreeze reveals pancreatic 

synthesis as common mechanism in polar fish freezing 

avoidance: Proceedings of the National Academy of 

Sciences of the USA, v. 103, no. 27, p. 10,491–10,496.

Chereshnev, I., Nazarkin, M.V., Skopets, M.B., Pitruk, D., 

Shestakov, A.V., Yabe, M., and others, 2001, Annotated list 

of fish-like vertebrates and fish in Tauisk Bay (northern part 

of the Sea of Okhotsk), in Andreev, A.V., and Bergmann, 

H.H., eds., Biodiversity and ecological status along the 

northern coast of the Sea of Okhotsk—A collection of 

study reports: Dalnauka Vladivostok, Russia, Institute of 

Biological Problems of the North, p. 64–86.

Chereshnev, I., Shestakov, A.V., and Skopets, M.B., 1999, On 

the distribution of silver smelts of the genus Hypomesus

(Osmeridae) in the northern part of the Sea of Okhotsk: 

Journal of Ichthyology, v. 39, no. 7, p. 498–503. 

Chernova, N.V., 1991, Aquarium observations of the snail 

fishes Careproctus reinhardti and Liparis liparis: Journal of 

Ichthyology, v. 31, no. 9, p. 65–70.

Chernova, N.V., 2008, Systematics and phylogeny of fish of 

the genus Liparis (Liparidae, Scorpaeniformes): Journal of 

Ichthyology, v. 48, no. 10, p. 831–852.

Chernova, N.V., 2011, Distribution patterns and chronological 

analysis of fish fauna of the Arctic region: Journal of 

Ichthyology, v. 51, no. 10, p. 825–924.

Cheung, W.W.L., Lam, V.W.Y., Sarmiento, J.L., Kearney, K., 

Watson, R., and Pauly, D., 2009, Projecting global marine 

biodiversity impacts under climate change scenarios: Fish 

and Fisheries, v. 10, p. 235–251.

Christiansen, J.S., ed., 2003, TUNU-1 Expedition—The fish 

fauna of the NE Greenland fjord systems—Technical report: 

Tromsø, Norway, University of Tromsø, Norwegian College 

of Fishery Science, Institute of Aquatic Resources, 33 p.
Chapter 8 705

Christiansen, J.S., Chernitsky, A.G., and Karamushko, O.V., 

1995, An Arctic teleost fish with a noticeably high body 

fluid osmolality: a note on the navaga, Eleginus navaga

(Pallas 1811), from the White Sea: Polar Biology, v. 15, 

no. 4, p. 303–306.

Christiansen, J.S., Fevolden, S.E., Karamushko, O.V., and 

Karamushko, L.J., 1998, Maternal output in polar fish 

reproduction, in di Prisco, G., Pisano, E., and Clarke, eds., 

Fishes of Antractica—A biological overview: Milan, Italy, 

Springer, p. 41–52.

Christiansen, J.S., Karamushko, O.V., Fevolden, S-E., and 

Præbel, K., 2008, Marine fishes of North East Greenland—

Diversity and adaptation: International Council for the 

Exploration of the Sea, ICES2008/B, 19 p. 

Christiansen, J.S., Mecklenburg, C.W., and Karamushko, O.V., 

2014, Arctic marine fishes and their fisheries in light of 

global climate change: Global Change Biology, v. 20, 

p. 352–359.

Christiansen, J.S., Reist, J.D., Brown, R.J., Brykov, V.A., 

Christensen, G., Christoffersen, K., Cott, P., Crane, P., 

Dempson, J.B., Docker, M., and others, 2013, Fishes, in

Meltofte, H., ed. Arctic biodiversity assessment—Status 

and trends in Arctic biodiversity: Akureyri, Iceland, 

Conservation of Arctic Flora and Fauna, p. 193–245, plus 

apps., http://www.arcticbiodiversity.is/the-report/chapters/

fishes.

Christensen, V., and Pauly, D., 1992, ECOPATH II—A 

software for balancing steady-state ecosystem models and 

calculating network characteristics: Ecological Modeling, 

v. 61, nos. 3–4, p. 169–185.

Chu, P.B., Wang, Q., and Bourke, R.H., 1999, A geometric 

model for the Beaufort/Chukchi Sea thermohaline structure: 

Journal of Atmospheric and Oceanic Technology, v. 16, 

p. 613–632.

Chuchukalo, V.I., Efimkin, A.Y., and Lapko, V.V., 1995, 

Feeding of some planktivorous fishes in the Sea of Okhotsk 

during the summer: Russian Journal of Marine Biology, 

v. 21, p. 112–115.

Clement, J.P., Bengtson, J.L., and Kelly, B.P., 2013, Managing 

for the future in a rapidly changing Arctic—A report to the 

President: Washington, D.C., Interagency Working Group 

on Coordination of Domestic Energy Development and 

Permitting in Alaska (Hayes, D.J., Chair): 59 p.

Close, D.A., Fitzpatrick, M.S., and Li, H.W., 2002, The 

ecological and cultural importance of a species at risk of 

extinction—Pacific lamprey: Fisheries, v. 27, no. 7, 

p. 19–25.

Clow, G.D., DeGange, A.R., Derksen, D.V., and Zimmerman, 

C.E., 2011, Climate change considerations, chap. 4 of

Holland-Bartels, Leslie, and Pierce, Brenda, eds., An 

evaluation of the science needs to inform decisions on Outer 

Continental Shelf energy development in the Chukchi and 

Beaufort Seas, Alaska: U.S. Geological Survey Circular 

1370, p. 81–109.

Coachman, L.R., Aagaard, K., and Tripp, R.B., 1975, 

Bering Strait-the regional physical oceanography: Seattle, 

University of Washington Press, 172 p.

Coad, B.W., and Reist, J.D., 2004, Annotated list of the Arctic 

marine fishes of Canada: Canadian Manuscript Report 

of Fisheries and Aquatic Sciences, Fisheries and Oceans 

Canada, no. 2674, 112 p.

Coad, B.W., Waszczuk, H., and Labignan, I., 1995, 

Encyclopedia of Canadian fishes: Canadian Museum of 

Nature and Canadian Sportfishing Productions, Inc., 928 p.

Coggan, R., 1996, Growth—Ration relationships in the 

Antarctic fish Notothenia coriiceps R. maintained under 

different conditions of temperature and photoperiod: Journal 

of Experimental Marine Biology and Ecology, v. 210, 

p. 23–35.

Cohen, D.M., Inada, T., Iwamoto, T., and Scialabba, N., 

1990, FAO species catalogue, volume 10—Gadiform 

fishes of the world (Order Gadiformes)—An annotated and 

illustrated catalogue of cods, hakes, grenadiers and other 

gadiform fishes known to date: Rome, Food and Agriculture 

Organization of the United Nations, FAO Fisheries 

Synopsis, no. 125, v. 10, 442 p.

Collette, B.B., and Klein-MacPhee, G., 2002, Bigelow 

and Schroeder’s fishes of the Gulf of Maine (3rd ed.): 

Washington, D.C., Smithsonian Institution Press, 882 p.

Collette, B.B., and Vecchione, M., 1995, Interactions between 

fisheries and systematics: Fisheries, v. 20, no. 1, p. 20–25.

Colonell, J.M., and Gallaway, B.J., 1997, Wind-driven 

transport and dispersion of age-0 Arctic ciscoes along 

the Alaska Beaufort coast: American Fisheries Society 

Symposium, no. 19, p. 90–103.

Committee on the Status of Endangered Wildlife in Canada, 

2004, COSEWIC assessment and update status report on 

the Bering cisco Coregonus laurettae in Canada: Ottawa, 

Canada, Committee on the Status of Endangered Wildlife in 

Canada, 19 p.

Cook, A.M., and Bundy, A., 2012, Use of fishes as sampling 

tools for understanding biodiversity and ecosystem 

functioning in the ocean: Marine Ecology Progress Series, 

v. 454, p. 1–18.
706 Alaska Arctic Marine Fish Ecology Catalog

Coombs, S., and Montgomery, J., 1994, Function and 

evolution of superficial neuromasts in an Antarctic 

notothenioid fish: Brain Behavior and Evolution, v. 44, 

p. 287–298.

Cooper, D.W., Maslenikov, K.P., and Gunderson, D.R., 2007, 

Natural mortality rate, annual fecundity, and maturity at 

length for Greenland halibut (Reinhardtius hippoglossoides) 

from the northeastern Pacific Ocean: Fishery Bulletin, 

v. 105, no. 2, p. 296–304.

Cossins, A.R., and Prosser, C.L., 1978, Evolutionary 

adaptation of membranes to temperature: Proceedings of the 

National Academy of Sciences of the USA, v. 75, no. 4, 

p. 2,040–2,043.

Courcelles, D., 2011, Re-evaluation of the length-weight 

relationship of Pacific halibut (Hippoglossus stenolepis), 

in IPHC Report of Assessment and Research Activities 

2011: International Pacific Halibut Commission (IPHC), 

p. 459–470 [Also available at http://www.iphc.int/library/

raras/254-rara-2011.html.]

Courtney, M., 2014, Dispersal patterns and summer 

distribution of adult Dolly Varden from the Wulik River, 

Alaska, evaluated with satellite telemetry [abs.]: Anchorage, 

Alaska Marine Science Symposium, January 20–24, 2014, 

p. 15.

Coyle, K.O., Gillispie, J.A., Smith, R.L., and Barber, W.E. 

1997, Food habits of four demersal Chukchi Sea fishes, in 

Reynolds, J.B., ed., Symposium 19, Proceedings of the Fish 

Ecology in Arctic North America Symposium: American 

Fisheries Society, Fairbanks, Alaska, May 19–21, 1992, 

p. 310–318.

Craig, P.C., 1984, Fish use of coastal waters of the Alaskan 

Beaufort Sea—A review: Transactions of the American 

Fisheries Society, v. 113, no. 3, p. 265–282.

Craig, P.C., 1987, Subsistence fisheries at coastal villages in 

the Alaskan Arctic, 1970–1986: Department of the Interior, 

Minerals Management Service, OCS Study MMS 87-6044, 

Technical Report No. 129, 63 p.

Craig, P.C., 1989a, An introduction to anadromous fishes in 

the Alaskan Arctic: Biological Papers of the University of 

Alaska, v. 24, p. 27–54.

Craig, P.C., 1989b, Subsistence fisheries at coastal villages 

in the Alaskan Arctic, 1970–1986: Biological Papers 

University of Alaska, no. 24, p. 131–152.

Craig, P.C., Griffiths, W.B., Haldorson, L., and McElderry, H., 

1982, Ecological studies of Arctic cod (Boreogadus saida) 

in Beaufort Sea coastal waters, Alaska: Canadian Journal of 

Fisheries and Aquatic Sciences, v. 39, no. 3, p. 395–406. 

Craig, P.C., Griffiths, W.B., Haldorson, L., and McElderry, H., 

1984a, Distributional patterns in an Alaska Arctic lagoon: 

Polar Biology, v. 4, no. 1, p. 9–18.

Craig, P.C., Griffiths, W.B., Johnson, S.R., and Schell, D.M., 

1984b, Trophic dynamics in an Arctic lagoon, in Barnes, 

P.W., and Reimnitz, E., eds., The Alaskan Beaufort Sea—

Ecosystems and environments: New York, Academic Press, 

p. 347–380.

Craig, P.C., and Haldorson, L.J., 1981, Beaufort Sea Barrier 

Island Lagoon ecological process studies—Final report, 

Simpson Lagoon—Fish: U.S. Department of Commerce and 

U.S. Department of the Interior, Biological Studies, 

p. 384–649.

Craig, P.C., and Haldorson, L.J., 1986, Pacific salmon in the 

North American Arctic: Arctic, v. 39, no. 1, p. 2–7. 

Craig, P.C., and McCart, P.J., 1974, Fall spawning and 

overwintering areas of fish populations along routes of 

proposed pipeline between Prudhoe Bay and the Mackenzie 

Delta 1972–1973, chap. 3 of McCart, P.J., ed., Life histories 

of anadromous and freshwater fish in the Western Arctic: 

Canadian Arctic Gas Study Ltd./Alaskan Arctic Gas Study 

Company, Biological Report Series, v. 20, p. 229.

Craig, P.C., and McCart, P.J., 1975, Fish utilization of 

nearshore coastal waters between the Colville and 

Mackenzie Rivers with an emphasis on anadromous species, 

in Craig, P.C., ed., Fisheries investigations in a coastal 

region of the Beaufort Sea: Calgary, Alberta, Canadian 

Arctic Gas Study Ltd. and Alaskan Arctic Gas Study Co., 

Biological Report Series 34: p. 172–219.

Craig, P.C., and Schmidt, D.R., 1985, Fish resources at 

Point Lay, Alaska: Barrow, Alaska, LGL Alaska Research 

Associates, Inc., North Slope Borough, Materials Source 

Division: 105 p.

Crane, P., Viavant, T., and Wenburg, J., 2005, Overwintering 

patterns of Dolly Varden Salvelinus malma in the 

Sagavanirktok River in the Alaska North Slope inferred 

using mixed-stock analysis: U.S. Fish and Wildlife Service, 

Alaska Fisheries Technical Report, no. 84, 25 p.

Crawford, R.E., and Jorgenson, J.K., 1996, Quantitative 

studies of Arctic cod (Boreogadus saida) schools—

Important energy stores in the Arctic food web: Arctic, 

v. 49, no. 2, p. 181–193.

Crawford, R.E., Vagle, S., and Carmack, E.C., 2012, Water 

mass and bathymetric characteristics of polar cod habitat 

along the continental shelf and slope of the Beaufort and 

Chukchi Seas: Polar Biology, v. 35, no. 2, p. 179–190.
Chapter 8 707

Criscione, C.D., Watral, V., Whipps, C.M., Blouin, M.S., 

Jones, S.R.M., and Kent, M.L., 2002, Ribosomal DNA 

sequences indicate isolated populations of Ichthyophonus 

hoferi in geographic sympatry in the north-eastern Pacific 

Ocean: Journal of Fish Disease, v. 25, no. 10, p. 575–582.

Cross, J.N., Fresh, K., Miller, B.S., Simenstad, C.A., 

Steinfort, S.N., and Fegley, J.C., 1978, Nearshore fish 

and invertebrates assemblages along the Strait of Juan de 

Fuca including food habits of the common inshore fishes: 

U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, Environmental Research 

Laboratories, NOAA Technical Memorandum ERL 

MESA-32.

Cui, X., Grebmeier, J.M., Cooper, L.W., Lovvorn, J.R., 

North, C.A., and Kolts, J.M., 2009, Spatial distributions 

of groundfish in the northern Bering Sea in relation to 

environmental variation: Marine Ecology Progress Series, 

v. 19, no. 6, p. 1,596–1,613.

Cziko, P.A., Evans, C.W., Cheng, C.-H.C., and DeVries, A.L., 

2006, Freezing resistance of antifreeze-deficient larval 

Antarctic fish: The Journal of Experimental Biology, v. 209, 

p. 407–420.

Datsky, A.V., 2015, Fish fauna of the Chukchi Sea and 

perspectives of its commercial use: Journal of Ichthyology, 

v. 55, p. 185–209.

Datsky A.V., and Andronov, P. Yu., 2007, Upper shelf 

ichtyocenose in the northwestern part of the Bering Sea, 

Magadan: NASC FEB RAS, 261 p. [In Russian.]

Davenport, J., 1992, Animal life at low temperature: London, 

Chapman and Hall, 246 p.

Davies, P.L., Baardsnes, J., Kuiper, M.J., and Walker, V.K., 

2002, Structure and function of antifreeze proteins: 

Physiological Transactions of the Royal Society London B, 

Biological Science, v. 357, no. 1423, p. 927–935.

Davies, P.L., Hew, C.L., and Fletcher, G.L., 1988, Fish 

antifreeze proteins—Physiology and evolutionary biology: 

Canadian Journal of Zoology, v. 66, no. 12, p. 2,611–2,617.

Davison, W., 2005, Antarctic fish skeletal muscle and 

locomotion, in Farrell, A.P., and Steffensen, J.F., eds., The 

physiology of polar fishes: Amsterdam, Elsevier, 

p. 317–349.

Davison, W., Axelsson, M., Nilsson, S., and Forster, M.E., 

1997, Cardiovascular function in Antarctic notothenioid 

fishes: Comparative and Biochemical Physiology, v. 118A, 

p. 1,001–1,008.

DeCicco, A.L., 1992, Long-distance movements of 

anadromous Dolly Varden between Alaska and the U.S.S.R.: 

Arctic, v. 45, no. 2, p. 120–123.

DeCicco, A.L., 1996, Abundance of Dolly Varden 

overwintering in the Wulik River, Northwest Alaska, during 

1994/1995: Alaska Department of Fish and Game, Fishery 

Data Series, No. 96-3, 24 p.

DeCicco, A.L., 1997, Movements of postsmolt anadromous 

Dolly Varden in northwestern Alaska, in Reynolds, J.B., ed., 

Fish ecology in Arctic North America: American Fisheries 

Society Symposium no. 19, Fairbanks, Alaska, May 19–21, 

1992 [Proceedings], p. 175–183.

DeGange, A.R., and Thorsteinson, L.K., 2011, Ecological 

and subsistence context, chap. 3 of Holland-Bartels, L., 

and Pierce, B., eds., An evaluation of the science needs 

to inform decisions on Outer Continental Shelf energy 

development in the Chukchi and Beaufort Seas, Alaska: 

U.S. Geological Survey Circular 1370, p. 41–80.

DeGraff, D., 1974, The life history of the pond smelt 

Hypomesus olidus Pallas (Osmeridae) in a small unnamed 

lake in the northern Yukon Territory: Arctic Gas Biological 

Report Series, v. 18, no. 2, p. 89.

DeGraaf, D.A., 1986, Aspects of the life history of the pond 

smelt (Hypomesus olidus) in the Yukon and Northwest 

Territories: Arctic, v. 39, no. 3, p. 260–263.

DeGraaf, D.A., and Machniak, K., 1977, Fisheries 

investigations along the cross delta pipeline route in the 

MacKenzie Delta, in McCart, P.J., ed., Studies to determine 

the impact of gas pipeline development on aquatic 

ecosystems: Calgary, Alberta, Canadian Arctic Gas Study, 

Ltd. and Alaskan Arctic Gas Study Co., Biological Report 

Series 39, p. 1–169.

Deibel, D., and Daly, L.L., 2007, Zooplankton processes in 

Arctic and Antarctic polynyas, in Smith, W.O., Jr., and 

Barber, D., eds., Polynyas—Windows into Polar oceans: 

Elsevier Oceanography Series 74, p. 271–322.

DeMartini, E.E., 1970, Reproductive colorations, paternal 

behavior and egg masses of kelp greenling, Hexagrammos 

decagrammus, and whitespotted greenling, H. stelleri: 

Northwest Science, v. 60, no. 1, p. 32–35.

 Denstad, J.-P., Aunaas, T., Börseth, J.B., Vollan Aarset, A., 

and Zachariassen, K.E., 1987, Thermal hysteresis antifreeze 

agents in fishes from Spitsbergen waters: Polar Research, 

v. 5, no. 2, p. 171–174.

Deriso, R.B., 1980, Harvesting strategies and parameter 

estimation for an age-structured model: Canadian Journal of 

Fisheries and Aquatic Sciences, v. 37, no. 2, p. 268–282.

Desjardins, M., Le François, N.R., Fletcher, G.L., and Blier, 

P.U., 2006, Seasonal modulation of plasma antifreeze 

protein levels in Atlantic (Anarhichas lupus) and spotted 

wolfish (A. minor): Journal of Experimental Marine Biology 

and Ecology, v. 335, no. 1, p. 142–150.
708 Alaska Arctic Marine Fish Ecology Catalog

Desjardins, M., Le François, N.R., Fletcher, G.L., and Blier, 

P.U., 2007, High antifreeze protein levels in wolfish 

(Anarchichas lupus) make them an ideal candidate for 

culture in cold, potentially ice laden waters: Aquaculture, 

v. 272, no. 1–4, p. 667–674.

DeVries, A.L., 1980, Biological antifreezes and survival in 

freezing environments, in Gilles, B., ed., Animals and 

environmental fitness, vol. 1: Oxford, England, Pergamon 

Press, p. 583–607,

DeVries, A.L., and Cheng, C.-H. C., 2005, Antifreeze proteins 

and organismal freezing avoidance in polar fishes, in

Farrell, A.P., and Steffensen, J.F., eds., The physiology of 

Polar fishes: Amsterdam, Elsevier, p. 155–201.

DeVries, A.L., and Lin, Y., 1977, The role of glycoprotein 

antifreezes in the survival of Antarctic fishes, in Llano, 

G.A., ed., Adaptations within Antarctic ecosystems: 

Houston, Texas, Gulf Publishing, p. 439–458.

DeVries, A.L., and Steffensen, J.F., 2005, The Arctic and 

Antarctic polar marine environments, in Farrell, A.P., 

and Steffensen, J.F., eds., The physiology of polar fishes: 

Amsterdam, Elsevier, p. 1–24.

Dick, M.H., and Warner, I.M., 1982, Pacific sand lance, 

Ammodytes hexapterus Pallas, in the Kodiak island group, 

Alaska: Syesis, v. 15, p. 43–50.

Dobbs, G.H., and DeVries, A.L., 1975, Renal function in 

Antarctic teleosts fish—Serum and urine composition: 

Marine Biology, v. 29, p. 59–70.

Dodson, J.J., Tremblay, S., Colombani, F., Carscadden, J.E., 

and Lecomte, F., 2007, Trans-Arctic dispersals and the 

evolution of a circumpolar marine fish species complex, the 

capelin (Mallotus villosus): Molecular Ecology, v. 16, 

p. 5,030–5,043.

Doney, S.C., Fabry, V.J., Feely, R.A., and Klepas, J.A., 2009, 

Ocean acidification—The other CO2

 problem: Annual 

Review of Marine Science, v. 1, p. 169–192.

Doney, S.C., and others, 2012, Climate change impacts on 

marine ecosystems: Annual Review of Marine Science, v. 4, 

p. 11–37.

Dorn, M., Aydin, K., Barbeaux, S., Guttormsen, M., Megrey, 

B., Spalinger, K., and others, 2007, Gulf of Alaska walleye 

pollock, in The Plan Team for the Groundfish Fisheries 

of the Gulf of Alaska, compilers, Stock assessment and 

fishery evaluation report for the groundfish resources of the 

Gulf of Alaska: Anchorage, Alaska, North Pacific Fishery 

Management Council, p. 51–168.

Doyle, M.J., Mier, K.L., Busby, M.S., and Brodeur, R.D., 

2002, Regional variation in springtime ichthyoplankton 

assemblages in the northeast Pacific Ocean: Progress in 

Oceanography, v. 53, no. 2, p. 247–281.

 Driedzic, W.R., Clow, K.A., Short, C.E., and Ewart, K.V., 

2006, Glycerol production in rainbow smelt (Osmerus 

mordax) may be triggered by low temperature alone and 

is associated with the activation of glycerol-3-phosphate 

dehydrogenase and glycerol-3-phosphatase: Journal of 

Experimental Biology, v. 209, no. 6, p. 1,016–1,023.

Driedzic, W.R., and Short, C.E., 2007, Relationship between 

food availability, glycerol and glycogen levels in lowtemperature challenged rainbow smelt Osmerus mordax: 

Journal of Experimental Biology, v. 210, no. 16, 

p. 2,866–2,872.

Dudnik, Y.I., and Dolganov, V.N., 1992, Distribution and 

abundance of fish on the continental slopes of the Sea of 

Okhotsk and of the Kuril Islands during the summer of 

1989: Journal of Ichthyology, v. 32, no. 9, p. 58–76.

Dudnik, Y.I., and Usol’tsev, E.A., 1964, The herrings of the 

eastern part of the Bering Sea, in Moiseev, P.A., ed., Soviet 

fisheries investigations in the Northeastern Pacific, part 1: 

Jerusalem, Israel Program for Scientific Translations [1968], 

p. 236–240.

Duffy-Anderson, J.T., Doyle, M.J., Mier, K.L., Stabeno, P.J., 

and Wilderbuer, T.K., 2010, Early life ecology of Alaska 

plaice (Pleuronectes quadrituberculatus) in the eastern 

Bering Sea— Seasonality, distribution, and dispersal: 

Journal of Sea Research, v. 64, no. 1–2, p. 3–14.

Dunbar, M.J., and Hildebrand, H.H., 1952, Contributions 

to the study of the fishes of Ungava Bay: Journal of the 

Fisheries Research Board of Canada, v. 9, no. 2, p. 83–126.

Duncker, B.P., Koops, M.D., Walker, V.K., and Davies, P.L., 

1995, Low temperature persistence of type I antifreeze 

protein is mediated by cold-specific mRNA stability: FEBS 

Letters v. 377, no. 2, p. 185–188.

Dunn, J.R., and Matarese, A.C., 1987, A review of the early 

life history of northeast Pacific gadoid fishes: Fisheries 

Research, v. 5, nos. 2–3, p. 163–184. 

Dunn, J.R., and Sample, T.M., 1976, Greenland turbot (family 

Pleuronectidae) in Pereya, W.T., Reeves, J.E., and Bakkala, 

R.G., eds., Demersal fish and shellfish resources of the 

eastern Bering Sea in the baseline year 1975: Seattle, 

Washington, National Marine Fisheries Service, Northwest 

and Alaska Fisheries Science Center, Processed Report, 

p. 475–487.

Dunn, J.R., and Vinter, B.M., 1984, Development of larvae of 

the saffron cod, Eleginus gracilis, with comments on the 

identification of gadid larvae in Pacific and Arctic waters 

contiguous to Canada and Alaska: Canadian Journal of 

Fisheries and Aquatic Sciences, v. 41, no. 2, p. 304–318.
Chapter 8 709

Dunton, K.H., Goodall, J.L., Schonberg, S.V., Grebmeier, 

J.M., and Maidment, D.R., 2005. Multi-decadal synthesis 

of benthic–pelagic coupling in the western Arctic: role of 

cross-shelf advective processes, Deep-Sea Research II, 

v. 52, p. 3,462–3,477.

Dunton, K.H., Reimnitz, E., and Schonberg, S., 1982, An 

Arctic kelp community in the Alaskan Beaufort Sea: Arctic, 

v. 35, no. 4, p. 465–484.

Dunton, K.H., Schonberg, S.V., and Cooper, L.W., 2012, Food 

web structure of the Alaskan nearshore shelf and estuarine 

lagoons of the Beaufort Sea: Estuaries and Coasts, v. 35, 

no. 2, p. 416–435.

Dunton, K.H., Weingartner, T., and Carmack, E.C., 2006, The 

nearshore western Beaufort Sea ecosystem: Progress in 

Oceanography, v. 71, no. 2–4, p. 362–378.

D’yakov, Y.P., 1982, The fecundity of the Greenland halibut, 

Reinhardtius hippoglossoides (Pleuronectidae), from the 

Bering Sea: Journal of Ichthyology, v. 22, no. 5, p. 59–64.

Dykstra, C.L., Olsen, W., Williams, G.H., and Gregg, J.L., 

2012, Ichthyophonus prevalence in Pacific halibut, in

IPHC Report of Assessment and Research Activities 2012: 

International Pacific Halibut Commission (IPHC), 

p. 437–446, [Also available at http://www.iphc.int/

publications/rara/2012/rara2012437_ichthyophonus.pdf.]

Eastman, J.T., 1997, Comparison of the Antarctic and Arctic 

fish faunas: Cybium, v. 21, no. 4, p. 335–352.

Ege, R., and Krogh, A., 1914, On the relation between 

the temperature and the respiratory exchange in fishes: 

Internationale Revue der gesamten Hydrobiologie und 

Hydrographie, v. 7, no. 1, p. 48–55.

Egginton, S., 1996, Blood rheology of Antarctic fishes—

Viscosity adaptations at very low temperatures: Journal of 

Fish Biology, v. 48, p. 513–521.

Ehrlich, P., Dobkin, D.S., and Wheye, D., 1988, The birder’s 

handbook: New York, Simon and Schuster, Inc., 816 p.

Elith, J., and Leathwick, J.R., 2009, Species distribution 

models—Ecological explanation and prediction across 

space and time: Annual Review of Ecology, Evolution, and 

Systematics, v. 40, p. 677–697.

Ellis, D.V., 1962, Observations on the distribution and ecology 

of some Arctic fish: Arctic, v. 15, no. 3, p. 179–189.

Emmett, R.L., Stone, S.L., Hinton, S.A., and Monaco, 

M.E., 1991, Distribution and abundance of fishes and 

inverteprates in west coast estuaries, Volume II—Species 

life history summaries: National Oceanic and Atmospheric 

Administration/NOS Strategic Environmental Assessments 

Division, ELMR Report no. 8, 327 p.

Enevoldsen, L.T., Heiner, I., DeVries, A.L., and Steffensen, 

J.F., 2003, Does fish from the Disko Bay area of Greenland 

possess antifreeze proteins during the summer?: Polar 

Biology, v. 26, no. 6, p. 365–370.

Ennis, G.P., 1968, Occurrences of the staghorn sculpin 

(Gymnocanthus tricuspis) in Newfoundland waters: Journal 

of the Fisheries Research Board of Canada, v. 25, no. 12, 

p. 2,729–2,731.

Ennis, G.P., 1969, The biology of the shorthorn sculpin 

Myoxocephalus scorpius (L.) in Newfoundland waters: 

Saint John’s, Newfoundland, Memorial University, Master’s 

thesis.

Ennis, G.P., 1970a, Age, growth, and sexual maturity 

of the shorthorn sculpin, Myoxocephalus scorpius, in 

Newfoundland waters: Journal of the Fisheries Research 

Board of Canada v. 27, no. 12, p. 2,155–2,158.

Ennis, G.P., 1970b, Reproduction and associated behaviour 

in the shorthorn sculpin, Myoxocephalus scorpius in 

Newfoundland waters: Journal of the Fisheries Research 

Board of Canada, v. 27, no. 11, p. 2,037–2,045.

Eschmeyer, W.N., Frick, R., Fong, J.D., and Polack, D.A., 

2010, Marine fish diversity—History of knowledge and 

discovery (Pisces): Zootaxa 2525, p. 19–50.

Eschmeyer, W.N., Frick, R., and van der Laan, R., eds, 2015, 

Catalog of fishes—Genera, species, references: California 

Academy of Sciences, Web site accessed June 17, 2016, at 

http://researcharchive.calacademy.org/research/ichthyology/

catalog/fishcatmain.asp.

Essington, T.E., Kitchell, J.F., and Walters, C.J., 2001, The 

von Bertalanffy growth function, bioenergetics, and 

consumption rates of fish: Canadian Journal of Fisheries 

and Aquatic Sciences, v. 58, no. 11, p. 2,129–2,138.

Evans, C.L., Reist, J.D., and Minns, C.K., 2002, Life history 

characteristics of freshwater fishes occurring in the 

Northwest Territories and Nunavut, with major emphasis on 

riverine habitat requirements: Canadian Manuscript Report 

of Fisheries and Aquatic Sciences, no. 2614, 169 p.

Evans, C.W., Gubala, V., Nooney, R., Williams, D.E., 

Brimble, M.A., and DeVries, A.L., 2011, How do Antarctic 

notothenioid fishes cope with internal ice? A novel function 

for antifreeze glycoproteins: Antarctic Science, v. 23, no. 1, 

p. 57–64.

Fabry, V.J., McClintock, J.B., Mathis, J.T., and Grebmeier, 

J.M., 2009, Ocean acidification at high latitudes—The 

bellweather: Oceanography, v. 22, no. 4, p. 160.
710 Alaska Arctic Marine Fish Ecology Catalog

Fadeev, N.S., 1965, Comparative outline of the biology of 

flatfishes in the southeastern part of the Bering Sea and 

condition of their resources, in Moiseev, P.A., ed., Soviet 

fisheries investigations in the northeastern Pacific, Part IV: 

Jerusalem, Israel Program for Scientific Translations, 

p. 112–146. 

Fadeev, N.S., 1970, The fishery and biological characteristics 

of yellowfin soles in the eastern part of the Bering Sea in

Moiseev, P.A., ed., Soviet fisheries investigations in the 

northeastern Pacific, Part V: Jerusalem, Israel Program for 

Scientific Translations, p. 332–396.

Falk-Petersen, I.-B., Frivoll, V., Gulliksen, B., and Haug, 

T., 1986, Occurrence and size/age relations of polar cod, 

Boreogadus saida (Lepechin), in Spitsbergen coastal 

waters: Sarsia, v. 71, no. 3–4, p. 235–245.

Falk-Petersen, I.-B., Frivoll, V., Gulliksen, B., Haug, T., 

and Vader, W., 1988, Age/size relations and food of two 

snailfishes, Liparis gibbus and Careproctus reinhardtii

(Teleostei, Liparididae) from Spitsbergen coastal waters: 

Polar Biology, v. 8, no. 5, p. 353–358.

Farwell, M.K., Green, J.M., and Pepper, V.A., 1976, 

Distribution and known life history of Stichaeus punctatus

in the northwest Atlantic: Copeia, v. 1976, no. 3, 

p. 598–602.

Fechhelm, R.G., Baker, J.S., Griffiths, W.B., and Schmidt, 

D.R., 1989, Localized movement patterns of least cisco 

(Coregonus sardinella) and Arctic cisco (C. autumnalis) in 

the vicinity of a solid-fill causeway: Biological Papers of 

the University of Alaska, v. 24, p. 75–106.

Fechhelm, R.G., Craig, P.C., Baker, J.S., and Gallaway, 

B.J., 1984, Fish distribution and use of nearshore waters 

in the northeastern Chukchi Sea: Final Report, LGL 

Ecological Research Associates Inc., National Oceanic and 

Atmospheric Administration, OMPA/OCSEAP, 190 p.

Fechhelm, R.G., and Fissel, D.B., 1988, Wind-aided 

recruitment of Canadian Arctic cisco (Coregonus 

autumnalis) into Alaskan waters: Canadian Journal of 

Fisheries and Aquatic Sciences, v. 45, no. 5, p. 906–910.

Fechhelm, R.G., and Griffiths, W.B., 1990, The effect of wind 

on the recruitment of Canadian Arctic cisco (Coregonus 

autumnalis) into the central Alaskan Beaufort Sea: 

Canadian Journal of Fisheries and Aquatic Sciences, v. 47, 

p. 2,164–2,171.

Fechhelm, R.G., Griffiths, W.B., Bryan, J.D., Gallaway, 

B.J., and Wilson, W.J., 1995a, Application of an in situ

growth model—Inferred instance of interspecific trophic 

competition between anadromous fishes of Prudhoe Bay, 

Alaska: Transactions of the American Fisheries Society, v. 

124, no. 1, p. 55–69.

Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Gallaway, B.J., 

and Bryan, J.D., 1995b, Intra- and interseasonal changes in 

the relative condition and proximate body composition of 

Broad Whitefish from the Prudhoe Bay region of Alaska: 

Transactions of the American Fisheries Society, v. 124, 

no. 4, p. 508–519.

Fechhelm, R.G., Griffiths, W.B., Wilson, W.J., Trimm, B.A., 

and Colonell, J.M., 1996, The 1995 fish and oceanography 

study in Mikkelsen Bay, Alaska: Anchorage, Alaska, 

Prepared by LGL Alaska Research Associates and 

Woodward-Clyde Consultant for BP Exploration (Alaska) 

Inc., 102 p. plus apps.

Fechhelm, R.G., Streever, B., and Gallaway, B.J., 2007, 

The Arctic cisco (Coregonus autumnalis) subsistence and 

commercial fisheries, Colville River, Alaska—A conceptual 

model: Arctic, v. 60, no. 4, p. 421–429.

Fechhelm, R.G., Wade, G.D., and Link, M.R., 2006, Nearshore 

Beaufort Sea fish monitoring in the Prudhoe Bay region, 

2005: Anchorage, Alaska, Report to British Petroleum 

(Alaska) Inc. by LGL Alaska Research Associates, Inc., 

72 p. plus apps.

Fedorov, V.V., 1986, Cottidae, in Whitehead, P.J.P., Bauchot, 

M.-L., Hureau, J.-C., Nielsen, J., and Tortonese, E., eds., 

Fishes of the North-eastern Atlantic and the Mediterranean: 

UNESCO, Paris, v. 3, p. 1,243–1,260.

Fedorov, V.V., 2000, Species composition, distribution and 

habitation depths of the northern Kuril Islands fish and 

fish like species, in Koteneva, B.N., ed., Commercial and 

biological studies of fishes in the Pacific waters of the Kuril 

Islands and adjacent areas of the Okhotsk and Bering Seas 

in 1992–1998: Moscow, VNIRO Publishing, collected 

papers, p. 7–41. [In Russian.]

Field, L.J., 1988, Pacific sand lance, Ammodytes hexapterus, 

with notes on related Ammodytes species, in Wilimovsky, 

N.J., Incze, L.S., and Westrheim, S.J., eds., Species 

synopses, life histories of selected fish and shellfish of the 

northeast Pacific and Bering Sea: Seattle, University of 

Washington, Washington Sea Grant Program and Fisheries 

Research Institute.

Finley, K.J., and Gibb, E.J., 1982, Summer diet of the narwhal 

(Monodon monoceros) in Pond Inlet, northern Baffin Island: 

Canadian Journal of Zoology, v. 60, no. 12, p. 3,353–3,363.

Fletcher, G.L., 1975, The effects of capture, “stress,” and 

storage of whole blood on the red blood cells, plasma 

proteins, glucose, and electrolytes of the winter flounder 

(Pseudopleuronectes americanus): Canadian Journal of 

Zoology, v. 53, no. 2, p. 197–206.
Chapter 8 711

Fletcher, G.L., 1977, Circannual cycles of blood plasma 

freezing point and Na+ and CL– concentrations in 

Newfoundland winter flounder (Pseudopleuronectes 

americanus)—Correlation with water temperature and 

photoperiod: Canadian Journal of Zoology, v. 55, no. 5, 

p. 789–795.

Fletcher, G.L., 1981, Effects of temperature and photoperiod 

on the plasma freezing point depression, Cl ̄ concentration, 

and protein “antifreeze” in winter flounder: Canadian 

Journal of Zoology, v. 59, no. 2, p. 193–201.

Fletcher, G.L., Addison, R.F., Slaughter, D., and Hew, C.L., 

1982, Antifreeze proteins in the Arctic shorthorn sculpin 

(Myoxocephalus scorpius): Arctic, v. 35, no. 2, p. 302–306.

Fletcher, G.L., Haya, K., King, M.J., and Reisman, H.M., 

1985a, Annual antifreeze cycles in Newfoundland, 

New Brunswick and Long island winter flounder, 

Pseudopleuronectes americanus: Marine Ecology Progress 

Series, v. 21, p. 205–212.

Fletcher, G.L., Hew, C.L., and Davis, P.L., 2001, Antifreeze 

proteins of teleost fishes: Annual Review of Physiology, 

v. 63, p. 359–390.

Fletcher, G.L., Hew, C.L., Li, X.M., Haya, K., and Kao, 

M.H., 1985b, Year-round presence of high levels of 

plasma antifreeze peptides in a temperate fish, ocean pout 

(Macrozoarces americanus): Canadian Journal of Zoology, 

v. 63, no. 3, p. 488–493.

Fletcher, G.L., Idler, D.R., Vaisius, A., and Hew, C.L., 1989, 

Hormonal regulation of AFP gene expression in winter 

flounder: Fish Physiology and Biochemistry, v. 7, no. 1–6, 

p. 387–393.

Fletcher, G.L., Kao, M.H., and Haya, K., 1984, Seasonal and 

phenotypic variations in plasma protein antifreeze levels 

in a population of marine fish, sea raven (Hemitripterus 

americanus): Canadian Journal of Fisheries and Aquatic 

Sciences, v. 41, no. 5, p. 819–824.

Fletcher, G.L., King, M.J., and Kao, M.H., 1987, Low 

temperature regulation of antifreeze glycopeptides levels in 

Atlantic cod (Gadus morhua): Canadian Journal of Zoology, 

v. 65, no. 2, p. 227–233.

Fluharty, D., 2012, Recent developments at the Federal 

level in ocean policymaking in the United States: Coastal 

Management, v. 40, no. 2, p. 209–221.

Fordham, D.A., and others, 2013, Population dynamics can be 

more important than physiological limits for determining 

range shifts under climate change, Global Change Biology, 

v. 19, p. 3,224–3,237.

Forrest, A., Sampei, M., Hatton, H., Makabe, R., Sasaki, H., 

Fukuchi, M., Wassman, P., and Fortier, L., 2007, Particulate 

organic carbon fluxes on the slope of the MacKenzie Shelf 

(Beaufort Sea)—Physical and biological forcing of shelfbasin exchanges: Journal of Marine Systems: v. 68, no. 1–2, 

p. 39–54.

Forrester, C.R., 1964, Laboratory observations on embryonic 

development and larvae of the Pacific cod (Gadus 

macrocephalus Tilesius): Journal of the Fisheries Research 

Board of Canada, v. 21, no. 1, p. 9–16.

Forsberg, J.E., 2011, Age distribution of Pacific halibut in the 

2011 IPHC stock assessment setline survey, in International 

Pacific Halibut Commission (IPHC) Report of Assessment 

and Research Activities 2011: International Pacific Halibut 

Commission, p. 553–560. [Also available at http://www.

iphc.int/library/raras/254-rara-2011.html.]

Fortier, L., Sirois, P., Michaud, J., and Barber, D., 2006, 

Survival of Arctic cod larvae (Boreogadus saida) in relation 

to sea ice and temperature in the Northeast Water Polynya 

(Greenland Sea): Canadian Journal of Fisheries and Aquatic 

Sciences, v. 63, no. 7, p. 1,608–1,616.

Fourney, R.M., Fletcher, G.L., and Hew, C.L., 1984, The 

effects of long day lengths on liver antifreeze mRNA in 

winter flounder: Canadian Journal of Zoology, v. 62, no. 8, 

p. 1,456–1,460.

Foy, R.J., Crapo, C.A., and Kramer, D.E., 2006, Investigating 

the roles of temperature and exercise in the development of 

chalkiness in Pacific halibut: International Pacific Halibut 

Commission, Technical Report No. 50.

Fresh, K.L., 1979, Distribution and abundance of fishes 

occurring in the nearshore surface waters of northern Puget 

Sound, Washington: Seattle, University of Washington, 

Master’s thesis.

Frey, K.E., Arrigo, K.R., and Gradinger, R.R., 2011, Arctic 

report card, update for 2011—Arctic Ocean primary 

productivity: National Oceanic and Atmospheric 

Administration Arctic Theme Page Web site, accessed 

October 25, 2012, at http://www.arctic.noaa.gov/report11/

primary_productivity.html.

Froese, R., and Pauly, D., eds., 2012, FishBase—Global 

information system on fishes: FishBase database, accessed 

July 8, 2012, at http://www.fishbase.org.

Frost, K.J., and Lowry, L.F., 1980, Feeding of ribbon seals 

(Phoca fasciata) in the Bering Sea in spring: Canadian 

Journal of Zoology, no. 58, v. 9, p. 1,601–1,607.

Frost, K.J., and Lowry, L.F., 1981, Trophic importance of 

some marine gadids in Northern Alaska and their bodyotolith size relationships: Fishery Bulletin, v. 79, no. 1, 

p. 187–192.
712 Alaska Arctic Marine Fish Ecology Catalog

Frost, K.J., and Lowry, L.F., 1983, Demersal fishes and 

invertebrates trawled in the northeastern Chukchi and 

western Beaufort Seas 1976–1977: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration, Technical Report NMFS-SSRF-764, 22 p.

Frost, K.J., and Lowry, L.F., 1984, Trophic relationships 

of vertebrate consumers in the Alaskan Beaufort Sea, in

Barnes, P.W., Schell, D.M., and Reimnitz, E., eds., The 

Alaska Beaufort Sea—Ecosystems and environments: San 

Diego, Academic Press, p. 382–401.

Fruge, D.J., and Wiswar, D.W., 1991, First records of the 

Bering wolffish, Anarhichas orientalis, for the Alaskan 

Beaufort Sea: Canadian Field-Naturalist, v. 105, no. 1, 

p. 107–109.

Fruge, D.J., Wiswar, D.W., Dugan, L.J., and Palmer, D.E., 

1989, Fish population characteristics of Arctic National 

Wildlife Refuge coastal waters, summer 1988: Fairbanks, 

Alaska, U.S. Fish and Wildlife Service, Fishery Assistance 

office, Progress Report, 73 p.

Frydenberg, O., Møller, D., Névdal, G., and Sick, K., 1965, 

Hemoglobin polymorphism in Norwegian cod populations: 

Hereditas, v. 53, no. 1, p. 257–271.

Furniss, R.A., 1975, Inventory and cataloging of Arctic area 

waters: Alaska Department of Fish and Game, Federal Aid 

in Fish Restoration, Annual Report of Performance, Project 

F-9-7, Study G-1-1, v. 16, 47 p.

Gallaway, B.J., and Fechhelm, R.G., 1997, Anadromous and 

amphidromous fishes, in Truett, J.C., and Johnson, S.R., 

eds., The natural history of an Arctic oil field: San Diego, 

Academic Press, p. 349–369.

Gallaway, B.J., Griffiths, W.B., Craig, P.C., Gazey, W.J., and 

Helmericks, J.W., 1983, An assessment of the Colville 

River Delta stock of Arctic cisco—Migrants from Canada?: 

Biological Papers of the University of Alaska, no. 21, 

p. 4–23.

Gallaway, B.J., and Norcross, B.L., eds., 2011, A synthesis 

of diversity, distribution, abundance, age, size and diet 

of fishes in the lease sale 193 Area of the northeastern 

Chukchi Sea: Anchorage, Alaska, Final report prepared 

for CononcoPhillips Alaska, Inc., Shell Exploration and 

Production Co., and StatOil USA E&P, Inc., 218 p. 

Gamito, S., 1998, Growth models and their use in ecological 

modeling—An application to a fish population: Ecological 

Modeling, v. 113, nos. 1–3, p. 83–94.

Garrett, D.L., Pietsch, T.W., Utter, F.M., and Hauser, L., 

2007, The hybrid sole, Inopsetta ischyra (Teleostei: 

Pleuronectiformes: Pleuronectidae)—Hybrid or biological 

species?: Transactions of the American Fisheries Society, 

v. 136, no. 2, p. 460–468.

Garrison, K.J., and Miller, B.S., 1982, Review of the early 

life history of Puget Sound fishes: Seattle, University of 

Washington, Fisheries Research Institute, 729 p.

Gaston, A.J., Cairns, D.K., Elliot, R.D., and Noble, D.G., 

1985, A natural history of Digges Sound: Report Series 

Canadian Wildlife Service no. 46, Canadian Government 

Publishing Centre, Canada Communication Group, 63 p.

George, C., Moulton, L.L., and Johnson, M., 2007, A field 

guide to the common fishes of the North Slope of Alaska: 

Alaska Department of Wildlife Management, North Slope 

Borough, 93 p.

Georgette, S., and Shiedt, A., 2005, Whitefish—Traditional 

ecological knowledge and subsistence fishing in the 

Kotzebue Sound, Alaska: Alaska Department of Fish and 

Game, Technical Report No. 290, 148 p.

Gieseg, S., Cuddihy, S., Hill, J.V., and Davison, W., 2000, 

A comparison of plasma vitamin C and E levels in 

two Antarctic and two temperate water fish species: 

Comparative Biochemistry and Physiology, Part B—

Biochemistry and Molecular Biology, v. 125, no. 3, 

p. 371–378.

Gillispie, J.G., Smith, R.L., Barbour, E., and Barber, W.E., 

1997, Distribution, abundance, and growth of Arctic cod 

in the Northeastern Chukchi Sea, in Reynolds, J.B., ed., 

Symposium 19, Proceedings of the Fish Ecology in Arctic 

North America Symposium, Fairbanks, Alaska, May 19–21, 

1992: American Fisheries Society, p. 81–89.

Gillman, D.V., and Kristofferson, A.H., 1984, Biological 

data on Pacific herring (Clupea harengus pallasi) from 

Tuktoyaktuk Harbour and the Liverpool Bay area, 

Northwest Territories, 1981 to 1983: Winnipeg, Manitoba, 

Canada, Canadian Data Report of Fisheries and Aquatic 

Sciences, Department of Fisheries and Oceans, Western 

Region, no. 485, 22 p.

Gjøsaeter, J., 1981, Growth, production and reproduction 

of the Myctophid fish Benthosema glaciale from western 

Norway and adjacent seas: Fiskeridirektoratets skrifter, 

Serie Havundersøkelse, v. 17, no. 3, p. 79–108.

Goddard, S.V., and Fletcher, G.L., 2002, Physiological 

ecology of antifreeze proteins—A northern perspective, in

Ewart, K.V., and Hew, C.L., eds., Molecular aspects of fish 

and marine biology, volume 1—Fish antifreeze proteins: 

Singapore, World Scientific, p. 17–60.

Goddard, S.V., Kao, M.H., and Fletcher, G.L., 1999, 

Population differences in antifreeze production cycles of 

juvenile Atlantic cod reflect adaptations to overwintering 

environment: Canadian Journal of Fisheries and Aquatic 

Sciences, v. 56, no. 11, p. 1,991–1,999.
Chapter 8 713

Goetz, F.W., McCauley, L., Goetz, G.W., and Norberg, B., 

2005, Using global genome approaches to address problems 

in cod mariculture: ICES Journal of Marine Science, v. 63, 

no. 2, p. 393–399.

Gomelyuk, V.E., 2000, Comparative analysis of everyday 

behavior and mode of the life of three species of 

Hexagrammos (Hexagrammidae, Scorpaeniformes) in 

summer: Journal of Ichthyology, v. 40, no. 1, p. 74–85.

Gong, H., Croft, K., Driedzic, W.R., and Ewart, K.V., 2011, 

Chemical chaperoning action of glycerol on the antifreeze 

protein of rainbow smelt: Journal of Thermal Biology, v. 36, 

no. 1, p. 78–83.

Gong, Z., Fletcher, G.L., and Chew, C.L., 1992, Tissue 

distribution of fish antifreeze protein mRNAs: Canadian 

Journal of Zoology, v. 70, no. 4, p. 810–814.

Grant, W.S., Bakkala, R.G., Utter, F.M., Teel, D.J., and 

Kobayashi, T., 1983, Biochemical genetic population 

structure of yellowfin sole, Limanda aspera, of the North 

Pacific Ocean and Bering Sea: Fishery Bulletin, v. 81, no. 4, 

p. 667–677.

Grebmeier, J.M., Cooper, L.W., Ashjian, C.A., Bluhm, B.A., 

Campbell, R.B., Dunton, K.E., Moore, J., Okkonen, S., 

Sheffield, G., Trefry, J., and Pasternak, S.Y., 2015, Pacific 

Marine Arctic Regional Synthesis (PacMARS) Final 

Report: North Pacific Research Board, 259 p.

Grebmeier, J.M., Cooper, L.W., Feder, H.M., and Sirenko, 

B.I., 2006a, Ecosystem dynamics of the Pacific-influenced 

North Bering and Chukchi Seas in the Amerasian Arctic: 

Progress in Oceanography, v. 71, issues 2–4, p. 331–361.

Grebmeier, J.M. and Maslowski, W., eds., 2014, The Pacific 

Arctic Region: Ecosystem status and trends in a rapidly 

changing environment, Springer Science+Business Media 

Dordrecht, 450 p.

Grebmeier, J.M., Overland, J.E., Moore, S.E., Farley, 

E.V., Carmack, E.C., Cooper, L., Frey, K.E., Helle, J.J., 

McLaughlin, F.A., and McNutt, S.L., 2006b, A major 

ecosystem shift in the northern Bering Sea: Science, v. 311, 

no. 5766, p. 1,461–1,464.

Green, J.M., 1974, A localized mass winter kill of cunners in 

Newfoundland: Canadian Field-Naturalist, v. 88, p. 96–97.

Green, J.M., and Farwell, M., 1971, Winter habits of the 

cunner, Tautogolabrus adspersus (Walbaum 1792), in 

Newfoundland: Canadian Journal of Zoology, v. 49, no. 12, 

p. 1,497–1,499.

Green, J.M., and Mitchell, L.R., 1997, Biology of the fish 

doctor, an eelpout, from Cornwallis Island, Northwest 

Territories, Canada: American Fisheries Society Symposium 

19, p. 140–174.

Green, J.M., and Steele, D.H., 1975, Observations on marine 

life beneath sea ice, Resolute Bay, N.W.T., in Proceedings 

of the Circumpolar Conference on Northern Ecology, 

September 15–18, 1975: Ottawa, Ontario, National 

Research Council of Canada, p. II-77–II-86.

Greenwald, N., Callimanis, S., Garty, A., and Peters, E., 2006, 

The Bering Sea—A biodiversity assessment of vertebrate 

species: Portland, Oreg., Center for Biological Diversity, 

Pacific Environment, 67 p. plus ACCESS database.

Gregg, J.L., Anderi, D.M., and Kimura, D.K., 2006, 

Improving the precision of otolith-based age estimates for 

Greenland halibut (Reinhardtius hippoglossoides) with 

preparation methods adapted for fragile sagittae: Fishery 

Bulletin, v. 104, no. 4, p. 643–648.

Greiner, B., 2009, Estimation of Dolly Varden and 

documentation of water and weather parameters on the 

Hulahula River, Alaska 2008: Meadville, Pennsylvania, 

Allegheny College, Department of Environmental Science, 

Senior thesis, 36 p.

Griffiths, W., Craig, P., Walder, G., and Mann, G., 1975, 

Fisheries investigations in a coastal region of the Beaufort 

Sea (Nunaluk Lagoon, Yukon), in Craig, P.C., ed., Fisheries 

investigations in a coastal region of the Beaufort Sea: 

Calgary, Alberta, Canadian Arctic Gas Study, Ltd. and 

Alaskan Arctic Gas Study Co., Arctic Gas Biological Report 

Series 34 (2), 219 p.

Griffiths, W.B., Fechhelm, R.G., Gallaway, B.J., and Fissel, 

D.B., 1988, Yukon fish and effects of causeway: Prepared 

for Indian and Northern Affairs Canada, LGL Ltd. and 

Arctic Science Ltd., 124 p. 

Griffiths, W.B., Fechhelm, R.G., Gallaway, B.J., Martin, 

L.R., and Watson, W.J., 1998, Abundance of selected fish 

species in relation to temperature and salinity patterns in the 

Sagavanirktok Delta, Alaska, following construction of the 

Endicott Causeway: Arctic, v. 51, no. 2, p. 94–104.

Griffiths, W.B., Gallaway, B.J., Gazey, W.J., and Dillinger, 

R.E., Jr., 1992, Growth and condition of Arctic Cisco and 

Broad Whitefish as indicators of causeway-induced effects 

in the Prudhoe Bay Region, Alaska: Transactions of the 

American Fisheries Society, v. 121, no. 5, p. 557–577.

Grigorev, S.S., 1995, Characteristics of flatfish eggs and larva 

samples near eastern Kamchatka and in the western Bering 

Sea during June-July 1991, in Alaska Sea Grant Program, 

eds., Proceedings of the International Symposium on North 

Pacific Flatfish: University of Alaska, Fairbanks, Alaska Sea 

Grant Program, AK-SG-95-04, p. 101–116.
714 Alaska Arctic Marine Fish Ecology Catalog

Grigorev, S.S., and Fadeev, N.S., 1995, Distribution of Alaska 

plaice, flathead sole, and Bering flounder eggs in the 

eastern Bering Sea during April–July, 1988–1990, in Alaska 

Sea Grant Program, eds., Proceeding of the International 

Symposium on North Pacific Flatfish: Fairbanks, University 

of Alaska, Alaska Sea Grant Program, AK-SG-95-04.

Gritsenko, O.F., Churikov, A.A., and Rodionova, S.S., 1984, 

The reproductive ecology of the Arctic smelt, Osmerus 

mordax dentex (Osmeridae), in the rivers of Sakhalin Island: 

Journal of Ichthyology, v. 24, no. 3, p. 23–33.

Gross, M.R., 1987, Evolution of diadromy in fines: American 

Fisheries Society Symposium, no. 1, p. 14–25.

Grossberg, R., and Cunningham, C.W., 2001, Genetic 

structure in the sea from populations to communities, chap. 

3 of Bertness M.D., Gaines, S.D., and Hay, M.E., eds., 

Marine community ecology: Sunderland, Massachusetts, 

Sinauer Associates, Inc., p. 61–84.

Gunderson, D.R., 1980, Using r-K selection theory to predict 

natural mortality: Canadian Journal of Fisheries and Aquatic 

Sciences, v. 37, no. 12, p. 2,266–2,271.

Gustafson, R.G., Lenarz, W.H., McCain, B.B., Schmitt, 

C.C., Grant, W.S., Builder, T.L., and Methot, R.D., 2000, 

Status review of Pacific hake, Pacific cod, and walleye 

pollock from Puget Sound, Washington: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum NMFSNWFSC-44. 

Haertel, L., and Osterberg, C., 1967, Ecology of zooplankton, 

benthos and fishes in the Columbia River Estuary: Ecology, 

v. 48, no. 3, p. 459–472.

Haldorson, L.J., and Craig, P., 1984, Life history and ecology 

of a Pacific-Arctic population of rainbow smelt in coastal 

waters of the Beaufort Sea: Transactions of the American 

Fisheries Society, v. 113, no. 1, p. 33–38.

Haldorson, L., Pritchett, M., Sterritt, D., and Watts, J., 1993, 

Abundance patterns of marine fish larvae during spring in a 

southeastern Alaskan bay: Fishery Bulletin, v. 91, no. 1, 

p. 36–44.

Hamada, T., 1966, Studies on fluctuations in the abundance 

of larval sand-lance in the Harima-nada and Osaka Bay 

I—Relation between the progeny abundance and the age 

composition of parent fish: Bulletin of the Japanese Society 

of Fisheries Oceanography, v. 32, no. 5, p. 393–398.

Hamazaki, T., Fair, L., Watson, L., and Brennan, E., 2005, 

Analyses of Bering Sea bottom-trawl surveys in Norton 

Sound—Absence of regime shift effects on epifauna and 

demersal fish: ICES Journal of Marine Science, v. 62, no. 8, 

p. 1,597–1,602.

Hardie, D.C., 2003, The evolutionary ecology of Arctic fishes: 

InfoNorth v. 56, no. 4, p. 430–433.

Harris, C.K., and Hartt, A.C., 1977, Assessment of pelagic and 

nearshore fish in three bays on the east and south coasts of 

Kodiak Island, Alaska: Seattle, University of Washington, 

Fisheries Research Institute, FRI-UW-7719.

Harry, G.Y., and Hartley, J.R., 1981, Northern fur seals in 

the Bering Sea, in Hood, D.W., and Calder, J.A., eds., The 

Eastern Bering Sea shelf: Oceanography and Resources: 

Seattle, University of Washington Press, p. 847–867.

Hart, J.L., 1973, Pacific fishes of Canada: Ottawa, Fisheries 

Research Board of Canada, 740 p.

Hart, P.J.B., and Reynolds, J.D., eds., 2002, Handbook of fish 

biology and fisheries—2 volume set: Blackwell Science, 

Ltd., 856 p.

Harwood, L.A., Sandstrom, S., and Linn, E., 2009, Status 

of anadromous Dolly Varden (Salvelinus malma) of the 

Rat River, Northwest Territories, as assessed through 

sampling of the subsistence fishery (1995–2007): Canadian 

Manuscript Report of Fisheries and Aquatic Sciences 2891, 

52 p.

Harley, C.D.G., Hughs, A.R., Hutgren, K.M., Miner, B.G., 

Sorte, C.J.B., Thomber, C.S., Rodriguez, L.F., Tomanek, L., 

and Williams, S.L., 2006, The impacts of climate change in 

coastal systems: Ecological Letters, v. 9, p. 228–241.

Hatch, S.A., and Sanger, G.A., 1992, Puffins as samplers of 

juvenile pollock and other forage fish in the Gulf of Alaska: 

Marine Ecology Progress Series, v. 80, p. 1–14.

Hauser, L., Spies, I., and Loher, T., 2006, Microsatellite 

screening in Pacific halibut (Hippoglossus stenolepis) and 

a preliminary examination of population structure based on 

observed DNA variation: Seattle, Washington, International 

Pacific Halibut Commission Scientific Report, no. 81.

Hayes, D.L., and Kuletz, K.J., 1997, Decline of pigeon 

guillemot populations in Prince William Sound, Alaska, 

and apparent changes in distribution and abundance of their 

prey, in Forage fishes in marine ecosystems—Proceedings 

of the International Symposium on the Role of Forage 

Fishes in Marine Ecosystems: University of Alaska, 

Fairbanks, Alaska Sea Grant College Program, p. 699–702.

Healey, M.C., 1991, Life history of Chinook salmon 

(Oncorhynchus tshawytscha), in Groot, C., and Margolis, 

L., eds., Pacific salmon life histories, Vancouver, British 

Columbia, University of British Columbia Press, 

p. 313–393.

Heard, W.R., 1991, Life history of pink salmon 

(Oncorhynchus gorbuscha), in Groot, C., and Margolis, 

L., eds., Pacific salmon life histories: Vancouver, British 

Columbia, UBC Press, p. 121–230.
Chapter 8 715

Hebert P.D.N., and Wearing-Wilde J., eds., 2002, Canada’s 

Polar Life: Web page, CyberNatural Software, University of 

Guelph, accessed October 15, 2013, at http://www.polarlife.

ca.

Heikkinen, R.K., Luoto, M., Araújo, M.B., Virkkala, 

R.V., Thuiller, W., and Sykes, M.T., 2006, Methods and 

uncertainties in bioclimatic envelope modelling under 

climate change: Progress in Physical Geography, v. 30, 

no. 6, p. 1–17.

Hershberger, P.K., Pacheco, C.A., Gregg, J.L., Purcell, 

M.K., and LaPatra, S.E., 2008, Differential survival of 

Ichthyophonus isolates indicates parasite adaptation to its 

host environment: Journal of Parasitology, v. 94, no. 5, 

p. 1,055–1,059.

Hershberger, P., Rhodes, L., Kurath, K., and Winton, J., 2013, 

Infectious diseases of fishes in the Salish Sea: Fisheries, 

v. 38, no. 9, p. 402–409.

Hew, C.L., Fletcher, G.L., and Ananthanarayanan, V.S., 

1980, Antifreeze proteins from the shorthorn sculpin, 

Myoxocephalus scorpius: isolation and characterization: 

Canadian Journal of Biochemistry, v. 58, no. 5, p. 377–383.

Hew, C.L., Wang, N.-C., Joshi, S., Fletcher, G.L., Scott, G.K., 

Hayes, P.H., Buettner, B., and Davies, P.L., 1988, Multiple 

genes provide the basis for antifreeze protein diversity and 

dosage in ocean pout, Macrozoarces americanus: Journal of 

Biological Chemistry v. 263, p. 12,049–12,055.

Hildebrandt, N., Bergmann, M., and Knust, R., 2011, 

Longevity and growth efficiency of two deep-dwelling 

Arctic zoarcids and comparison with eight other zoarcid 

species from different climatic regions: Polar Biology, v. 34, 

no. 10, p. 1,523–1,533.

Hill, J.V., and Egginton, S., 2010, Control of branchial artery 

tone in fish—Effects of environmental temperature and 

phylogeny: Physiological Biochemistry and Zoology, v. 83, 

no. 1, p. 33–42.

Hjorth, J.P., 1974, Molecular and genetic structure of 

multiple hemoglobins in the eelpout, Zoarces viviparous L: 

Biochemical Genetics, v. 13, no. 5–6, p. 379–391.

Hobson, E.S.,1986, Predation on the Pacific sand lance, 

Ammodytes hexapterus (Pisces: Ammodytidae), during the 

transition between day and night in southeastern Alaska: 

Copeia, v. 1986, no. 1, p. 223–226.

Hoff, G.R., 2006, Biodiversity as an index of regime shift in 

the eastern Bering Sea: Fishery Bulletin, v. 104, no. 2, 

p. 226–237.

Hoff, G.R., and Britt, L.L., 2003, The 2002 Eastern Bering 

Sea upper continental slope survey of groundfish and 

invertebrate resources: U.S. Department of Commerce, 

National Oceanic and Atmospheric Administration 

Technical Memorandum, NMFS-AFSC-141, Department of 

Commerce, 286 p.

Hoff, G.R., Stevenson, D.E., and Orr, J.W., 2015, Guide 

to the gadiform fishes of the eastern North Pacific: 

U.S. Department of Commerce, National Oceanic and 

Atmospheric Aadministration Technical Memorandum 

NMFS-AFSC-309, 68 p. doi:10.7289/V5/TM-AFSC-309, 

available at http://www.afsc.noaa.gov/Publications/

AFSC-TM/NOAA-TM-AFSC-309.pdf.

Hoff, G.R., and Britt, L.L., 2005, Results of the 2004 Eastern 

Bering Sea upper continental slope survey of groundfish and 

invertebrate resources: National Oceanic and Atmospheric 

Administration Technical Memorandum, 300 p.

Hognestad, P.T., 1968, Observations on polar cod in the 

Barents Sea: Rapports et procès-verbaux des reunions: 

Conseil permanent international pour l’exploration de la 

mer., v. 158, p. 126–130.

Holeton, G.F., 1974, Metabolic cold adaptation of polar fish—

Fact or artefact?: Physiological Zoology, v. 47, no. 3, 

p. 137–152.

Holladay, B.A., and Norcross, B.L., 1995, Diet diversity 

as a mechanism for partitioning nursery grounds of 

pleuronectids: Proceedings of the International Symposium 

on North Pacific Flatfish, Alaska Sea Grant AK-SG-95-04, 

p. 177–203.

Holland-Bartels, L., and Pierce, B., eds., 2011, An evaluation 

of the science needs to inform decisions on Outer 

Continental Shelf energy development in the Chukchi and 

Beaufort Seas, Alaska: U.S. Geological Survey Circular 

1370, 278 p.

Hollowed, A.B., Planque, B., and Loeng, H., 2013, Potential 

movement of fish and shellfish stocks from the sub-Arctic to 

the Arctic Ocean: Fisheries Oceanography, v. 22, no. 5, 

p. 355–370.

Holmes, W.N., and Donaldson, E.M., 1969, The body 

compartments and distribution of electrolytes: in Hoar, W.S. 

and Randall, D.J., eds., Fish physiology, volume 1: New 

York, Academic Press, p. 1–89,

Hop, H., and Graham, M., 1995, Respiration of juvenile 

Arctic cod (Boreogadus saida)—Effects of acclimation, 

temperature, and food intake: Polar Biology, v. 15, no. 5, 

p. 359–367.
716 Alaska Arctic Marine Fish Ecology Catalog

Hop, H., Welch, H.E., and Crawford, R.E., 1997, Population 

structure and feeding ecology of Arctic cod schools in 

the Canadian High Arctic: American Fisheries Society 

Symposium 19, p. 13–39.

Hopky, G.E., and Ratynski, R.A., 1983, Relative abundance, 

spatial and temporal distribution, age and growth of fishes 

in Tuktoyaktuk Harbour, N.W.T., 28 June to 5 September, 

1981: Canadian Manuscript Report of Fisheries and Aquatic 

Sciences 1983, 77 p.

Hopcroft, R., Bluhm, B., and Gradinger, R., eds., 2008, Arctic 

Ocean synthesis—Analysis of climate change impacts in 

the Chukchi and Beaufort Seas with strategies for future 

research: Fairbanks, University of Alaska, 184 p.

Houston, J., and McAllister, D.E., 1990a, Status of the Bering 

wolfish, Anarhichas orientalis, in Canada: The Canadian 

Field-Naturalist, v. 104, no. 1, p. 20–23.

Houston, J., and McAllister, D.E., 1990b, Status of the 

blackline prickleback, Acantholumpenus mackayi, in 

Canada: The Canadian Field-Naturalist, v. 104, no. 1, 

p. 24–28.

Howard, E.I., Blakeley, M.P., Haertlein, M., Petit-Haertlein, 

I., Mitschler, A., Fisher, S.J., Cousido-Siah, A., Salvay, 

A.G., Popov, A., Muller-Dieckmann, C., Petrova, T., and 

Podjarny, A., 2010, Neutron structure of type-III antifreeze 

protein allows the reconstruction of AFP-ice interface: 

Journal of Molecular Recognition, v. 24, no. 4, p. 724–732.

Howland, K.L., Gendron, M., Tonn, W.M., and Tallman, R.F., 

2004, Age determination of a long-lived coregonid from 

the Canadian North—Comparison of otoliths, fin rays and 

scales in inconnu (Stenodus leucichthys): Annales Zoologici 

Fennici, Finnish Zoological and Botanical Publishing 

Board, p. 205–214.

Hufford, G.L., 1974, WEBSEC 71-72—An ecological survey 

in the Beaufort Sea, August–September 1971–1972: U.S. 

Coast Guard Oceanographic Unit report No. CG 373-64, 

268 p.

Hunt, B.M., Hoefling, K., and Cheng, C.-H. C., 2003, Annual 

warming episodes in seawater temperatures in McMurdo 

Sound in relationship to endogenous ice in notothenioid 

fish: Antarctic Science, v. 15, no. 3, p. 333–338.

Hunt G.L. Jr., and McKinnell, S., 2006, Interplay between 

top-down, bottom-up, and wasp-waist control in marine 

ecosystems: Progress in Oceanography, v. 68, no. 2–4, 

p. 115–124

Hunt, J.L., Jr., Blanchard, A.L., Boveng, Peter, Dalpadado, 

Padmini, Drinkwater, K.F., Eisner, Lisa, Hopcroft, R.R., 

Kovacs, K.M., Norcross, B.L., Renaud, Paul, Reigstad, 

Marit, Renner, Martin, Skjoldal, H.R., Whitehouse, Andy, 

and Woodgate, R.A, 2013, The Barents and Chukchi Seas—

Comparison of two Arctic shelf ecosystems: Journal of 

Marine Systems, v. 109–110, p. 43–68. 

Huntington, H.P., 2000, Traditional knowledge of the ecology 

of belugas, Delphinapterus leucas, in Cook Inlet, Alaska: 

Marine Fisheries Review, v. 62, p. 134–140.

Huse, G., and Ellingsen, I., 2008, Capelin migrations and 

climate change—A modeling analysis: Climatic Change, 

v. 87, no. 1–2, p. 177–197.

Ianelli, J.N., Wilderbuer, T.K., and Nichol, Dan, 2007, 

Assessment of Greenland turbot in the eastern Bering Sea 

and Aleutian Islands: North Pacific Fisheries Management 

Council, Bering Sea and Aleutian Islands SAFE Document, 

p. 513–562, accessed April 6, 2014, at http://www.afsc.

noaa.gov/refm/docs/2007/BSAIturbot.pdf.

Imsland, A.K., Foss, A., Naevdal, G., Johansen, T., Folkvord, 

A., Stefansson, S.O., and Jonassen, T.M., 2004, Variations 

in growth in haemoglobin genotypes of Atlantic cod: Fish 

Physiology and Biochemistry, v. 30, no. 1, p. 47–55.

International Pacific Halibut Commission, 1998, The Pacific 

halibut—Biology, fishery, and management: Seattle, 

Washington, International Pacific Halibut Commission 

Technical Report, no. 40.

Irvine, J.R., Macdonald, R.W., Brown, R.J., Godbout, L., 

Reist, J.D., and Carmack, E.C., 2009, Salmon in the Arctic 

and how they avoid lethal low temperatures: North Pacific 

Anadromous Fish Commission, Bulletin no. 5, p. 39–50.

Irvine, G., and Meyer, R., 1989, Overview of the fishes 

occurring in the Alaska Arctic, in Meyer, R.M., 

and Johnson, T.M., eds., Fishery oceanography—A 

comprehensive formulation of technical objectives for 

offshore application in the Arctic: Anchorage, Alaska, U.S. 

Department of the Interior, Minerals Management Service, 

OCS Study MMS 88-0042. p. 6–25.

Isakson, J.S., Houghton, J.P., Rogers, D.E., and Parker, S.S., 

1986, Fish use of inshore habitats north of the Alaska 

Peninsula, June–September 1984 and June–July 1985—

Final report: U.S. Department of the Interior, Minerals 

Management Service, Alaska OCS Region, OCS EIS/EA, 

380 p.

Jakobsson, M., Polyak, L., Edwards, M., Kleman, J., 

and Coakley, B., 2008, Glacial geomorphology of the 

Central Arctic Ocean—The Chukchi Borderland and the 

Lomonosov Ridge: Earth Surface Processes and Landforms, 

v. 33, p. 526–545.
Chapter 8 717

Jarvela, L.E., and Thorsteinson, L.K., 1999, The epipelagic 

fish community of Beaufort Sea coastal waters, Alaska: 

Arctic, v. 52, no. 1, p. 80–94.

Jay, C.V., Marcot, B.G., and Douglas, D.C., 2011, Projected 

status of the Pacific walrus (Odobenus rosmarus divergenes) 

in the twenty-first century: Polar Biology, v. 34, no. 7, 

p. 1,065–1,084.

Jensen, A.S., 1948, A contribution to the ichthyofauna of 

Greenland: Spoila Zoologica Musei Hauniensis, v. 9, 

p. 27–57.

Jewett, S.C., and Feder, H.M., 1980, Autumn food of adult 

starry flounders, Platichthys stellatus, from the northeastern 

Bering Sea and the southeastern Chukchi Sea: Journal de 

Conseil International pour l’Exploration de la Mer., v. 39, 

no. 1, p. 7–14.

Johnson, L., 1997, Living with uncertainty, in Reynolds, 

J., ed., Fish ecology in Arctic North America: American 

Fisheries Society Symposium 19, p. 340–345.

Johnson, M.L., Fiscus, C.H., Ostenson, B.T., and Barbour, 

M.L., 1966, Marine mammals, in Wilimovsky, N.J., and 

Wolfe, J.N., eds., Environment of the Cape Thompson 

Region, Alaska: Oak Ridge, Tennessee, United States 

Atomic Energy Commission, Division of Technical 

Information, p. 877–924.

Johnson, S.W, Neff, A.D., Lindeberg, J.F., and Maselko, J.M., 

2012, Atlas of nearshore fishes of Alaska—A synthesis 

of marine surveys from 1998 to 2011: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration, Technical Memorandum NMFS-AFSC-239, 

261 p.

Johnson, S.W., Neff, A.D., and Thedinga, J.F., 2005, An atlas 

on the distribution and habitat of common fishes in shallow 

nearshore waters of southeastern Alaska: Alaska Fisheries 

Science Center, Technical Memorandum NMFS-AFSC-157, 

98 p.

Johnson, S.W., Thedinga, J.F., and Lindeberg, M.R., 2012, 

Nearshore fish atlas of Alaska: National Oceanic and 

Atmospheric Administration Fisheries, accessed February 

2012 at http://www.fakr.noaa.gov/habitat/fishatlas/. 

Johnson, S.W., Thedinga, J.F., Neff, A.D., and Hoffman, C.A., 

2010, Fish fauna in nearshore waters of a Barrier Island 

in the western Beaufort Sea, Alaska: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration, Technical Memorandum, NMFSAFSC-210, 

28 p.

Johnson, T.P., and Johnston, I.A., 1991, Temperature 

adaptation and the contractile properties of live muscle 

fibres from teleosts fish: Journal of Comparative Physiology 

B, v. 161, no. 1, p. 27–36.

Johnston, I.A., 1985, Sustained force development—

Specializations and variation among the vertebrates: Journal 

of Experimental Biology, v. 115, p. 239–251.

Johnston, I.A., and Altringham, J.D., 1985, Evolutionary 

adaptation of muscle power output to environmental 

temperature—Force-velocity characteristics of skinned 

fibres isolated from Antarctic, temperate and tropical marine 

fish: Pflügers Archiv, v. 405, no. 2, p. 136–140.

Johnston, I.A., and Battram, J., 1993, Feeding energetics 

and metabolism in demersal fish species from Antarctic, 

temperate and tropical environments: Marine Biology, 

v. 115, no. 1, p. 7–14. 

Johnston, I.A., Calvo, J., Guderley, H., Fernandez, D., and 

Palmer, L., 1998, Latitudinal variation in the abundance and 

oxidative capacities of muscle mitochondria in perciform 

fishes: The Journal of Experimental Biology, v. 201, no. 1, 

p. 1–12.

Jones, M.L., and Den Beste, J., 1977, Tuft Point and adjacent 

coastal areas fisheries projects: Calgary, Alberta, Aquatic 

Environments Ltd., 152 p.

Jordan, D.S., 1891, Relations of temperature to vertebrae 

among fishes: Proceedings of The United States National 

Museum: v. 14, no. 845, p. 107–120.

Jordan, D.S., and Evermann, B.W., 1896, The fishes of North 

and Middle America—A descriptive catalogue of the 

species of fish-like vertebrates found in the waters of North 

America, North of the Isthmus of Panama, part 1: Bulletin 

of the United States National Museum, no. 47, 1,240 p.

Jørgensen, O.A., 1997, Pelagic occurrence of Greenland 

halibut, Reinhardtius hippoglossoides (Walbaum), in west 

Greenland waters: Journal of Northwest Atlantic Fisheries 

Science, v. 21, p. 39–50.

Junquera, S., Román, E., Morgan, J., Sainza, M., and Ramilo, 

G., 2003, Time scale of ovarian maturation in Greenland 

halibut (Reinhardtius hippoglossoides, Walbaum): ICES 

Journal of Marine Science, v. 60, no. 4, p. 767–773.

Kajimura, H., and Loughlin, T.R., 1988, Marine mammals 

in the oceanic food web of the eastern subarctic Pacific: 

University of Tokyo, Bulletin of the Ocean Research 

Institute, v. 26, no. 2, p. 187–233.

Karamushko, L.I., Shatunovskii, M.I., and Christiansen, I. Sh., 

2004, Metabolic rate and metabolic adaptations in fishes of 

different latitudes: Journal of Ichthyology, v. 44, 

p. 647–654.
718 Alaska Arctic Marine Fish Ecology Catalog

Karpenko, V.I., and Vasilets, P.M., 1996, Biology of smelt 

(Osmeridae) in the Korf-Karagin coastal area of the 

southwestern Bering Sea, in Mathisen, O.A., and Coyle, 

K.O., eds., Ecology of the Bering Sea—A review of Russian 

literature: Fairbanks, University of Alaska, Alaska Sea 

Grant Program, AK-SG-96-01, p. 217–235.

Karpov, A.K., and Novikov, G.G., 1981, Hemoglobin 

alloforms in cod, Gadus morhua (Gadiformes, Gadidae), 

their functional characteristics and occurrence in 

populations: Journal of Ichthyology, v. 6, p. 45–49.

Kartavtsev, Y.P., Sharina, S.N., Goto, T., Chichvarkhin, 

A.Y., Balanov, A.A., Vinnikoc, K.A., Invankov, V.N., and 

Hanzawa, N., 2008, Cytochrome oxidase 1 gene sequence 

analysis in six flatfish species (Teleostei Pleuronectidae) 

of Far East Russia with inferences in phylogeny and 

taxonomy: DNA Sequence v. 19, no. 6, p. 479–489.

Katsanevakis, S., 2006, Modelling fish growth—Model 

selection, multi-modal inference and model selection 

uncertainty: Fisheries Research v. 81, p. 229–235.

Kawasaki, T., 1980, Fundamental relations among the 

selections of life history in marine teleosts: Bulletin of the 

Japanese Society of Scientific Fisheries, v. 46, no. 3, 

p. 289–293.

Kendel, R.E., Johnston, R.A.C., Kozak, M.D., and Lobsiger, 

U., 1974, Movements, distribution, populations and 

food habits of fish in the western coastal Beaufort Sea: 

Department of the Environment, Canada, Interim Report of 

Beaufort Sea Project, Study B1, 64 p.

Kendel, R.E., Johnston, R.A.C., Lobsiger, U., and Kozak, 

M.D., 1975, Fishes of the Yukon coast: Victoria, British 

Columbia, Department of the Environment (Canada), 

Beaufort Sea Project, Technical Report 6, 114 p.

Kennett, J.P., 1982, Marine geology: Englewood Cliffs, New 

Jersey, Prentice-Hall, 813 p.

Ketchen, K.S., 1975, Age and growth of dogfish Squalus 

acanthias in British Columbia waters: Journal of the 

Fisheries Research Board of Canada, v. 32, no. 1, p. 43–59.

Kim, S.T., 2001, Winter migrations of shelf fish to the 

continental slope: Journal of Ichthyology, v. 41, no. 8, 

p. 564–574.

Kim, S.T., and Biryukov, I.A., 1998, Distribution and some 

features of the biology of Reinhardtius hippoglossoides 

matsuurae (Pleuronectidae) and Sebastolobus macrochir

(Scorpaenidae) from the eastern coast of Sakhalin: Journal 

of Ichthyology, v. 38, no. 1, p. 143–146.

Kim, S.T., Biryukov, I.A., and Pometeev, E.V., 1999, Species 

composition, distribution, and structural changes in the 

flatfish community of the Tatarskii Strait during the fishing 

period 1994–1997: Journal of Ichthyology, v. 39, no. 6, 

p. 433–440.

Kim, S.T., and Shepeleva, O.N., 2001, The structure of shelf 

ichthyocenoses of northeastern Sakhalin and Terpeniya Bay: 

Journal of Ichthyology, v. 41, no. 9, p. 711–722.

Kinder, T.H., and Schumacher, J.D., 1981, Circulation 

over the continental shelf of the southeastern Bering 

Sea, in Hood, D.W., and Calder, J.A., eds., The Eastern 

Bering Sea shelf—Oceanography and resources, vol. 1: 

U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, Office of Marine Pollution 

Assessment, p. 53–75.

King, J.R., and McFarlane, G.A., 2003, Marine fish life history 

strategies—Applications to fishery management: Fisheries 

Management and Ecology, v. 10, no. 4, p. 249–264.

Knipowitsch, N., 1906, Ichthyologische Untersuchungen 

im Eismeer, I. Lycodes und Lycenchelys: Mémoires de 

l’Academie Imperiale des Sciences de Saint Petersbourg, 

v. 19, no. 1, p. 1–130, 1 pl.

Kobelev, E.A., 1989, Some biological characteristics of the 

Arctic flounder, Liopsetta glacialis, in the southeastern area 

of the Barents Sea: Journal of Ichthyology, v. 29, no. 6, 

p. 16–21.

Kodolov, L.S., and Matveychuk, S.P., 1995, Stock condition of 

Greenland turbot (Reinhardtius hippoglossoides matsuurae

Jordan et Snyder) in the northwestern Bering Sea: 

Anchorage, Alaska Sea Grant Program AK-SG-95-04.

Kolpakov, N.V., 2005a, Diversity and seasonal dynamics of 

ichthyocenosis of the Circumlittoral of Russkaya Bight 

(Northern Primor’s): Journal of Ichthyology, v. 45, no. 9, 

p. 744–753.

Kolpakov, N.V., 2005b, On the biology of Platichthys stellatus

(Pleuronectidae) in coastal waters of northern Primorye: 

Journal of Ichthyology, v. 45, no. 8, p. 594–606.

Kostow, K., 2002, Oregon lampreys—Natural history, status, 

and analysis of management issues—Information reports: 

Corvallis, Oregon Department of Fish and Wildlife, Oregon 

State University, Klamath Waters Digital Library, accessed 

at http://digitallib.oit.edu/cdm/ref/collection/kwl/id/3483. 

Knight, C.A., Cheng, C.C., and DeVries, A.L., 1991, 

Adsorption of alpha-helical antifreeze peptides on specific 

ice crystal surface planes: Biophysical Journal, v. 59, no. 2, 

p. 409–418.

Krogh, A., 1916, The respiratory exchange of animals and 

man: London, Longmans, 173 p.
Chapter 8 719

Krylov, A.A., Andreeva, I.A., Vogt, C., Backman, J., 

Krupskaya,V.V., Grikurov, G.E., and others, 2008, A shift 

in heavy and clay mineral provenance indicates a middle 

Miocene onset of a perennial sea ice cover in the Arctic 

Ocean: Paleoceanography 23, PA1S06.

Kucheryavyi, A.V., Savvaitova, K.A., Gruzdeva, M.A., 

and Pavlov, D.S., 2007, Sexual dimorphism and some 

special traits of spawning behavior of the Arctic lamprey 

Lethenteron camtschaticum: Journal of Ichthyology, v. 47, 

no. 7, p. 481–485.

Kucheryavyi, A.V., Savvaitova, K.A., Pavlov, D.S., Gruzdeva, 

M.A., Kuzishchin, K.V., and Stanford, J.A., 2007, 

Variations of life history strategy of the Arctic lamprey 

Lethenteron camtschaticum from the Utkholok River 

(Western Kamchatka): Journal of Ichthyology, v. 47, no. 1, 

p. 37–52.

Kulka, D., Hood, C., and Huntington, J., 2007, Recovery 

strategy for northern wolffish (Anarhichas denticulatus) 

and spotted wolffish (Anarhichas minor), and management 

plan for Atlantic Wolffish (Anarhichas lupus) in Canada: St. 

John’s, Fisheries and Oceans Canada, Newfoundland and 

Labrador Region, 103 p.

Kupriyanov, S.V., 1996, Distribution and biological indices 

of yellowfin sole (Pleuronectes asper) in the southwestern 

Bering Sea, in Mathisen, O.A., and Coyle, K.O., eds., 

Ecology of the Bering Sea—A review of Russian literature: 

Fairbanks, University of Alaska, Alaska Sea Grant Report 

no. 96-01, 306 p.

Lacho, G., 1991, Stomach content analyses of fishes from 

Tuktoyaktuk Harbour, N.W.T., 1981: Winnipeg, Manitoba, 

Canadian Data Report of Fisheries and Aquatic Sciences, 

Central and Arctic Region, Department of Fisheries and 

Oceans, no. 853, 15 p.

Laevastu, T., Marasco, R., Bax, N., Honkalehto, T., Fredin, 

R., Fukuhara, F., Gallagher, A., Ingraham, J., Livingston, 

P., Miyahara, R., and Pola, N., 1985, Evaluation of the 

effects of oil development on commercial fisheries in the 

eastern Bering Sea: Anchorage, Alaska, U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration, Outer Continental Shelf Environmental 

Assessment Program, Final Report for Research Unit 643, 

47 p.

Lang, G.M., Livingston, P.A., and Dodd, K., 2005, Groundfish 

food habits and predation on commercially important 

prey species in the eastern Bering Sea from 1997 through 

2001: U.S. Department of Commerce, National Oceanic 

and Atmospheric Administration Technical Memorandum 

NMFS-AFSC-158, 249 p.

Lang, G.M., Livingston, P.A., Pacunski, R.E., Parkhurst, 

J., and Yang, M.-S., 1991, Groundfish food habits and 

predation on commercially important prey species in the 

eastern Bering Sea from 1984 to 1986: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration Technical Memorandum NMFS F/NWC207, 240 p.

Lannig, G., Eckerle, L.G., Serendero, I., Sartoris, F.-J., 

Fischer, T., Knust, R., Johansen, T., and Pörtner, H.-O., 

2003, Temperature adaptation in eurythermal cod (Gadus 

morhua)—A comparison of mitochondrial enzyme 

capacities in boreal and Arctic populations: Marine Biology, 

v. 142, no. 3, p. 589–599.

Lawrence, M.J., Lacho, G., and Davies, S., 1984, A survey 

of the coastal fishes of the southeastern Beaufort Sea: 

Canadian Technical Report of Fisheries and Aquatic 

Sciences, no. 1220, 178 p.

Lecklin, T., Nash, G., and Egginton, S., 1995, Do fish 

acclimated to low temperature improve microcirculatory 

perfusion by adapting red cell rheology: Journal of 

Experimental Biology, v. 198, no. 8, p. 1,801–1,808.

Leong, H., Brander, K., Carmack, E., Denisenko, S., 

Drinkwater, K., Hansen, B., Kovacs, K., Livingston, P., 

McLaughlin, F., and Sakshaug, E., 2005, Marine systems, 

in Arctic climate impact assessment: Cambridge, United 

Kingdom, Cambridge University Press, p. 453–538.

Levin, P.S., Fogarty, M.J., Murawski, S.A., and Fluharty, D., 

2009, Integrated ecosystem assessments—Developing the 

scientific basis for ecosystem-based management of the 

ocean: PLoS ONE, v. 7, no. 1, p. 23–28.

LGL Limited, 2014, LGL and Greeneridge publications on 

fish: LGL Limited Web page, accessed May 15, 2014, at 

http://lgl.com/en/fish.

Lindeman, R.L., 1942, The trophic-dynamic aspect of 

ecology: Ecology, v. 23, p. 399–418.

Litzow, M.A., 2007, Alaska Fisheries Science Center quarterly 

research report, January–March, 2007: National Oceanic 

and Atmospheric Administration Fisheries, Alaska Fisheries 

Science Center Web page, accessed April 2, 2014, at http://

www.afsc.noaa.gov/Quarterly/jfm2007/divrptsRACE4.htm.

Litzow, M.A., Piatt, J.F., Prichard, A.K., and Roby, D.D., 

2002, Response of pigeon gullemots to variable abundance 

of high-lipid and low-lipid prey: Oecologia, v. 132, no. 2, 

p. 286–295.
720 Alaska Arctic Marine Fish Ecology Catalog

Livingston, P.A., Ward, A., Lang, G.M., and Yang, 

M.-S., 1993, Groundfish food habits and predation on 

commercially important prey species in the eastern Bering 

Sea from 1987 to 1989: National Oceanic and Atmospheric 

Administration Technical Memorandum NMFS-AFSC-11, 

202 p.

Logerwell, E., Busby, M., Carothers, C., Cotton, S., 

Duffy-Anderson, J., Farley, E., Goddard, P., Heintz, R., 

Holladay, B., Horne, J., Johnson, S., Lauth, B., Moulton, 

L., Neff, D., Norcross, B., Parker-Stetter, S., Seigle, J., 

and Sformo, T., 2015, Fish communities across a spectrum 

of habitats in the western Beaufort Sea and Chukchi Sea: 

Progress in Oceanography, no. 136, p. 115–132.

Logerwell, E., Rand, K., Parker-Setter, S., Horne, J., 

Weingartner, T., and Bluhm, B., 2009, Beaufort Sea marine 

fish monitoring 2008—Pilot survey and test of hypotheses: 

Anchorage, Alaska, Final Report from U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration/National Marine Fisheries Service to U.S. 

Department of the Interior, Minerals Management Service, 

Alaska OCS Region, OCE Study MMS 2008-062, 258 p.

Logerwell, E., Rand, K., and Weingartner, T.J., 2011, 

Oceanographic characteristics of the habitat of benthic fish 

and invertebrates in the Beaufort Sea: Polar Biology, v. 34, 

p. 1,783–1,796.

Loher, T., 2012, Potential mechanisms, research avenues, 

and management action associated with size at age and 

growth of Pacific halibut, in IPHC Report of Assessment 

and Research Activities 2012: International Pacific Halibut 

Commission (IPHC) online report, p. 457–486, accessed 

February 16, 2014, at http://www.iphc.int/publications/

rara/2012/rara2012457_sizeatage.pdf.

Loher, T., and Seitz, A.C., 2006, Seasonal migration and 

environmental conditions of Pacific halibut Hippoglossus 

stenolepis, elucidated from pop-up archival transmitting 

(PAT) tags: Marine Ecology Progress Series, v. 317, 

p. 259–271.

Longshan, L., Yongjun, C., Yungchih, L., Jing, Z., Punqing, 

S., Xingguang, Y., Risheng, W., and Kwang-tsao, S., 2014, 

Composition of fish species in the Bering and Chukchi 

Seas and their responses to changes in the ecological 

environment, Acta Oceanologica Sinica, v. 33, no. 6, 

p. 63-73.

Lourie, S.A., and Vincent, A.C.J., 2004, Using biogeography 

to help set priorities in marine conservation: Conservation 

Biology, v. 18, no. 4, p. 1,004–1,020.

Love, M.S., 2011, Certainly more than you wanted to know 

about the fishes of the Pacific coast: Santa Barbara, Calif., 

Really Big Press, 649 p.

Love, M.S., Mecklenburg, C.W., Mecklenburg, T.A., and 

Thorsteinson, L.K., 2005, Resource inventory of marine and 

estuarine fishes of the West Coast and Alaska—A checklist 

of North Pacific and Arctic Ocean species from Baja 

California to the Alaska-Yukon border: U.S. Geological 

Survey, OCS Study MMS 2005-030 and USGS/NBII 2005-

001, 276 p.

Low, W.-K., Lin, Q., Ewart, K.V., Fletcher, G. L., and Hew, 

C.L., 2002, The skin-type antifreeze polypeptides—A new 

class of Type I AFPs, in Ewart, K.V., and Hew, C.L., eds., 

Molecular aspects of fish and marine biology, volume 1—

Fish antifreeze proteins: New Jersey, World Scientific, 

p. 161–186.

Lowry, L.F., and Frost, K.J., 1981, Distribution, growth, and 

foods of Arctic cod (Boreogadus saida) in the Bering, 

Chukchi, and Beaufort Seas: Canadian Field-Naturalist, 

v. 95, no. 2, p. 186–191.

Lowry, L.F., Frost, K.J., and Burns, J.J., 1980, Variability in 

the diet of ringed seals, Phoca hispida, in Alaska: Canadian 

Journal of Fisheries and Aquatic Sciences, v. 37, no. 12, p. 

2,254–2,261.

Luksenburg, J.A., Pedersen, T., and Falk-Petersen, I.-B., 2004, 

Reproduction of the shorthorn sculpin Myoxocephalus 

scorpius in northern Norway: Journal of Sea Research, 

v. 51, no. 2, p. 157–166.

Luzier, C.W., Schaller, H.A., Brostrom, J.K., Cook-Tabor, 

C., Goodman, D.H., Nelle, R.D., and others, 2011, Pacific 

Lamprey (Entosphenus tridentatus)—Assessment and 

template for conservation measures: U.S. Fish and Wildlife 

Service, 282 p.

Lynghammar, A., Christiansen, J.S., Mecklenburg, C.W., 

Karamushko, O.V., Møller, P.R., Gallucci, V.F., and 

Moller, P.R., 2012, Species richness and distribution of 

chondrichthyan fishes in the Arctic Ocean and adjacent seas: 

Biodiversity, v. 14, no. 1, p. 57–66.

Macdonald, J.A., 1981, Temperature compensation in the 

peripheral nervous systems—Antarctic vs temperate 

poikilotherms: Journal of Comparative Physiology, v. 142, 

p. 411–418.

Macdonald, J.A., and Montgomery, J.C., 1982, Thermal limits 

of neuromuscular function in an Antarctic fish: Journal of 

Comparative Physiology, v. 147, p. 237–250.

Macdonald, J.A., and Montgomery, J., 2005, The nervous 

system, in Farrell, A.P., and Steffensen, J.F., eds., The 

physiology of polar fishes: Amsterdam, Elsevier, 

p. 351–383.
Chapter 8 721

Magdanz, J.S., Smith, H., Braem, N., Fox, P., and Koster, 

D.S., 2011, Patterns and trends in subsistence fish harvests, 

Northwest Alaska, 1994–2004: Alaska Department of Fish 

and Game, Technical Paper no. 366, 104 p.

Majewski, A.R., Sareault, J.E., and Reist, J., 2006, Fish catch 

data from offshore sites in the Mackenzie River estuary and 

Beaufort Sea during the open water season, August 2004 

aboard the CCGS Nahidik: Winnipeg, Manitoba, Fisheries 

and Oceans Canada, no. 2771, 42 p.

Maksimenkov, V.V., and Tokranov, A.M., 1993, The diet of 

the pond smelt, Hypomesus olidus, in the Bol’shaya River 

Estuary (Western Kamchatka): Journal of Ichthyology, 

v. 33, no. 9, p. 11–21.

Mann, G., 1974, Life history types of the least cisco 

(Coregonus sardinella, Valenciennes) in the Yukon 

Territory, North Slope and eastern Mackenzie River Delta 

drainages, in McCart, P.J., ed., Life histories of three species 

of freshwater fishes in Beaufort Sea drainages, Yukon 

Territory,: Arctic Gas Biological Report Series, v. 18, no. 3, 

132 p.

Markon, C.J., Trainor, S.F., and Chapin, F.S., III, eds., 2012, 

The United States National Climate Assessment—Alaska 

Technical Regional Report: U.S. Geological Survey 

Circular 1379, 148 p.

Marliave, J.B., 1975, The behavioral transformation from the 

planktonic larval stage of some marine fishes reared in the 

laboratory: Vancouver, University of British Columbia, 

Ph.D. dissertation.

Marshall, N.B. 1953, Egg size in Arctic, Antarctic and deepsea fishes: Evolution, v. 7, p. 328–341.

Martin, D.J., Glass, D.R., Whitmus, C.J., Simenstad, C.A., 

Milward, D.A., Volk, E.C., Stevenson, M.L., Nunes, P., 

Savoie, M., and Grotefendt, R.A., 1986, Distribution, 

seasonal abundance, and feeding dependencies of 

juvenile salmon and non-salmonid fishes in the Yukon 

River Delta: Outer Continental Shelf Environmental 

Assessment Program, Reports of Principal Investigators, 

U.S. Department of Commerce and U.S. Department of the 

Interior, 388 p.

Martin, M.H., and Clausen, D.M., 1995, Data report—1993 

Gulf of Alaska Bottom Trawl Survey: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration, NOAA Technical Memorandum, NMFSAFSC-59, 217 p.

Marz, S., 2010, Arctic sea ice ecosystem—A summary of 

species that depend on and associate with sea ice and 

projected impacts from sea ice changes: Report to the Arctic 

Council Working Group on the Conservation of Arctic Flora 

and Fauna, 64 p.

Maslenikov, K.P., Orr, J.W., and Stevenson, D.E., 2013, Range 

extensions and significant distributional records for eighttwo species of fishes in Alaska marine waters: Northwestern 

Naturalist, v. 94, no. 1, p. 1–21.

Matarese, A.C., Blood, D.M., Picquelle, S.J., and Benson, 

J.L., 2003, Atlas of abundance and distribution patterns 

of ichthyoplankton from the northeast Pacific Ocean and 

Bering Sea ecosystems based on research conducted by the 

Alaska Fisheries Science Center (1972–1996): National 

Oceanic and Atmospheric Administration Professional 

Paper NMFS 1, 281 p.

Matarese, A.C., Kendall, J., Arthur W., Blood, D.M., and 

Vinter, B.M., 1989, Laboratory guide to early life history 

stages of northeast Pacific fishes: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration, National Marine Fisheries Service 

Technical Report NMFS 80, 652 p.

Mathis, J.T., and Questel, J.M., 2013, Assessing seasonal 

changes in carbonate parameters across small spatial 

gradients in the Northeastern Chukchi Sea: Continental 

Shelf Research, v. 67, p. 42–51, accessed February 3, 

2014, at http://www.sciencedirect.com/science/article/pii/

S0278434313001416.

Maule, A.G., and Thorsteinson, L.K., 2012, Feasibility plan 

to assess the effects of dispersed oil on Arctic Cod: Final 

report prepared by the U.S. Department of the Interior, 

U.S. Geological Survey for the Bureau Ocean Energy 

Management, OCS Study BOEM 2011-062, 52 p.

McAllister, D.E., 1959, Records of marine fishes from fresh 

water in British Columbia: The Canadian Field-Naturalist, 

v. 73, no. 1, p. 13–14.

McAllister, D.E., 1975, Ecology of the marine fishes of Arctic 

Canada, in Proceedings of the Circumpolar Conference 

on Northern Ecology, September 15–18, 1975: Ottawa, 

National Research Council of Canada, p. II-49–II-65.

McAllister, D.E., Anderson, M.E., and Hunter, J.G., 1981, 

Deep-water eelpouts, Zoarcidae, from Arctic Canada 

and Alaska: Canadian Journal of Fisheries and Aquatic 

Sciences, v. 38, no. 7, p. 821–839.

McArdle, H.J., and Johnston, I.A., 1980, Evolutionary 

temperature adaptation in fish muscle sarcoplasmic 

reticulum: Journal of Comparative Physiology, v. 135B, 

p. 157–164.

McCart, P.J., Craig, P.C., and Bain, H., 1972, Report 

of fisheries investigations in the Sagavanirtok River 

and neighboring drainages: Bellevue, Wash., Aquatic 

Environments Ltd., Report to Alyeska Pipeline Service 

Company, 165 p.
722 Alaska Arctic Marine Fish Ecology Catalog

McConnaughey, T., and McRoy, C.P., 1979, 13C label identifies 

eelgrass (Zostera marina) carbon in an Alaskan estuarine 

food web: Marine Biology, v. 53, no. 3, 

p. 263–269.

McDowall, R.M., 1987, The occurrence and distribution 

of diadromy among fishes: American Fisheries Society 

Symposium, no. 1, p. 1–13.

McDowall, R.M., 2008, Jordan’s and other ecogeographical 

rules, and the vertebral number in fishes: Journal of 

Biogeography, v. 35, p. 501–508.

McGregor, S.B., 1990, Seasonal and ontogenetic changes 

in meiofauna in the diets of postmetamorphic flatfishes: 

University of Alaska, Fairbanks, Master’s thesis.

McKechnie, R.J., and Fenner, R.B., 1971, Food habits of 

white sturgeon, Acipenser transmontanus in San Pablo and 

Suisun Bays, California: California Fish and Game, v. 57, 

p. 209–212.

McKinney, M.L., 1997, Extinction vulnerability and 

selectivity—Combining ecological and paleontological 

views: Annual Review of Ecology and Systematics, v. 28, 

p. 495–516. 

McLaren, I.A., 1958, The biology of the ringed seal (Phoca 

hispida Schreber) in the Eastern Canadian Arctic: Bulletin 

Fisheries Research Board of Canada 118, 97 p.

McPhail, J.D., and Lindsey, C.C., 1970, Freshwater fishes of 

northwestern Canada and Alaska: Bulletin of the Fisheries 

Research Board of Canada Bulletin 173, 381 p.

Mecklenburg, C.W., 2003a, Family Anarhichadidae Bonaparte 

1846—Wolffishes: California Academy of Sciences 

Annotated Checklists of Fishes, no. 10, 6 p.

Mecklenburg, C.W., 2003b, Family Hemitripteridae Gill 

1872—Sea ravens or sailfin sculpins: California Academy 

of Sciences Annotated Checklists of Fishes no. 5, 6 p.

Mecklenburg, C.W., 2003c, Family Zaproridae Jordan and 

Evermann 1898—Prowfishes: California Academy of 

Sciences Annotated Checklists of Fishes No. 13.

Mecklenburg, C.W., Byrkjedal, I., Christiansen, J.S., 

Karamushko, O.V., Lynghammar, A., and Møller, P.R., 

2013, List of marine fishes of the Arctic Region annotated 

with common names and zoogeographic characterizations: 

Akureyri, Iceland, Conservation of Arctic Flora and Fauna, 

35 p.

Mecklenburg, C.W., Byrkjedal, I., Karamushko, O.V., 

and Møller, P.R., 2014, Atlantic fishes in the Chukchi 

Borderland: Marine Biodiversity, v. 44, no. 1, p. 127–150, 

doi 10./1007/s12526-013-0192-1.

Mecklenburg, C.W., and Mecklenburg, T.A., 2009, Arctic 

marine fish museum specimens, 2nd ed., Metadata report 

and database submitted to ArcOD, Institute of Marine 

Science: University of Alaska, Fairbanks, by Point Stephens 

Research, metadata report accessed August 8, 2012, at 

http://www.arcodiv.org/Database/Fish_datasets.html.

Mecklenburg, C., and Mecklenburg, T.A., 2011a, Longhead 

Dab—Limanda proboscidea Gilbert, 1896: Arctic Ocean 

Diversity—Census of Marine Life, accessed April 2011 at 

http://www.arcodiv.org/Fish/Limanda_proboscidea.html. 

Mecklenburg, C.W., and Mecklenburg, T.A., 2011b, Polar 

Eelpout—Lycodes polaris (Sabine, 1824): Arctic Ocean 

Diversity Web site, accessed 2011 at http://www.arcodiv.

org/Fish/Lycodes_polaris.html.

Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., 

and Chernova, N.V., 2006, Arctic marine fish museum 

specimens—Metadata report and database submitted to 

ArcOD, Institute of Marine Science: University of Alaska 

Fairbanks by Point Stephens Research, metadata report, 

http://dw.sfos.uaf.edu/rest/metadata/ArcOD/2007F1.

Mecklenburg, C.W., Mecklenburg, T.A., Sheiko, B.A., 

and Steinke, D., 2016, Pacific Arctic marine fishes: 

Akureyri, Iceland, Conservation of Arctic Flora and Fauna, 

Monitoring Series Report No. 23, 406 p., accessed May 10, 

2016, at http://caff.is/monitoring-series/370-pacific-arcticmarine-fishes. 

Mecklenburg, C.W., Mecklenburg, T.A., and Thorsteinson, 

L.K., 2002, Fishes of Alaska: Bethesda, Maryland, 

American Fisheries Society, 1,116 p.

Mecklenburg, C.W., Møller, P.R., and Steinke, D., 2011, 

Biodiversity of Arctic marine fishes—Taxonomy and 

zoogeography: Marine Biodiversity, v. 41, no. 1, p. 109–

140, Online Resources 1–5.

Mecklenburg, C.W., Norcross, B.L., Holladay, B.A., and 

Mecklenburg, T.A., 2008, Fishes, in Hopcroft, R., Bluhm, 

B., and Gradinger, R., eds., Arctic Ocean Synthesis—

Analysis of climate change impacts in the Chukchi and 

Beaufort Seas with strategies for future research: Fairbanks, 

Alaska, Institute of Marine Science, University of Alaska, 

p. 66–79.

Mecklenburg, C.W., and Sheiko, B.A., 2003, Family 

Cyclopteridae Bonaparte 1831—Lumpsuckers: California 

Academy of Sciences Annotated Checklists of Fishes no. 6, 

17 p.

Mecklenburg, C.W., Stein, D.L., Sheiko, B.A., Chernova, 

N.V., Mecklenburg, T.A., and Holladay, B.A., 2007, 

Russian–American long-term census of the Arctic—Benthic 

fishes trawled in the Chukchi Sea and Bering Strait, August 

2004: Northwestern Naturalist, v. 88, no. 3, p. 168–187.
Chapter 8 723

Mecklenburg, C.W., and Steinke, D., 2015, Ichthyofaunal 

baselines in the Pacific Arctic region and RUSALCA study 

area, Oceanography, v. 28, no. 3, p. 158–189.

Mellilo, J.M., Richmond, T.C., and Yohe, G.W., eds., 2014, 

Climate change impacts in the United States—The third 

annual national climate assessment: U.S. Global Climate 

Research Program, 841 p.

Meyer Ottesen, C.A., Hop, H., Christiansen, J.S., and FalkPetersen, S., 2011, Early life history of the daubed shanny 

(Teleostei: Leptoclinus maculatus) in Svalbard waters: 

Marine Biodiversity, v. 41, no. 3, p. 383–394.

Mikhail, M.Y., and Welch, H.E., 1989, Biology of Greenland 

cod, Gadus ogac, at Saqvaqjuac, northwest coast of Hudson 

Bay: Environmental Biology of Fishes, v. 26, p. 49–62.

Miller, B.S., 1967, Stomach contents of adult starry flounder 

and sand sole in East Sound, Orcas Island, Washington: 

Journal of the Fisheries Research Board of Canada, v. 24, 

no. 12, p. 2,515–2,521.

Miller, B.S., and Kendall, A.W., 2009, Early life history of 

marine fishes: Berkley, University of California Press, 

437 p.

Miller, B.S., Simenstad, C.A., Moulton, L.L., Fresh, K.L., 

Funk, F.C., Karp, W.A., and others, 1977, Puget Sound 

baseline program—Nearshore fish survey: University of 

Washington, Fishery Research Institute, FRI-UW-7710, 

219 p.

Miller, D.J., and Lea, R.N., 1976, Guide to the coastal marine 

fishes of California: California Fisheries Bulletin no. 157, 

255 p. 

Miller, D.J., and Schmidtke, J., 1956, Report on the 

distribution and abundance of Pacific herring (Clupea 

pallasi) along the coast of central and southern California: 

California Fish and Game, v. 42, no. 3, p. 163–187.

Miller, J.M., Burke, J.S., and Fitzhugh, G.R., 1991, Early 

life history patterns of Atlantic North Atlantic American 

flatfish—Likely (and unlikely) factors controlling 

recruitment: Netherlands Journal of Sea Research, v. 27, 

p. 261–275.

Miller, T.J., 2007, Contribution of individual-based coupled 

physical-biological models to understanding recruitment in 

marine fish populations: Marine Ecology Progress Series, 

v. 347, p. 127–138.

Milliman, J.D., and Meade, R.H., 1983, World-wide delivery 

of river sediment to the oceans: Journal of Geology, v. 91, 

no. 1, p. 1–21.

Miner, P., and Rivasplata, A., 1994, Thresholds of significance; 

criteria for defining significance: Sacramento, Calif., CEQA 

Technical Advice Series, Governor’s Office of Planning and 

Research, 12 p.

Minerals Management Service, 2008, Beaufort Sea and 

Chukchi Sea planning areas—Oil and gas lease sales 

209, 212, 217, and 221: U.S. Department of the Interior, 

Minerals Management Service Alaska OCS Region, OCS 

EIS/EA: MMS 2008-0055, 538 p. 

Mineva, T.A., 1964, On the biology of some flatfishes in the 

eastern Bering Sea, in Moiseev, P.A. ed., Soviet fisheries 

investigations in the northeastern Pacific, Part II: Jerusalem, 

Israel Program for Scientific Translations, p. 227–235.

Mito, K.-I., Nishimura, A., and Yanagimoto, T., 1999, Ecology 

of groundfishes in the eastern Bering Sea, with emphasis 

on food habits, in Loughlin, T.R., and Ohtani, K., eds., 

Dynamics of the Bering Sea: Fairbanks, University of 

Alaska Sea Grant, p. 537–580.

Moiseev, P.A., 1953, Cod and flounders of far-eastern seas: 

Izvestiya TINRO, Proceedings Pacific Science Research 

Institute of Marine Fisheries and Oceanography, v. 40, 

p. 1–287.

Moles, A., and Norcross, B.L., 1995, Sediment preference 

in juvenile Pacific flatfishes: Netherlands Journal of Sea 

Research, v. 34, no. 1–3, p. 177–182.

Møller, P.R., 2000, Restoration of the taxon Lycodes 

marisalbi, with notes on its disjunct Arctic distribution: 

Journal of Fish Biology, v. 57, no. 6, p. 1,404–1,415.

Møller, P.R., 2001, Redescription of the Lycodes pallidus

species complex (Pisces, Zoarcidae), with a new species 

from the Arctic/North Atlantic Ocean: Copea, v. 2001, 

no. 4, p. 972–996. 

Møller, P.R., and Jørgensen, O.A., 2000, Distribution and 

abundance of eelpouts (Pisces, Zoarcidae) off West 

Greenland: Sarsia, v. 85, no. 1, p. 23–48.

Møller, P.R., Nielsen, J.G., Knudsen, S.W., Poulsen, J.Y., 

Sünksen, K., and Jørgensen, O.A., 2010, A checklist of the 

fish fauna of Greenland waters: Zootaxa, v. 2378, p. 1–84.

Monterio, R.S.O., 2002, Fish growth modeling: growth of the 

European anchovy (Engraulis encrasicolus) in the Tagus 

Estuary, Portugal: European Diploma Eruditionum Altarum 

Modelling of the Marine Environment, 53 p.

Montgomery, J.C., and Macdonald, J.A., 1984, Performance of 

motor systems in Antarctic fishes: Journal of Comparative 

Physiology, v. 154A, p. 241–248.
724 Alaska Arctic Marine Fish Ecology Catalog

Montgomery, J.C., and Sutherland, K.B.W., 1997, Sensory 

development of the Antarctic silverfish Pleurogramma 

antarcticum—A test for the ontogenetic shift hypothesis: 

Polar Biology, v. 18, p. 112–115.

Moran, C., and Melan, R., 2001, Temperature-dependent 

conduction properties in Arctic fish peripheral nerves: Polar 

Biology, v. 24, p. 9–15.

Mork, J., Giskeödegård, R., and Sundnes, G., 1983, 

Haemoglobin polymorphism in Gadus morhua—Genotypic 

differences in maturing age and within-season gonad 

maturation: Helgoländer Meeresunters, v. 36, p. 313–322.

Morris, R.W. and Schneider, M.J., 1969, Brain fatty acids of 

an Antarctic fish, Trematomus bernacchii: Comparatuve 

Biochemistry and Physiology, v. 28, p. 1,461–1,465.

Morrow, J.E., 1980, The freshwater fishes of Alaska: 

Anchorage, Alaska Northwest Publishing Company, 248 p.

Moser, H.G., 1996, The early stages of fishes in the California 

current region: Atlas, California Cooperative Oceanic 

Fisheries Investigations, no. 33, 1,505 p.

Mouillot, D., Bellwood, D.R., Baraloto, C., Chave, J., and 

Galzin, R., 2013, Rare species support vulnerable functions 

in high-diversity ecosystems: PLoS Biology, v. 11, no. 5, 

e1001569, doi:10.1371/journal.pbio.1001569.

Moulton, L.L., and Penttila, D.E., 2000, Forage fish spawning 

distribution in San Juan County and protocols for 

sampling intertidal and nearshore regions: Mount Vernon, 

Washington, San Juan County Forage Fish Assessment 

Project, Northwest Straits Commission, 53 p.

Moulton, L.L., Seavey, B., and Pausanna, J., 2010, History of 

an under-ice subsistence fishery for Arctic Cisco and Least 

Cisco in the Colville River, Alaska: Arctic, v. 63, no. 4, 

p. 381–390.

Moulton, L.L., and Tarbox, K.E., 1987, Analysis of Arctic cod 

movements in the Beaufort Sea nearshore region, 1978–79: 

Arctic, v. 40, no. 1, p. 43–49.

Moyle, P.B., Kiernan, J.D., Crain, P.K., and Quiñones, R.M., 

2013, Climate change vulnerability of native and alien 

freshwater fishes of California—A systematic assessment 

approach: PLoS ONE, v. 8, no. 5, e63883, doi:10.1371/

journal.pone.0063883.

Murray, H.M., Hew, C.L., and Fletcher, G.L., 2002, Skin-type 

antifreeze protein expression in integumental cells of larval 

winter flounder: Journal of Fish Biology, v. 60, no. 6, 

p. 1,391–1,406.

Murray, H.M., Hew, C.L., and Fletcher, G.L., 2003, Spatial 

expression patterns of skin-type antifreeze protein in winter 

flounder (Pseudopleuronectes americanus) epidermis 

following metamorphosis: Journal of Morphology, v. 257, 

no. 1, p. 78–86.

Musick, J.A., 1999, Criteria to define extinction risk in marine 

fishes: Fisheries, v. 24, no. 12, p. 6–14.

Mueter, F.J., Bond, N.A., Ianelli, J.N., and Hollowed, A.B., 

2011, Expected declines in recruitment of walleye pollock 

(Theragra chalcogramma) in the eastern Bering Sea under 

future climate change: ICES Journal of Marine Science, 

v. 68, no. 6, p. 1,284–1,296.

Mueter, F.J., Broms, C., Drinkwater, K.F., Friedland, K.D., 

Hare, J.A., Hunt, G.L., Jr., Melle, W., and Taylor, M., 2009, 

Ecosystem response to recent oceanographic variability in 

high-latitude Northern Hemisphere ecosystems: Progress in 

Oceanography, v. 81, no. 1–4, p. 93–110.

Mueter, F.J., Lauth, R., Andrews, A., de Robertis, A., Britt, 

L., Farley, E., Norcross, B., Sigler, M., and Wilson, C., 

2013, The marine fish communities of the Bering and 

Chukchi Sea: Paper presented at the 2013 Sea Grant 

Lowell Wakefield Symposium, Responses of Arctic Marine 

Ecosystems to Climate Change, March 26–29, 2013, 

Anchorage, Alaska.

Mueter, F.J., and Litzow, M.A., 2008, Sea ice retreat alters 

the biogeography of the Bering Sea continental shelf: 

Ecological Applications, v. 18, no. 2, p. 309–320. 

Mueter, F.J., and Norcross, B.L., 2002, Spatial and temporal 

patterns in the demersal fish community on the shelf and 

upper slope regions of the Gulf of Alaska: Fishery Bulletin 

100, no. 3, p. 559–581.

Murphy, S.M., Mueter, F.J., and Braund, S.R., 2007, Variation 

in the abundance of Arctic Cisco in the Colville River—

Analysis of existing data and local knowledge, volume 

1—Final report: U.S. Department of the Interior, Minerals 

Management Service, OCS Study MMS 2007-042, 240 p.

Musienko, L.N., 1970, Reproduction and development of 

Bering Sea fishes, in Moiseev, P.A., ed., Soviet fisheries 

investigations in the northeastern Pacific—part V: 

Jerusalem, Israel Program for Scientific Translations, 

p. 161–224.

Nahrgang, J., Camus, L., Broms, F., Christiansen, J.S., and 

Hop, H., 2010, Seasonal baseline levels of physiological 

and biochemical parameters in polar cod (Boreogadus 

saida)—Implications for environmental monitoring: Marine 

Pollution Bulletin, v. 60, no. 8, p. 1,336–1,345.
Chapter 8 725

Napazakov, V.V., and Chuchukalo, V.I., 2003, Diet and trophic 

status of mass species of sculpins (Cottidae) in the western 

part of the Bering Sea in autumn: Journal of Ichthyology, 

v. 43, no. 3, p. 236–244.

National Aeronautics and Space Administration, 2012, 

Understanding earth—The icy Arctic: Greenbelt, Maryland, 

Goddard Space Flight Center, 12 p. 

National Oceanic and Atmospheric Administration, 2008, 

West coast of North America living marine resources data 

atlas: Rockville, Maryland, U.S. Department of Commerce, 

National Oceanic and Atmospheric Administration, 

Strategic Assessment Branch, Ocean Assessment Division, 

Office of Oceanography and Marine Assessment, National 

Ocean Service. 

National Oceanic and Atmospheric Administration, 2009, 

World ocean database 2009: National Oceanographic 

and Atmospheric Administration, National Centers for 

Environmental Information database, accessed October 

20, 2010, at http://www.nodc.noaa.gov/OC5/WOD09/

pr_wod09.html.

National Oceanic and Atmospheric Administration, 2010, Age 

and growth program: National Oceanic and Atmospheric 

Administration, accessed April 2011 at http://www.afsc.

noaa.gov/refm/age/Stats/max_age.htm.

 National Oceanic and Atmospheric Administration, 2012, 

Groundfish Assessment Program: National Oceanic and 

Atmospheric Administration database, accessed November 

10, 2013, at http://www.afsc.noaa.gov/RACE/groundfish/

default.php.

National Oceanic and Atmospheric Administration, 2013a, 

Effects of oil and gas activities in the Arctic Ocean—

Supplemental draft environmental impact statement: Silver 

Spring, Md., U.S. Department of Commerce, National 

Marine Fisheries Service, Office of Protected Resources, 

v. 1, chaps. 1–3, 445 p.

National Oceanic and Atmospheric Administration, 2013b, 

Effects of oil and gas development in the Arctic Ocean, 

Supplemental Draft Environmental Impact Statement: 

U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, National Marine Fisheries 

Service, 445 p., accessed April 2, 2013 at http://www.nmfs.

noaa.gov/pr/permits/eis/arctic_sdeis_vol1.pdf.

National Oceanic and Atmospheric Administration., 2014, 

Summer bottom surface temperatures—Eastern Bering Sea: 

Alaska Fisheries Science Center, accessed November 24, 

2014, at http://access.afsc.noaa.gov/reem/EcoWeb/content/

pdf/EBSSummerBottomTemps.pdf.

National Oceanic and Atmospheric Administration, 2014, 

Alaska Fisheries Science Center: National Oceanic and 

Atmospheric Administration Fisheries, Web site accessed 

November 24, 2014, at http://www.afsc.noaa.gov/.

National Research Council, 1996, The Bering Sea ecosystem: 

Washington D.C., National Academies Press, 307 p.

National Snow and Ice Data Center, 2011a, Monthly sea ice 

extent for 1980 and 2007: National Snow and Ice Data 

Center database, accessed April 15, 2011, at http://nsidc.

org/.

National Snow and Ice Data Center, 2011b, NSIDC BIST, 

Compare data, spreadsheet—Sea Ice Index—Extent and 

concentrations trends: accessed April 15, 2011, at http://

nsidc.org/cgi-bin/bist/bist.pl?config=seaice_extent_ 

trends&submit=Go%21. 

National Snow and Ice Data Center, 2011c, State of the 

cryosphere—Is the cryosphere sending signals about 

climate change?: Boulder, Colorado, University of 

Colorado, accessed April 15, 2011, at http://nsidc.org/sotc/ 

sea_ice.html.

Naumenko, E.A., 1996, Distribution, biological condition, 

and abundance of capelin (Mallotus villosus socialis) in 

the Bering Sea, in Mathisen, O.A., and Coyle, K.O., eds., 

Ecology of the Bering Sea: Fairbanks, University of Alaska, 

Alaska Sea Grant College Program, p. 237–256.

Nazarov, D.Y., Kucheryavyi, A.V., Savvaitova, K.A., 

Gruzdeva, M.A., Kuzishchin, K.V., and Pavlov, D.S., 

2011, Population structure of Arctic lamprey Lenthenteron 

camtschaticum from the Kol River (western Kamchatka): 

Journal of Ichthyology, v. 51, no. 4, p. 277–290.

Negulescu, P.A., Rubinsky, B., Fletcher, G.L., and Machen, 

T.E., 1992, Fish antifreeze proteins block Ca entry into 

rabbit parietal cells: American Journal of Physiology, Cell 

Physiology, v. 263, p. 1,310–1,313.

Nelson, J.S., 2006, Fishes of the World: Hoboken, New Jersey, 

John Wiley and Sons, Inc., 416 p.

Nelson, J.S., Crossman, E.J., Espinosa- Pérez, H., Findley, 

L.T., Gilbert, C.R., Lea, R.N., and Williams, J.D., 2004, 

Common and scientific names of fishes from the United 

States, Canada, and Mexico (6th ed.): American Fisheries 

Society Special Publication 29, 386 p. 

Nelson, R., and Bouchard, C., 2013, Arctic Cod (Boreogadus 

saida) population structure and connectivity as examined 

with molecular genetics: Final Report to the North Pacific 

Research Board, NRPB Project 1125, 39 p.
726 Alaska Arctic Marine Fish Ecology Catalog

Nichol, D.G., 1995, Spawning and maturation of female 

yellowfin sole in the eastern Bering Sea, in Alaska 

Sea Grant Program, Proceedings of the International 

Symposium on North Pacific Flatfish: Alaska Sea Grant 

Program, AK-SG-95-04, p. 35–50.

Nichol, D.G., 1997, Effects of geography and bathymetry on 

growth and maturity of yellowfin sole, Pleuronectes asper, 

in the eastern Bering Sea: Fishery Bulletin, v. 95, no. 3, 

p. 494–503.

Nichol, D.G., 1998, Annual and between-sex variability 

of yellowfin sole, Pleuronectes asper, spring-summer 

distributions in the eastern Bering Sea: Fishery Bulletin, v. 

96, no. 3, p. 547–561.

Nichol, D.G., and Acuna, E.I., 2001, Annual and batch 

fecundities of yellowfin sole, Limanda aspera, in the eastern 

Bering Sea: Fishery Bulletin, v. 99, no. 1, p. 108–122.

Nielsen, J.G., and Fosså, S.A., 1993, Lycodes adolfi, a new 

species of eelpout (Zoarcidae) from Greenland: Cybium, 

v. 17, p. 39–44.

Nielsen, J.L., Ruggerone, G.T., and Zimmerman, C.E., 2012, 

Adaptive strategies and life history characteristics in a 

warming climate—Salmon in the Arctic?: Environmental 

Biology of Fishes, v. 96, no. 10–11, p. 1,187–1,226.

Nikolsky, G.V., 1963, The ecology of fishes: London, 

Academic Press, 352 p.

Norcross, B.L., Blanchard, A., and Holladay, B.A., 1999, 

Comparison of models for defining nearshore flatfish 

nursery areas in Alaska waters: Fisheries Oceanography, 

v. 8, p. 50–67.

Norcross, B.L., Holladay, B.A., Busby, M.S., and Mier, K.L., 

2010, Demersal and larval fish assemblages in the Chukchi 

Sea: Deep-Sea Research II, v. 57, no. 1–2, p. 57–70.

Norcoss, B.L., Holladay, B.A., and Mecklenburg, C.W., 

2013a, Recent and historical distribution and ecology 

of demersal fishes in the Chukchi Sea Planning Area: 

Fairbanks, Alaska, OCS Study BOEM 2012-073, 210 p. 

Norcross, B.L., Müter, F.-J., and Holladay, B.A., 1997, Habitat 

models for juvenile pleuronectids around Kodiak Island, 

Alaska: Fishery Bulletin, v. 95, no. 3, p. 504–520.

Norcross, B.L., Raborn, S.W., Holladay, B.A., Gallaway, B.J., 

Crawford, S.T., Priest, J.T., Edenfield, L.E. and Meyer, 

R., 2013b, Northeastern Chukchi Sea demersal fishes and 

associated environmental characteristics, 2009–2010, 

Continental Shelf Research v. 67, p. 77–95.

North Pacific Fishery Management Council, 2009, Fishery 

management plan for fish resources of the Arctic 

management area: Anchorage, Alaska, North Pacific Fishery 

Management Council, 158 p.

North Pacific Fishery Management Council, 2010, Fishery 

management plan for groundfish of the Bering Sea and 

Aleutian Islands management area—Appendices—Fishery 

management plan for groundfish of the BSAI Management 

Area: North Pacific Fisheries Management Council, 233 p.

Novikov, N.P., 1964, Basic elements of the biology of the 

Pacific halibut (Hippoglossus hippoglossus stenolepis

Schmidt) in the Bering Sea, in Moiseev, P.A., ed., Soviet 

fisheries investigations in the northeastern Pacific, Part II: 

Jerusalem, Israel Program for Scientific Translations, 

p. 175–219.

Nozères, C., Archambault, D., Chouinard, P.-M., Gauthier, 

J., Miller, R., Parent, E., Schwab, P., Savard, L., and Dutil, 

J.-D., 2010, Identification guide for marine fishes of the 

estuary and northern Gulf of St. Lawrence and sampling 

protocols used during trawl surveys between 2004 and 

2008: Canadian Technical Report of Fisheries and Aquatic 

Sciences no. 2866.

Ochman, S., and Dodson, J.J., 1982, Composition and 

structure of the larval and juvenile fish community of the 

Eastman River and estuary, James Bay: Naturaliste Canada, 

v. 109, p. 803–813.

Odum, E.P., 1985, Ecologia: Rio de Janeiro, Brazil, 

Interamericana, 434 p.

Odum, W.E., and Heald, E.J., 1975, The detritus-based food 

web of an estuarine mangrove community, in Cronin, L.E., 

ed., Estuarine research: New York, Academic Press, Inc., 

p. 265–286.

Ofelt, C.H., 1975, Food habits of nesting bald eagles in 

southeast Alaska: The Condor, v. 77, p. 337–338.

O’Connell, M., Dillon, M.C., Wright, J.M., Bentzen, P., 

Merkouris, S.E., and Seeb, J.E., 1998, Genetic structuring 

among Alaskan Pacific herring populations identified using 

microsatellite variation: Journal of Fish Biology, v. 53, 

no. 1, p. 150–163.

O’Dea, N.R., and Haedrich, R.L., 2001, COSEWIC status 

report on the northern wolffish Anarhichas denticulatus

in Canada, in Committee on the Status of Endangered 

Wildlife in Canada, Status report on the northern wolffish 

Anarhichas denticulatus in Canada: Ottawa, Committee on 

the Status of Endangered Wildlife in Canada, p. 1–21.

O’Grady, S.M., and DeVries, A.L., 1982, Osmotic and ionic 

regulation in polar fishes: Journal of Experimental Marine 

Biology and Ecology, v. 57, no. 2–3, p. 219–228.

Olden, J.D., Lawler, J.J., and Poff, N.L., 2008, Machine 

learning methods without tears—A primer for ecologists: 

The Quarterly Review of Biology, v. 83, no. 2, p. 171–193.
Chapter 8 727

Orcutt, H.G., 1950, The life history of the starry flounder 

Platichthys stellatus (Pallas): Fish Bulletin, California Fish 

and Game, v. 78, 64 p.

Orlov, A.M., 1999, Some aspects of trophic relations among 

Pacific predatory fishes off the northern Kuril Islands and 

southeastern Kamchatka, in MacKinlay, D., and Houlihan, 

D., eds., Gutshop ‘98: American Fisheries Society, 

p. 41–52.

Orlov, A.M., 2003, Impact of eddies on spatial distributions 

of groundfishes along waters off the northern Kuril Islands, 

and southeastern Kamchatka (north Pacific Ocean): Indian 

Journal of Marine Sciences, v. 32, no. 2, p. 95–111.

Orlov, A.M., 2005, Bottom trawl-caught fishes and some 

features of their vertical distribution in the Pacific waters off 

the north Kuril Islands and south-east Kamchatka, 1993–

1999: Aqua, Journal of Ichthyology and Aquatic Biology of 

Fishes, v. 9, no. 4, p. 139–160.

Orr, A.J., Banks, A.S., Mellman, S., Huber, H.R., and DeLong, 

R.L., 2004, Examination of the foraging habits of Pacific 

harbor sea (Phoca vitulina richardsoni) to describe their 

use of the Umpqua River, Oregon, and their predation on 

salmonids: Fishery Bulletin, v. 102, no. 1, p. 108–117.

Orr, J.W., and Matarese, A.C., 2000, Revision of the genus 

Lepidopsetta Gill, 1862 (Teleostei:Pleuronectidae) based 

on larval and adult morphology, with a description of a 

new species from the North Pacific Ocean and Bering Sea: 

Fishery Bulletin, v. 98, p. 539–582.

Orr, J.W., Wildes, S., Kai, Y., Raring, N., Nakabo, T., Katugin, 

O., and Guyon, J., 2015, Systematics of North Pacific sand 

lances of the genus Ammodytes based on molecular and 

morphological evidence, with the description of a new 

species from Japan: Fishery Bulletin no. 113, p. 129–156.

Orsi, J.A., and Landingham, J.H., 1985, Numbers, species, and 

maturity stages of fish captured with beach seines during 

spring 1981 and 1982 in some nearshore marine waters of 

southeastern Alaska: National Oceanic and Atmospheric 

Administration, Technical Memorandum NMFS F/NWC86, 68 p.

Overland, J.E., and Wang, M., 2013, When will the summer 

Arctic be nearly sea ice free?: Geophysical Research 

Letters, v. 40, p. 2,097–2,101.

Pacunski, R.E., Livingston, P.A., and Miller, B.S., 1998, 

Food of flathead sole Hippoglossoides elassodon in the 

eastern Bering Sea: National Oceanic and Atmospheric 

Administration, Technical Memorandum NMFS-AFSC-90, 

36 p.

Paemelaere, E., and Dobson, S.F., 2011, Fast and slow life 

histories of carnivores: Canadian Journal of Zoology, v. 89, 

no. 8, p. 692–704.

Page, L.M., Espinosa-Pérez, H., Findley, L.T., Gilbert, C.R., 

Lea, R.N., Mandrak, N.E., Mayden, R.L., and Nelson, 

J.S., 2013, Common and scientific names of fishes from 

the United States, Canada, and Mexico: Bethesda, Md., 

American Fisheries Society Special Publication 34, 384 p.

Pahlke, K.A., 1985, Preliminary studies of capelin (Mallotus 

villosus) in Alaskan waters: Alaska Department of Fish and 

Game, Informational Leaflet No. 250, 64 p.

Paine, R.T., 1966, Food web complexity and species diversity: 

The American Naturalist, v. 100, no. 910, p. 65–75.

Palmer, D.E., and Dugan, L.J., 1990, Fish population 

characteristics of Arctic National Wildlife Refuge coastal 

waters, summer 1989: Fairbanks, Alaska, U.S. Fish and 

Wildlife Service, Progress Report, 83 p.

Palsson, W.A., 1990, Pacific cod (Gadus macrocephalus) 

in Puget Sound and adjacent waters—Biology and stock 

assessment: Olympia, State of Washington, Department of 

Fisheries, Technical Report no. 112, 137 p.

Panchenko, V.V., 2001, Reproduction peculiarities of plain 

sculpin Myoxocephalus jaok in Peter the Great Bay, Sea of 

Japan: Russian Journal of Marine Biology, v. 27, 

p. 111–112. 

Panchenko, V.V., 2002, Age and growth of sculpins of the 

genus Myoxocephalus (Cottidae) in Peter the Great Bay (the 

Sea of Japan): Journal of Ichthyology, v. 42, no. 7, 

p. 516–522.

Panchenko, V.V., 2005, Seasonal distribution of antlered 

sculpin, Enophrys diceraus (Cottidae) in Peter the Great 

Bay, Sea of Japan: Russian Journal of Marine Biology, 

v. 31, p. 273–278.

Parker-Stetter, S.L., Horne, J.K., and Weingartner, T.J., 2011, 

Distribution of polar cod and age-0 fish in the U.S. Beaufort 

Sea: Polar Biology v. 34, p. 1,543–1,557.

Parin, N.V., 2004, Ichtyofauna of the seas of Russia—

Biological diversity and fishery potential: Izvestiya TINRO, 

v. 137, p. 226–231.

Parin, N.V., Evseenko, S.A., and Vasil’eva, E.D., 2014, Ryby 

morei Rossii—Annotirovannyi catalog (Fishes of Russian 

seas—Annotated catalogue): Moscow: KMK Scientific 

Press, 733 p. [In Russian.]

Parrish, J.D., Braun, D.P., and Unnach, R.S., 2003, Are we 

conserving what we say we are?—Measuring ecological 

integrity within protected areas: BioScience, v. 53, no. 9, 

p. 851–860.
728 Alaska Arctic Marine Fish Ecology Catalog

Patton, J.C., Gallaway, B.J., Fechhelm, R.G., and Cronin, 

M.A., 1997, Genetic variation of microsatellite and 

mitochondrial DNA markers in Broad Whitefish (Coregonus 

nasus) in the Colville and Sagavanirktok Rivers in northern 

Alaska: Canadian Journal of Fisheries and Aquatic 

Sciences, v. 54 no. 7, p. 1,548–1,556.

Pauly, D., and Swartz, W., 2007, Marine fish catches in North 

Siberia (Russia, FAO Area 18), in Zeller, D., and Pauly, 

D., eds., Reconstruction of marine fisheries catches for 

key countries and regions (1950–2005): Fisheries Centre, 

University of British Columbia, Canada, Fisheries Centre 

Research Reports, v. 15, no. 2, p. 17–33.

Pauly, D., and Watson, R., 2005, Background and 

interpretation of the ‘Marine Trophic Index’ as a measure of 

biodiversity: Philosophical Translations, The Royal Society 

B, v. 360, no. 1454, p. 415–423.

Pauley, G.B., Bowers, K.L., and Thomas, G.L., 1988, Species 

profiles—Life histories and environmental requirements of 

coast fishes and invertebrates (Pacific Northwest)—Chum 

salmon: U.S. Fish and Wildlife Service and U.S. Army 

Corps of Engineers, Biological Report no. 82 (11.81), 

TR EL-82-4, 17 p.

Percy, R., 1975, Fishes of the outer Mackenzie Delta: Victoria, 

British Columbia, Beaufort Sea Project, Beaufort Sea 

Technical Report, no. 8, 114 p.

Pertseva-Ostroumova, T.A., 1961, The reproduction and 

development of far-eastern flounders: Moscow, Academy 

Nauk U.S.S.R. Institute Okeanol, 486 p. [In Russian.]

 Petersen, M.F., and Steffensen, J.F., 2003, Preferred 

temperature of juvenile Atlantic cod Gadus morhua with 

different haemoglobin genotypes at normoxia and moderate 

hypoxia: Journal of Experimental Biology, v. 206, 

p. 359–364.

Philo, L.M., George, J.C., and Moulton, L.L., 1993b, The 

occurrence and description of anadromous fish in the Dease 

Inlet/Admiralty Bay, Alaska area, 1988–1990: Barrow, 

Alaska, Department of Wildlife Management, North Slope 

Borough and MJM Research, 150 p.

Philo, L.M., George, J.C., and Moulton, L.L., 1993a, The 

occurrence and description of anadromous and freshwater 

fish in Teshekpuk Lake, Alaska, 1990–1992: Barrow, 

Alaska, Department of Wildlife Management, North Slope 

Borough and MJM Research, 98 p.

Piatt, J.F., and Springer, A.M., 2003, Advection, pelagic food 

webs and the biogeography of seabirds in Beringia: Marine 

Ornithology, v. 31, p. 141–154.

Piatt, J.F., Wells, J.L., MacCharles, A., and Fadely, B.S., 1989, 

The distribution of seabirds and fish in relation to ocean 

currents in the southeastern Chukchi Sea, in Montevecchi, 

W.A., and Gaston, A.J., eds., Studies of high-latitude 

seabirds 1—Behavioral, energetic, and oceanographic 

aspects of seabird feeding ecology: Canadian Wildlife 

Service, Occasional Papers, no. 68, p. 21–31.

Pietsch, T.W., 1993, Systematics and distribution of 

cottid fishes of the genus Triglops Reinhardt (Teleostei: 

Scorpaeniformes): Zoological Journal of the Linnean 

Society, v. 109, p. 335–393.

Pirtle, J.L., and Mueter, F.J., 2011, Beaufort Sea fish and 

their trophic linkages—Literature search and synthesis: 

Anchorage, Alaska, U.S. Department of the Interior, Bureau 

of Ocean Management, Regulation, and Enforcement, 

BOEMRE 2011-021, 41 p.

Pockett, S., and Macdonald, J.A., 1986, Temperature 

dependence of neurotransmitter release in the Antarctic fish 

Pagothenia borchgrevinki: Experientia, v. 42, no. 4, 

p. 414–415.

Policansky, D., and Sieswerda, P., 1979, Early life history of 

the starry flounder, Platichthys stellatus, reared through 

metamorphosis in the laboratory: Transactions of the 

American Fisheries Society, v. 108, no. 3, p. 326–327.

Polyak, L., Alley, R.B., Andrews, J.T., Brigham-Grette, J., 

Cronin, T.M., Darby, D.A. and others, 2010, History of sea 

ice in the Arctic: Quaternary Science Reviews v. 29, 

p. 1,757–1,778.

Ponomarenko, V.P., 1968, Some data on the distribution and 

migrations of polar cod in the seas of the Soviet Arctic: 

Rapports et Procès-Verbeaux des Réunions du Conseil 

International pour l’Exploration de la Mer, v. 158, 

p. 131–135.

Pörtner, H.O., 2002, Physiological basis of temperaturedependent biogeography—Trade-offs in muscle design and 

performance in polar ectotherms: Journal of Experimental 

Biology, v. 205, p. 2,217–2,230.

Pörtner, H.O., Berdal, B., Blust, R., Brix, O., Colosimo, A., De 

Wachter, B., Giuliani, A., Johansen, T., Fischer, T., Knust, 

R., Lannig, G., Naevdal, G., Nedenes, A., Nyhammer, G., 

Sartoris, F.J., Serendero, I., Sirabella, P., Thorkildsen, S., 

and Zakhartsev, M., 2001, Climate induced temperature 

effects on growth performance, fecundity of recruitment in 

marine fish—Developing a hypothesis for cause and effect 

relationships in Atlantic cod (Gadus morhua) and common 

eelpout (Zoarces viviparous): Continental Shelf Research, 

v. 21, no. 18–19, p. 1,975–1,997.
Chapter 8 729

Pörtner, H.O., Bock, C., Rainer, K., Lanning, G., Lucassen, 

M., Mark, F.C., and Satoris, F.J., 2008, Cod and climate in a 

latitudinal cline: physiological analyses of climate effects in 

marine fishes: Climate Research, v. 37, p. 253–270.

Pörtner, H.O., Lucassen, M., and Storch, D., 2005, Metabolic 

biochemistry—Its role in thermal tolerance and in the 

capacities of physiological and ecological function, in

Farrell, A.P., and Steffensen, J.F., eds., The physiology of 

polar fishes: Amsterdam, Elsevier, p. 79–154.

Power, G., 1997, A review of fish ecology in Arctic North 

America: American Fisheries Society Symposium, no. 19, 

p. 13–39.

Praebel, K., Hunt, B., Hunt, L., DeVries, A.L., 2009, The 

presence of quantification of splenic ice in the McMurdo 

Sound notothenioid fish, Pagothenia borchgrevinki

(Boulenger, 1902)—Comparative biochemistry and 

physiology part A: Molecular and Integrative Physiology, 

v. 154, no. 4, p. 564–569.

Praebel, K., and Ramlov, H., 2005, Antifreeze activity in the 

gastrointestinal fluids of Arctogadus glacialis (Peters, 1874) 

is dependent on food type: Journal of Experimental Biology, 

v. 208, p. 2,609–2,613.

Protection of the Arctic Marine Environment, 2013, Large 

Marine Ecosystems (LMEs) of the Arctic area—Revision of 

the LME map 15 May 2013, 2nd ed.: Arctic Council, 24 p.

Pruter, A.T., and Alverson, D.L., 1962, Abundance, 

distribution, and growth of flounders in the south-eastern 

Chukchi Sea: Journal de Conseil International pour 

l’Exploration de la Mer., v. 27, p. 79–99.

Quast, J.C., 1974, Density distribution of juvenile Arctic cod, 

Boreogadus saida, in the eastern Chukchi Sea in the fall of 

1970: Fishery Bulletin, v. 72, no. 4, p. 1,094–1,105.

R Core Team, 2013, R—A language and environment for 

statistical computing: Vienna, Austria, R Foundation for 

Statistical Computing, accessed November 5, 2013, at 

http://www.R-project.org/.

Rand, K.M., and Logerwell, E.A., 2011, The first demersal 

trawl survey of benthic fish and invertebrates in the 

Beaufort Sea since the late 1970s: Polar Biology, v. 34, 

no. 4, p. 475–488.

Rasmussen, C., Purcell, M.K., Gregg, J.L., LaPatra, S.E., 

Winton, J.R., and Hershberger, P.K., 2010, Sequence 

analysis of the internal transcribed spacer (ITS) region 

reveals a novel clade of Ichthyophonus sp. from rainbow 

trout: Diseases of Aquatic Organisms, v. 89, p. 179–183.

Rass, T.S., 1941, Analogous or parallel variations in structure 

and development of fishes in the northern and Arctic seas: 

Jubilee Publication of the Moscow Society of Naturalists 

1805–1940, p. 1–60. 

Rass, T.S., 1989, Regularity in the increase of egg mass size 

among marine fishes and invertebrates polewards from the 

Equator to high latitudes: Rapports et Procès-Verbeaux des 

Réunions du Conseil International pour l’Exploration de la 

Mer, v. 191, p. 443–444.

Ratynski, R.A., 1983, Mid-summer ichthyoplankton 

populations of Tuktoyaktuk Harbor, N.W.T.: Canadian 

Technical Report of Fisheries and Aquatic Sciences, 

no. 1218, 21 p.

Raymond, J.A., 1992, Glycerol is a colligative antifreeze in 

some northern fishes: Journal of Experimental Zoology, 

v. 262, no. 3, p. 347–352.

Raymond, J.A., 1994, Seasonal variations of trimethylamine 

oxide and urea in the blood of a cold-adapted marine teleost, 

the rainbow smelt: Fish Physiology and Biochemistry, v. 13, 

no. 1, p. 13–22.

Raymond, J.A., 1997, Responses of marine fishes to freezing 

temperatures—A new look at colligative mechanisms, in

Willis, J.S., ed., Advances in molecular and cell biology: 

Greenwich, Conn., JAI Press, p. 33–142.

Raymond, J.A., and DeVries, A.L., 1977, Adsorption 

inhibition as a mechanism of freezing resistance in polar 

fishes: Proceedings of the National Academy of Sciences of 

the USA, v. 74, no. 6, p. 2,589–2,593.

Raymond, J.A., and DeVries, A.L., 1998, Elevated 

concentrations and synthetic pathways of trimethylamine 

oxide and urea in some teleosts fishes of McMurdo Sound, 

Antarctica: Fish Physiology and Biochemistry, v. 18, no. 4, 

p. 387–398.

Raymond, J.A., Lin, Y., and DeVries, A.L., 1975, Glycoprotein 

and protein antifreezes in two Alaskan fishes: Journal of 

Experimental Zoology, v. 193, no. 1, p. 125–130.

Reist, J.D., and Chang-Kue, K.T.J., 1997, The life history and 

habitat usage of broad whitefish in the lower Mackenzie 

River Basin, in Tallman, R.F., and Reist, J.D., eds., The 

proceedings of the broad whitefish workshop—The biology, 

traditional knowledge and scientific management of 

broad whitefish (Coregonus nasus (Pallas)) in the Lower 

Mackenzie River: Canadian Technical Report of Fisheries 

and Aquatic Sciences, no. 2193, p. 63–84.

Reist, J.D., Wrona, F.J., Prowse, T.D., Power, M., Dempson, 

J.B., Beamish, R.J., King, J.R., Carmichael, T.J., and 

Sawatzky, C.D., 2006, General effects of climate change on 

Arctic fishes and fish populations, AMBIO: A Journal of the 

Human Environment v. 35, no. 7, p. 370–380.
730 Alaska Arctic Marine Fish Ecology Catalog

Renaud, P.E., Carroll, M.L., and Ambrose, W.G., Jr., 2008, 

Effects of global warming on Arctic sea-floor communities 

and its consequences for higher trophic levels, in Duarte, 

C.M., ed., Impacts of global warming on polar ecosystems: 

Bilbao, Spain, Fundación BBVA, p. 141–177.

Reynolds, W.W., 1977, Temperature as a proximate factor 

in orientation behavior: Journal of the Fisheries Research 

Board of Canada, v. 34, no. 5, p. 734–739.

Richardson, E.S., Reist, J.D., and Minns, C.K., 2001, Life 

history characteristics of freshwater fishes occurring in the 

Northwest Territories and Nunavut, with major emphasis on 

lake habitat requirements: Canadian Manuscript Report of 

Fisheries and Aquatic Sciences no. 2569, 156 p.

Ricker, W.E., 1975, Computation and interpretation of 

biological statistics of fish populations: Ottawa, Canada, 

Bulletin of Fisheries Research Board of Canada, no. 191, 

382 p.

Robards, M.D., and Piatt, J.F., 1999, Biology of the genus 

Ammodytes, the sand lances, in Robards, M.D., and others, 

eds., Sand land—A review of biology and predator relations 

and annotated bibliography: U.S. Forest Service, Pacific 

Northwest Research Station, Research Paper PNW-RP-521.

Robards, M.D., Piatt, J.F., and Rose, G.A., 1999, Maturation, 

fecundity, and intertidal spawning of Pacific sand lance in 

the northern Gulf of Alaska: Journal of Fish Biology v. 54, 

no. 5, p. 1,050–1,068.

Robards, M.D., Rose, G.A., and Piatt, J.F., 2002, Growth and 

abundance of Pacific sand lance, Ammodytes hexapterus, 

under differing oceanographic regimes: Environmental 

Biology of Fishes, v. 64, no. 4, p. 429–441.

Robinson, L.M., Elith, J., Hobday, A.J., Pearson, R.G., 

Kendall, B.E., Possingham, H.P., and Richardson, A.J., 

2011, Pushing the limits in marine species distribution 

modeling—Lessons from the land present challenges and 

opportunities: Global Ecology and Biogeography, v. 20, 

p. 789–802.

Robles, V., Cabrita, E., Fletcher, G.L., Shears, M.A., King, 

M.J., and Herráez, M.P., 2005, Vitrification assays with 

embryos from a cold tolerant sub-arctic fish species: 

Theriogenology v. 64, no. 7, p. 1,633–1,646.

Roessig, J.M., Woodley, C.M., Cech, J.J., Jr., and Hansen, 

L.J., 2004, Effects of global climate change on marine and 

estuarine fishes and fisheries: Reviews in Fish Biology and 

Fisheries, v. 14, no. 2, p. 251–275.

Rogers, D.E., Rabin, D.J., Rogers, B.J., Garrison, K.J., and 

Wangerin, M.E., 1979, Seasonal composition and food web 

relationships of marine organisms in the nearshore zone of 

Kodiak Island—including ichthyoplankton, meroplankton 

(shellfish), zooplankton, and fish: Seattle, University of 

Washington College of Fisheries, Fisheries Research 

Institute, Technical Report FRI-UW-7906, 123 p.

Romanuk, T.N., Hayward, April, and Hutchings, J.A., 2011, 

Trophic level scales positively with body size in fishes: 

Global Ecology and Biogeography, v. 20, p. 231–240, 

doi: 10.1111/j.1466-8238.2010.00579.x.

Ronholt, L.L., Wakabayashi, K., Wilderbuer, T.K., Yamaguchi, 

H., and Okada, K., 1986, Ground fish resource of the 

Aleutian Island waters based on the U.S.–Japan trawl 

survey, June–November 1980: Bulletin International North 

Pacific Fisheries Commission 48, p. 251.

Rooper, C.N., 2002, English sole transport during pelagic 

stages on the Pacific Northwest coast, and habitat use 

by juvenile flatfish in Oregon and Washington estuaries: 

Seattle, University of Washington, Ph.D. dissertation.

Rose, G.A., 2005a, Capelin (Mallotus villosus) distribution 

and climate: a sea “canary” for marine ecosystem change: 

ICES Journal of Marine Science, v. 62, no. 7, 

p. 1,524–1,530.

Rose, G.A., 2005b, On distributional responses of North 

Atlantic fish to climate change: ICES Journal of Marine 

Science, v. 62, no. 7, p. 1,360–1,374.

Rosenberg, A., Bigford, T.E., Leathery, S., Hill, R., and 

Bickers, K., 2000, Ecosystem approaches to fishery 

management through essential fish habitat: Bulletin of 

Marine Science, v. 66, no. 3, p. 535–542.

Rosenthal, R.J., 1980, Shallow water fish assemblages in the 

northeastern Gulf of Alaska—Habitat evaluation, species 

composition, abundance, spatial distribution and trophic 

interaction, in Bureau of Land Management, Environmental 

assessment of the Alaskan Continental Shelf, final reports 

of principal investigators: U.S. Department of Commerce, 

National Oceanic and Atmospheric Administration, 

p. 451–540.

Rosenthal, R.J., Haldorson, L.J., Field, L.J., MoranO’Connell, V., LaRiviere, M.G., Underwood, J., and others, 

1982, Inshore and shallow offshore bottomfish resources in 

the southeastern Gulf of Alaska (1981–82): Alaska Coastal 

Research and University of Alaska, Juneau, 166 p.

Roy, K., Jablonski, D., Vaelntine, J.W., and Rosenberg, 

G., 1998, Marine latitudinal gradients—Tests of causal 

hypotheses: Proceedings of the National Academy of 

Science, v. 95, no. 7, p. 3,699–3,702.
Chapter 8 731

Rubinsky, B., Arav, A., Mattioli, M., and DeVries, A.L., 

1990, The effect of antifreeze glycopeptides on membrane 

potential changes at hypothermic temperatures: Biochemical 

and Biophysical Research Communications, v. 173, no. 3, 

p. 1,369–1,374.

Russian-American Long-Term Census of the Arctic, 2009, 

Expedition to the Bearing Strait and Pacific side of 

the Arctic Ocean: National Oceanic and Atmospheric 

Administration Web site, accessed April 15, 2013, at http://

www.arctic.noaa.gov/rusalca/2009.

Sadorus, L.L., Lauth, R., and Ranta, A.M., 2012, Size and age 

composition of Pacific halibut in NMFS Bering Sea shelf 

trawl surveys, in IPHC Report of Assessment and Research 

Activities 2012: International Pacific Halibut Commission 

online report, p. 593–600, accessed February 16, 2014, at 

http://www.iphc.int/publications/rara/2012/rara2012593_

beringtrawlcomp.pdf.

Safronov, S.N., 1986, Peculiarities of reproduction and 

principles of change in the fecundity of Pacific navaga, 

Eleginus gracilis (Gadidae): Journal of Ichthyology, v. 26, 

no. 5, p. 59–68.

Salveson, S.J., 1976, Alaska plaice (family Pleuronectidae), 

in Pereya, W.T., Reeves, J.E., and Bakkala, R.G., eds., 

Demersal fish and shellfish resources of the Eastern Bering 

Sea in the baseline year 1975: National Oceanic and 

Atmospheric Administration, p. 489–495.

Salveson, S.J., and Alton, M.S., 1976, Yellowfin sole (family 

Pleuronectidae) in Pereya, W.T., Reeves, J.E., and Bakkala, 

R.G., eds.: Demersal fish and shellfish resources of the 

Eastern Bering Sea in the baseline year 1975: Seattle, 

Washington, National Marine Fisheries Service, Northwest 

and Alaska Fisheries Center, Processed Report, p. 439–459.

Sameoto, D.D., 1988, Feeding of lantern fish Benthosema 

glaciale off the Nova Scotia shelf: Marine Ecology Progress 

Series, v. 44, p. 113–129.

Sandercock, F.K., 1991, Life history of Coho salmon 

(Oncorhynchus kisutch), in Groot, C., and Margolis, L., 

eds., Pacific salmon life histories: Vancouver, British 

Columbia, UBC Press, p. 397–445.

Saunders, M.W., and McFarlane, G.A., 1993, Age and length 

of maturity of the female spiny dogfish, Squalus acanthias, 

in the Strait of Georgia, British Columbia, Canada: 

Environmental Biology of Fishes, v. 38, p. 49–57.

Schmidt, D.R., Griffiths, W.B., Beaubien, D.K., and 

Herlugson, C.J., 1991, Movement of young-of-the-year 

Arctic ciscoes across the Beaufort Sea coast, 1985–1988: 

American Fisheries Society Symposium, no. 11, 

p. 132–144.

Schmidt, D.R., Griffiths, W.B., and Martin, L.R., 1987, 

Importance of anadromous fish overwintering habitat in 

the Sagavanirktok River Delta, Alaska: Anchorage, Alaska, 

Report by Ecological Research Associates for Standard 

Alaska Production Company and North Slope Borough, 

71 p.

Schmidt, D.R., Griffiths, W.R., and Martin, L.R., 1989, 

Overwintering biology of anadromous fish in the 

Sagavanirktok River Delta, Alaska: Biological Papers of the 

University of Alaska, v. 24, p. 55–74.

Scholander, P.F., Flagg, W., Walters, V., and Irving, L., 1953, 

Climatic adaptations in Arctic and tropical poikilotherms: 

Physiological Zoology, v. 26, no. 1, p. 67–92.

Scott, G.K., Fletcher, G.L., and Davies, P.L., 1986, Fish 

antifreeze proteins—Recent gene evolution: Canadian 

Journal of Fisheries and Aquatic Science, v. 43, no. 5, 

p. 1,028–1,034.

Scott, G.K., Hayes, P.H., Fletcher, G.L., and Davies, P.L., 

1988, Wolffish antifreeze protein genes are primarily 

organized as tandem repeats that each contain two genes in 

inverted orientation: Molecular and Cellular Biology, v. 8, 

no. 9, p. 3,670–3,675.

Scott, W.B., and Scott, M.G., 1988, Atlantic fishes of Canada: 

Toronto, University of Toronto Press, 730 p.

Seitz, A.C., Loher, T., and Nielsen, J.L., 2008, Seasonal 

movements and environmental conditions experienced 

by Pacific halibut along the Aleutian Islands, examined 

by pop-up satellite tags: International Pacific Halibut 

Commission Scientific Report, no. 85, 24 p. 

Seitz, A.C., Norcross, B.L., and Nielsen, J.L., 2003, Pip-up 

archival transmitting (PAT) tags—A method to investigate 

the migration and behavior of Pacific halibut Hippoglossus 

stenolepsis in the Gulf of Alaska: Alaska Fishery Research 

Bulletin, v. 10, no. 2, p. 124–135.

Seitz, A.C., Norcross, B.L., Wilson, D., and Nielsen, 

J.L., 2005, Identifying spawning behavior in Pacific 

halibut, Hippoglossus stenolepis, using electronic tags: 

Environmental Biology of Fishes, v. 73, p. 445–451.

Sekerak, A.D., Stallard, N., and Griffiths, W.B., 1992, 

Distribution of fish and fish harvests in the nearshore 

Beaufort Sea and Mackenzie Delta during ice-covered 

periods, October–June: Environmental Studies Research 

Funds Report, LGS Ltd., no. 117, 524 p.

Shchetinnikov, A.S., 1983, Nutrition of Acantholumpenus 

mackayi (Stichaeidae) in the Gulf of Terpenium (Sakhalin 

Island): Journal of Ichthyology, v. 23, no. 6, p. 155–158.
732 Alaska Arctic Marine Fish Ecology Catalog

Shears, M.A., Kao, M.H., Scott, G.K., Davies, P.L., and 

Fletcher, G.L., 1993, Distribution of type III antifreeze 

in the Zoarcoidei: Molecular Marine Biology and 

Biotechnology, v. 2, p. 104–111.

Sheiko, B.A., and Fedorov, V.V., 2000, Class 

cephalaspidomorphi-lampreys, Class chondrichthyescartilaginous fishes, Class holocephali-chimaeras, Class 

osteichthyes-bony fishes, in Artuykhin, Yu. B, and Sheyko, 

B.A., eds., Catalog of vertebrates of Kamchatka and 

adjacent waters: Russia, Rossiyskaya Akademiya Nauk, 

p. 7–69. [In Russian.]

Sheiko, B.A., and Mecklenburg, C.W., 2004, Family Agonidae 

Swainson 1839—Poachers: California Academy of Sciences 

Annotated Checklists of Fishes, no. 30, 27 p.

Shestakov, A.V., and Nazarkin, M.V., 2006, On the biology of 

the white-spotted greenling Hexagrammos stelleri and the 

masked greenling H. octogrammus (Hexagrammidae) from 

Taui Bay of the Sea of Okhotsk: Journal of Ichthyology, 

v. 46, no. 8, p. 677–680.

Shuntov, V.P., 1970, Seasonal distribution of black and arrowtoothed halibuts in the Bering Sea: in Moiseev, P.A., ed., 

Soviet fisheries investigations in the northeastern Pacific: 

Jerusalem, Israel Program for Scientific Translations 1968, 

p. 397–408.

Sick, K., 1965, Hemoglobin polymorphism of cod in the North 

Sea and the North Atlantic Ocean: Heriditas, v. 54, no. 1, 

p. 49–68.

Siefert, D.L., Incze, L.S., and Ortner, P.B., 1988, Vertical 

distribution of zooplankton, including ichthyoplankton, in 

Shelikof Strait, Alaska—Data from fisheries-oceanography 

coordinated investigations (FOCI) cruise in May 1986: 

National Marine Fisheries Service, NWAFC Processed 

Report, v. 88, no. 28, 232 p.

Sigler, M.F., Renner, M., Danielson, S.L., Eisner, L.B., Lauth, 

R.R., Kuletz, K.J., Logerwell, E.A., and Hunt, G.L., Jr., 

2011, Fluxes, fins, and feathers—Relationships among the 

Bering, Chukchi, and Beaufort Seas in a time of climate 

change: Oceanography, v. 24, no. 3, p. 250–265.

Simenstad, C.A., Isakson, J.S., and Nakatani, R.E., 1977, 

Marine fish communities, in Merritt, M.L., and Fuller, R.G., 

eds., The environment of Amchitka Island, Alaska: National 

Technical Information Center, Energy Research and 

Development Administration TID-26712, p. 451–492.

Sinclair, E.H., and Zeppelin, T.K., 2002, Seasonal and spatial 

differences in diet in the western stock of Steller sea lions 

(Eumetopias jubatus): Journal of Mammology, v. 83, no. 4, 

p. 973–990. 

Slagstad, D., Ellingsen, I.H., and Wassmann, P., 2011, 

Evaluating primary and secondary production in an 

Arctic Ocean void of summer sea ice—An experimental 

simulation approach: Progressive Oceanography, v. 90, 

no. 1–4, p. 117–131.

Smetacek V., and Nicol, S., 2005, Polar ecosystems in a 

changing world: Nature, v. 437, p. 362–368.

Smith, K.R., Somerton, D.A., Yang, M.-S., and Nichol, 

D.G., 2004, Distribution and biology of prowfish (Zaprora 

silenus) in the northeast Pacific: Fishery Bulletin, v. 102, 

no. 1, p. 168–178.

Smith, M.A., 2010, Arctic marine synthesis—Atlas of the 

Chukchi and Beaufort Seas: Anchorage, Alaska, Audubon 

Alaska and Oceana.

Smith, R.E.H., Anning, J., Clement, P., and Cota, G., 1988, 

Abundance and production of ice algae in Resolute Passage, 

Canadian Arctic: Marine Ecology Progress Series, v. 48, 

p. 251–263.

Smith, R.L., Vallarino, M., Barbour, E., Fitzpatrick, E., and 

Barber, W.E., 1997, Population biology of the Bering 

flounder in the Northeastern Chukchi Sea: American 

Fisheries Society Symposium, v. 19, p. 127–132.

Smith, R.L., Barber, W.E., Vallarino, M., Gillispie, J., and 

Ritchie, A., 1997a, Population biology of the Arctic 

staghorn sculpin in the northeastern Chukchi Sea: American 

Fisheries Society Symposium 19, p. 133–139.

Smith, R.L., Vallarino, M., Barbour, E., Fitzpatrick, E., and 

Barber, W.E., 1997b, Population biology of the Bering 

flounder in the northeastern Chukchi Sea: American 

Fisheries Society Symposium, v. 19, p. 127–132.

Smith, R.T., 1936, Report on the Puget Sound otter trawl 

investigations: Washington Department of Fisheries, 

Biological Report No. 36B, 120 p.

Smith, S.L., and Schnack-Schiel, S.B., 1990, Polar 

zooplankton polar oceanography part B—Chemistry 

biology and geology: San Diego, Academic Press, 

p. 527–598.

Sohn, D., Ciannelli, L., and Duffy-Anderson, J.T., 2010, 

Distribution and drift pathways of Greenland halibut 

(Reinhardtius hippoglossoides) during early life stages 

in the eastern Bering Sea and Aleutian Islands: Fisheries 

Oceanography, v. 19, no. 5, p. 339–353.

Sokolovskaya, T.G., Sokolovskii, A.S., and Sobolevskii, E.I., 

1998, A list of fishes of Peter the Great Bay (the Sea of 

Japan): Journal of Ichthyology, v. 38, no. 1, p. 1–11.
Chapter 8 733

Sokolovskii, A.S., and Sokolovskaya, T.G., 1999, The early 

ontogenesis of Enophrys diceraus (Cottidae) from the 

waters of Peter the Great Bay (the Sea of Japan): Journal of 

Ichthyology, v. 39, no. 3, p. 268–272.

Sokolovskii, A., and Sokolovskaya, T., 2009, Distribution of 

ichthyoplankton in the eastern Peter the Great Bay, Sea of 

Japan, in June and July of 2007: Russian Journal of Marine 

Biology, v. 35, no. 3, p. 250–254.

Somero, G.N., 1992, Adapting to water stress—Convergence 

on common solutions, in Somero, G.N., Osmond, C.B., and 

Bolis, C.L., eds., Water and life, comparative analysis of 

water relationships at the organismic, cellular and molecular 

levels: Berlin, Springer-Verlag, p. 3–18.

Sorenson, T.F., and Ramlov, H., 2001, Variations in antifreeze 

activity and serum inorganic ions in the eelpout Zoarces 

viviparous—Antifreeze activity in the embryonic state: 

Comparative Biochemistry and Physiology Part A—

Molecular and Integrative Physiology, v. 130, no. 1, 

p. 123–132.

Spalding, M.D., Fox, H.E., Allen, G.R., Davidson, N., 

Ferdaña, Z.A., Finlayson, M., Halpern, B.S., Martin, 

K.D., Mcmanus, E., Molnar, J., Recchia, C.A., and 

Robertson, J., 2007, Marine ecosystems of the world—A 

bioregionalization of coastal and shelf areas: BioScience, 

v. 57, no. 7, p. 573–583.

Speer, L., and Laughlin, T.L., 2011, IUNC/NRDC workshop 

to identify areas of ecological and biological significance 

or vulnerability in the Arctic marine environment: IUNC/

NRDC Workshop Report, La Jolla, Calif., November 2–4, 

2010, 16 p.

Springer, A.M., Murphy, E.C., Rosenau, D.G., Peter McRoy, 

C., and Cooper, B.A., 1986, The paradox of pelagic food 

webs in the northern Bering Sea-I—Seabird food habits: 

Continental Shelf Research, v. 7, no. 8, p. 895–911.

St.-Pierre, G., 1984, Spawning locations and season for Pacific 

halibut: Seattle, Washington, International Pacific Halibut 

Commission Scientific Report no. 70, 46 p.

Stabeno, P.J., Kachel, N.B., Moore, S.E., Napp, J.M., Sigler, 

M., Yamaguchi, A., and Zerbini, A.N., 2012b, Comparison 

of warm and cold years on the southeastern Bering Sea 

shelf and some implications for the ecosystem, Deep-Sea 

Research II: Tropical Studies in Oceanography. v. 65–70, 

p. 31–45.

Stabeno, P., and others, 2012a, A comparison of the physics 

of the northern and southern shelves of the eastern Bering 

Sea and some implications for the ecosystem, Deep Sea 

Research Part II: Topical Studies in Oceanography, 

v. 65–70, p. 14–30.

Stark, J.W., 2007, Geographic and seasonal variations in 

maturation and growth of female Pacific cod (Gadus 

macrocephalus) in the Gulf of Alaska and Bering Sea: 

Fishery Bulletin, v. 105, no. 3, p. 396–407.

Stauffer, G., comp., 2004, NOAA protocols for groundfish 

bottom trawl surveys of the Nation’s fishery resources: 

U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration Technical Memorandum 

NMFS-F/SPO-65, 205 p.

Stearns, S.C., 1977, The evolution of life history traits—A 

critique of the theory and a review of the data: Annual 

Review of Ecology and Systematics, v. 8, p. 145–171.

Steffensen, J.F., 2002, Metabolic cold adaptation of polar fish 

based on measurements of aerobic oxygen consumption—

Fact or artefact? Artefact!: Comparative Biochemistry and 

Physiology Part A—Molecular and Integrative Physiology, 

v. 132, no. 4, p. 789–795.

Steffensen, J.F., 2005, Respiratory systems and metabolic 

rates, in Farrell, A.P., and Steffensen, J.F., eds., The 

physiology of polar fishes: Amsterdam, Elsevier, 

p. 203–238.

Steffensen, J.F., Bushnell, P.G., and Schurmann, H., 1994, 

Oxygen consumption in four species of teleosts from 

Greenland—No evidence of metabolic cold adaptation: 

Polar Biology, v. 14, no. 1, p. 49–54.

Stehmann, M., and Bürkel, D.L., 1984, Rajidae, in Whitehead, 

P.J.P., and others, eds., Fishes of the North-eastern Atlantic 

and the Mediterranean: Paris, Unesco, p. 163–196.

Stephenson, S.A., 2006, A review of the occurrence of Pacific 

Salmon (Oncorhynchus spp.) in the Canadian Western 

Arctic: Arctic, v. 59, no. 1, p. 37–46.

Stewart, D.B., Ratynski, R.A., Bernier, L.M.J., and Ramsey, 

D.J., 1993, A fishery development strategy for the Canadian 

Beaufort Sea-Amundsen Gulf area: Canadian Technical 

Report Fisheries and Aquatic Sciences 1910, 135 p.

Stewart, H., 1977, Indian fishing—Early methods on the 

northwest coast: Seattle, University of Washington Press, 

182 p.

Stockwell, D., Horner, R., Saitoh, S., and Hill, V., 2008, 

Phytoplankton, in Hopcroft, R., Bluhm, B., and Gradinger, 

R., eds., Arctic Ocean synthesis—Analysis of climate 

change impacts in the Chukchi and Beaufort Seas with 

strategies for future research: Institute of Marine Sciences, 

University of Alaska Fairbanks, p. 33–37.

Stoner, A.W., and Abookire, A.A., 2002, Sediment preferences 

and size-specific distribution of young-of-the-year Pacific 

halibut in an Alaska nursery: Journal of Fish Biology, v. 61, 

no. 3, p. 540–559. 
734 Alaska Arctic Marine Fish Ecology Catalog

Stout, H.A., Gastafson, R.G., Lenarz, W.H., McCain, B.B., 

VanDoornik, D.M., Builder, T.L., and Methot, R.D., 2001, 

Status review of Pacific herring (Clupea pallasi) in Puget 

Sound, Washington: National Oceanic and Atmospheric 

Administration Technical Memorandum NMFSNWFSC-45, 175 p.

Suryan, R.M., Sab, V.S., Wallace, B.P., Hatch, S.A., 

Frederiksen, M., and Wanless, S., 2009, Environmental 

forcing on life history strategies—Evidence for multitrophic level responses at ocean basin scales: Progress in 

Oceanography, v. 81, no. 1–4, p. 214–222.

Swanson, J.A., and Hoppe, A.D., 2004, The coordination of 

signaling during Fc receptor-mediated phagocytosis: Journal 

of Leukocyte Biology, v. 76, no. 6, p. 1,093–1,103.

Tack, S.L., 1970, The summer distribution and standing 

stock of the fishes of Izembek Lagoon, Alaska: Fairbanks, 

University of Alaska, Master’s thesis.

Tallman, R.F., Abrahams, M.V., and Chudobiak, D.H., 2002, 

Migration and life history alternatives in a high latitude 

species, the broad whitefish, Coregonus nasus Pallas: 

Ecology of Freshwater Fish, v. 11, no. 2, p. 101–111.

Tarasyuk, S.N., 2002, Survival of yellowfin sole (Limanda 

aspera Pallas) in the northern part of the Tatar Strait 

(Sea of Japan) during the second half of the 20th century, 

in Batchelder, H.P., and others, eds., PICES-Globe 

International Program on Climate Change and Carrying 

Capacity—Report of 2110 BASS/MODEL, MONITOR and 

REX workshops, and The 2002 MODEL/REX workshop: 

Sidney, British Columbia, North Pacific Marine Science 

Organization (PICES).

Thompson, G., Ianelli, J., Dorn, M., Nichol, D., Gaichas, S., 

and Aydin, K., 2007, Assessment of the Pacific Cod stock 

in the eastern Bering Sea and Aleutian Islands Area, chap. 

2 of Plan team for groundfish fisheries of the Bering Sea/

Aleutian Islands (comps.): Anchorage, Alaska, North Pacific 

Fishery Management Council, Stock assessment and fishery 

evaluation report for groundfish resources of the Bering 

Sea/Aleutian Islands regions, p. 209–328. 

Thorsteinson, F.V., Sample, T., and Thorsteinson, L.K., 

1984, Fishery resources, chap. 6 of Jarvela, L.E., ed., The 

Navarin Basin environment and possible consequences of 

planned offshore oil and gas development: Juneau, Alaska, 

U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, OCSEAP, 157 p.

Thorsteinson, L.K., 1996, National Biological Service OCS 

environmental studies program—research and monitoring 

in Arctic Alaska, in Newbury, T., ed., Proceedings of the 

1995 Arctic Synthesis Meeting, October 23–25, 1995, 

Anchorage, Alaska: U.S. Department of the Interior, 

Minerals Management Service, Alaska OCS Region, OCS 

Study MMS 95-0065, 206 p. plus appendixes.

Thorsteinson, L.K., Becker, P.R., and Hale, D.A., 1989, The 

Yukon Delta synthesis of information: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration, OCS Study MMS 89-0081, 89 p.

Thorsteinson, L.K., Jarvela, L.E., and Hale, D.A., 1990, Arctic 

fish habitat use investigations—Nearshore studies in the 

Alaskan Beaufort Sea, summer 1988: Final Report, Alaska 

Office, Ocean Assessments Division, National Oceanic and 

Atmospheric Administration, Research Unit 682, 125 p.

Thorsteinson, L.K., Jarvela, L.E., and Hale, D.A., 1991, Arctic 

fish habitat use investigations—Nearshore studies in the 

Alaskan Beaufort Sea, summer 1990: Anchorage, Alaska, 

U.S. Department of Commerce, National Oceanic and 

Atmospheric Administration, OCSEAP Final Report to U.S. 

Department of the Interior, Mineral Management Service, 

166 p.

Tokranov, A.M., 1981, Distribution of sculpins (Pisces, 

Cottidae) on the west Kamchatka shelf in summer: 

Zoologicheskii Zhurnal, v. 60, no. 2, p. 229–237.

Tokranov, A.M., 1987a, Feeding of giant sculpin 

Myoxocephalus polyacanthocephalus Pallas and plain 

sculpin M. jaok (Cuvier) (Cottidae) in coastal waters of 

Kamchatka: Journal of Ichthyology, v. 27, p. 104–114.

Tokranov, A.M., 1987b, Species composition and distribution 

of sea poachers (Pisces, Agonidae) in nearshore waters of 

Kamchatka: Zoologicheskii Zhurnal, v. 66, p. 385–392.

Tokranov, A.M., 1991, Diet of the Pacific cod, Gadus morhua 

macrocephalus, and its position in the food chain in 

Kamchatkan coastal waters: Journal of Ichthyology, v. 31, 

no. 5, p. 84–98.

Tokranov, A.M., 1995, The size and sex structure of sculpins 

of the genus Triglops (Cottidae) in the coastal waters of 

Kamchatka: Journal of Ichthyology, v. 35, no. 4, 

p. 140–144.

Tokranov, A.M., 1999, Some features of biology of the 

prowfish Zaprora silenus (Zaproridae) in the Pacific waters 

of the northern Kuril Islands and southeastern Kamchatka: 

Journal of Ichthyology, v. 39, no. 6, p. 475–478.

Tokranov, A.M., 2007, Distribution and some biological 

features of the Pacific sand lance Ammodytes hexapterus

(Ammodytidae) in waters off Kamchatka in the Sea of 

Okhotsk: Journal of Ichthyology, v. 47, no. 4, p. 288–295.

Tokranov, A.M., and Maksimenkov, V.V., 1994, Feeding of the 

starry flounder, Platichthys stellatus, in the Bol’shaya River 

estuary (Western Kamchatka): Journal of Ichthyology, v. 34, 

no. 1, p. 76–83.
Chapter 8 735

Tokranov, A.M., and Orlov, A.M., 2005, Some features of the 

biology of Icelus spatula (Cottidae) in Pacific waters off the 

northern Kuril Islands: Journal of Ichthyology, v. 45, no. 3, 

p. 229–236.

Tokranov, A.M., Orlov, A.M., and Biryukov, I.A., 2004, 

Distribution and size-weight composition of some rare fish 

species in Pacific waters off the northern Kurile Islands and 

southeast Kamchatka: Journal of Ichthyology, v. 44, no. 3, 

p. 208–216.

Tokranov, A.M., and Polutov, V.I., 1984, Fish distribution 

in the Kronitsky Bay in relation to abiotic factors: 

Zoologicheskii Zhurnal, v. 63, p. 1,363–1,373.

Tokranov, A.M., and Zavarina, S.V., 1992, Size-age 

structure and sex ratios of Alaska plaice, Pleuronectes 

quadrituberculatus, on the Western Kamchatka Shelf: 

Journal of Ichthyology, v. 32, no. 7, p. 118–127.

Treble, M.A., Campana, S.E., Wastle, R.J., Jones, C.M., and 

Boje, J., 2008, Growth analysis and age validation of a 

deepwater Arctic fish, the Greenland halibut (Reinhardtius 

hippoglossoides): Canadian Journal of Fisheries and 

Aquatic Sciences, v. 65, no. 6, p. 1,047–1,059.

Trevor, H., 2013, GAM—Generalized Additive Models: R 

package version 1.09, accessed April 15, 2014, at http://

CRAN.R-project.org/package=gam.

Turner, L.M., 1886, Contributions to the natural history of 

Alaska—Arctic series of publications, no. 2: Washington, 

D.C., U.S. Government Printing Office, 226 p.

Tyler, A.V., 1972, Food resource division among northern, 

marine, demersal fishes: Journal of the Fisheries Research 

Board of Canada, v. 29, no. 7, p. 997–1,003.

Underwood, T.J., Gordon, J.A., Millard, M.J., Thorpe, L.A., 

and Osborne, B.M., 1995, Characteristics of selected fish 

populations of Arctic National Wildlife Refuge coastal 

waters, final report, 1988–91: Fairbanks, Alaska, U.S. Fish 

and Wildlife Service, Fairbanks Fishery Resource Office, 

Alaska Fisheries Technical Report, no. 28, 590 p.

United Nations Environment Programme, 2013, What future 

for the Arctic?: Nairobi, Kenya, Background paper for 

the Arctic Side Event at the Governing Council/Global 

Environmental Ministerial Forum, 12 p.

Vadstein, O., Mo, T.A., and Bergh, Ø., 2004, Microbial 

interactions, prophylaxes and diseases, in Moksness, E., 

Kjørsvik, E., and Olsen, Y., eds., Culture of cold-water 

marine fish: Victoria, Australia, Blackwell Publishing, 

p. 28–72.

Valerio, P.F., Kao, M.H., and Fletcher, G.L., 1990, Thermal 

hysteresis activity in the skin of the cunner, Tautogolabrus 

adspersus: Canadian Journal of Zoology, v. 68, no. 5, 

p. 1,065–1,067.

Valerio, P.F., Kao, M.H., and Fletcher, G.L., 1992, Fish skin—

An effective barrier to ice crystal propagation: Journal of 

Experimental Biology, v. 164, p. 135–151.

VanGerwen-Toyne, M., Walker-Larsen, J., and Tallman, 

R.F., 2008, Monitoring spawning populations of migratory 

coregonids in the Peel River, NT—The Peel River fish 

study 1998–2002: Canadian Manuscript Report of Fisheries 

and Aquatic Sciences no. 2851, 56 p.

Van Guelpen, L., 1986, Hookear sculpins (genus Artediellus) 

of North American Atlantic—Taxonomy, morphological 

variability, distribution, and aspects of life history: Canadian 

Journal of Zoology, v. 64, no. 3, p. 677–690.

Van Voorhies, W.V., Raymond, J.A., and DeVries, A.L., 1978, 

Glycoproteins as biological antifreeze agents in the cod, 

Gadus ogac (Richardson): Physiological Zoology, v. 51, 

no. 4, p. 347–353.

Verde, C., Carratore, V., Riccio, A., Tamburrini, M., Parisi, 

E., and di Prisco, G., 2002, The functionally distinct 

hemoglobins of the Arctic spotted wolffish Anarhichas 

minor: Journal of Biological Chemistry, v. 277, no. 39, 

p. 36,312–36,320.

Verde, C., de Rosa, M.C., Giordanao, D., Mosca, D., De 

Pascale, D., Raiola, L., Cocca, E., Carratore, V., Giardina, 

B., and Di Prisco, G., 2005, Structure, function and 

molecular adaptations of haemoglobins of the polar 

cartilaginous fish Bathyraja eatonii and Raja hyperborea: 

Biochemical Journal, v. 389, no. 2, p. 297–306.

Verde, C., Parisi, E., and di Prisco, G., 2006, The evolution of 

thermal adaptation in polar fish: Gene, v. 385, p. 137–145.

Vermeij, G.J., 1991, Marine extinctions and their implications 

for conservation and biogeography, in The Unity 

of Evolutionary Biology, Proceedings of the Fourth 

International Congress of Systematic and Evolutionary 

Biology, v. 1, p. 143–148.

Viavant, T., 2001, Eastern North Slope Dolly Varden 

spawning and over-wintering assessment feasibility: 

Anchorage, Alaska, U.S. Fish and Wildlife Service, 

Office of Subsistence Management, Fisherie Information 

Services Division, Federal Subsistence Fisheries Resource 

Monitoring Program, Final Report for Study 00-002, 25 p.

Viavant, T., 2009, Aerial monitoring of Dolly Varden 

overwintering abundance in the Anaktuvuk, Ivishak, 

Canning, and Hulahula rivers, 2006-2008, Alaska 

Department of Fish and Game, Division of Sport Fish, 

Research and Technical Services, Fishery Data Series No. 

09-21, 9 p.

Vollen, T., and Albert, O.T., 2008, Pelagic behavior of adult 

Greenland halibut (Reinhardtius hippoglossoides): Fishery 

Bulletin, v. 106, no. 4, p. 457–470.
736 Alaska Arctic Marine Fish Ecology Catalog

von Biela, V.R., Zimmerman, C.E., and Moulton, L.L., 2011, 

Long-term increases in young-of-the-year growth of Arctic 

cisco Coregonus autumnalis and environmental influences: 

Journal of Fish Biology, v. 78, no. 1, p. 39–56.

Walkusz, W., Paulic, J.E., Williams, W.J., Kwasniewski, S., 

and Papst, M.H., 2011, Distribution and diet of larval and 

juvenile Arctic cod (Boreogadus saida) in the shallow 

Canadian Beaufort Sea: Journal of Marine Systems, v. 84, 

p. 78–84.

Wang, M., and Overland, J.E., 2009, A sea ice free summer 

Arctic within 30 years?: Geophysical Research Letters, 

v. 36, no. 7, L07502.

Wassmann, P., Duarte, C.M., Agust, S., and Sejr, M.K., 

2011, Footprints of climate change in the Arctic marine 

ecosystem: Global Change Biology, v. 17, no. 2, 

p. 1,235–1,249.

Weingartner, T.J.,1997, A review of the physical oceanography 

of the northeastern Chukchi Sea, in Reynolds, J., ed., Fish 

ecology in Arctic North America: Bethesda, Md., Special 

Publication (No. 19) of the American Fisheries Society, 

p. 40–59.

Weingartner, T., Shimada, K., McLaughlin, F., and 

Proshutinsky, A., 2008, Physical oceanography, in

Hopcroft, R., Bluhm, B., and Gradinger, R., eds., Arctic 

Ocean synthesis—Analysis of climate change impacts 

in the Chukchi and Beaufort Seas with strategies for 

future research: Institute of Marine Science, University of 

Fairbanks, Alaska, p. 6–17.

Weslawski, J.M., Kwaśniewski, S., Stempniewicz, L., and 

Blachowiak-Samolyk, K., 2006, Biodiversity and energy 

transfer to top trophic levels in two contrasting Arctic 

fjords: Polish Polar Research, v. 27, no. 3, p. 259–278. 

Welch, H.E., Crawford, R.E., and Hop, H., 1993, Occurrence 

of Arctic cod (Boreogadus saida) schools and their 

vulnerability to predation in the Canadian High Arctic: 

Arctic, v. 46, no. 4, p. 331–339. 

Wells, R.M.G., 2005, Blood-gas transport and hemoglobin 

function in polar fishes—Does low temperature explain 

physiological characters?, in Farrell, A.P. and Steffensen, 

J.F., eds., The physiology of polar fishes: Amsterdam, 

Elsevier, p. 281–316.

Wernberg T., Smale, D.A., and Thomsen, M.S., 2012, 

A decade of climate change experiments on marine 

organisms—Procedures, patterns, and problems: Global 

Change Biology, v. 18, no. 5, p. 1,491–1,498.

West, R.L., and Fruge, D.J., 1989, A review of coastal plain 

fish surveys and the results of 1986 fish surveys of selected 

coastal lakes and streams Arctic National Wildlife Refuge, 

Alaska: Alaska Fisheries Technical Report, no. 4, 22 p.

Wheeler, A., 1992, A list of common and scientific names of 

fishes of the British Isles, Fisheries Society of the British 

Isles, Academic Press, London.

White House, 2013, National strategy for the Arctic Region: 

Washington D.C., Office of the President, 13 p. https://

www.whitehouse.gov/sites/default/files/docs/nat_arctic_

strategy.pdf.

Whiteley, N.M., Christiansen, J.S., and Egginton, S., 

2006, Polar cod, Boreogadus saida (Gadidae), show 

an intermediate stress response between Antarctic 

and temperate fishes: Comparative Biochemistry and 

Physiology Part A: Molecular and Integrative Physiology, 

v. 145, no. 4, p. 493–501.

Wienerroither, R., Johannesen, E., Langøy, H., Børve Eriksen, 

K., de Lange Wenneck, T., Høines, Å., Bjelland, O., 

Aglen, A., Prokhorova, T., Murashko, P., Prozorkevich, D., 

Konstantin Byrkjedal, I., Langhelle Drevetnyak, G., and 

Smirnov, O., 2011, Atlas of the Barents Sea fishes: IMR/

PINRO Joint Report Series 1-2011, ISSN 1502-8828, 274 p.

Wiese, F.K., Merrick, R., Auad, G., Williams, D., Stabeno, 

P., and Kelly, B., 2013, Strawman conceptual ecosystem 

model for the Chukchi and Beaufort Seas, appendix B 

in Developing a conceptual model of the Arctic marine 

ecosystem: Interagency Arctic Research Policy Committee 

workshop, April 30–May 2, 2013, Washington, D.C., 

p. B1–B29.

Wiese, F., Auad, G., Williams, D., Merrick, R., Jeffries, M., 

Kelly, B., Thornhill, A., Thorsteinson, L., and D. Dickson, 

D., 2015, Framing Arctic marine research initiatives: A 

Framework for coordinated marine ecosystem research 

in the U.S. Chukchi and Beaufort Seas, North Pacific 

Research Board, Anchorage, AK., 34 p., http://www.

iarpccollaborations.org/uploads/cms/documents/framing_

arctic_marine_research_initiatives_report_p2b_lowres.pdf.

Wildermuth, D.A., 1983, Length-weight regression analysis 

for thirty-eight species of sport caught marine fishes: 

Western Department of Fisheries Progress Report, no. 189, 

7 p.

Wilderbuer, T.K., Walters, G.E., and Bakkala, R.G., 1992, 

Yellowfin sole, Pleuronectes asper, of the eastern Bering 

Sea—Biological characteristics, history of exploitation, and 

management: Marine Fisheries Review, v. 54, no. 4, 

p. 1–18.

Wilimovsky, N.J., 1974, Fishes of the Bering Sea—The state 

of existing knowledge and requirements for future effective 

efforts, in Hood, D.W., and Kelly, E.J., eds., Oceanography 

of the Bering Sea with emphasis on renewable resources: 

Proceedings of the International Symposium, Bering Sea 

Study, Hakodate, Japan, January 31–February 4, 1972, 

Institute Marine Science, University of Alaska, Fairbanks, 

Occasional Publication No. 2., p. 243–256.
Chapter 8 737

Williams, B.K., Nichols, J.D., and Conroy, M.H., 2002, 

Analysis and management of animal populations: San 

Diego, Academic Press, 817 p. 

Willson, M.F., Armstrong, R.H., Robards, M.D., and Piatt, 

J.F., 1999, Sand lance as cornerstone prey for predator 

populations, in Robards, M.D., and others, eds., Sand 

land—A review of biology and predator relations and 

annotated bibliography: U.S. Forest Service, Pacific 

Northwest Research Station, Research Paper PNW-RP-521.

Wilson, W.J., and Ormseth, O.A., 2009, A new management 

plan for Arctic waters of the United States: Fisheries, v. 34, 

no. 11, p. 555–558.

Winemiller, K.O., 2004, Life history strategies, population 

regulation, and implications for fisheries management: 

Canadian Journal of Fisheries and Aquatic Sciences, v. 62, 

no. 4, p. 872–885.

Winemiller, K.O, 2005, Life history strategies, population 

regulation, and implications for fisheries management:

Canadian Journal of Fisheries and Aquatic Sciences, v. 62, 

p. 872–885.

Winemiller, K.O., and Rose, K.A., 1992, Patterns of 

life-history diversification in North American fishes—

Implications for population regulation: Canadian Journal of 

Fisheries and Aquatic Sciences, v. 49, no. 10, 

p. 2,196–2,218.

Wing, B.L., 1985, Salmon stomach contents from the 

Alaska troll logbook program 1977–84: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration National Marine Fisheries Service, 

Technical Memorandum NMFS F/NWC-91, 43 p.

Winters, G.H., 1970, Biological changes in coastal capelin 

from the over-wintering to the spawning condition: 

Journal of the Fisheries Board of Canada, v. 27, no. 12, 

p. 2,215–2,224.

Wiswar, D.W., 1994, Summer distribution of Arctic fishes 

in the 1002 Area of the Arctic National Wildlife Refuge, 

Alaska, 1991, with special emphasis on selected lakes, 

tundra streams, and the Sadlerochit River drainage: U.S. 

Fish and Wildlife Service, Alaska Fisheries Technical 

Report Number 27, 47 p.

Wiswar, D.W., and Frugé, D.J., 2006, Fisheries investigations 

in western Camden Bay, Arctic National Wildlife Refuge, 

Alaska, 1987: U.S. Fish and Wildlife Service, Alaska 

Fisheries Data Series 2006-10, 49 p.

Wiswar, D.W., West, R.L., and Winkleman, W.N., 1995, 

Fisheries investigation in Oruktalik Lagoon, Arctic National 

Wildlife Refuge, Alaska, 1986: U.S. Fish and Wildlife 

Service, Alaska Fisheries Technical Report, no. 30, 38 p. 

Wohlschlag, D.E., 1960, Metabolism of an Arctic fish and the 

phenomenon of cold adaptation: Ecology, v. 41, no. 2, 

p. 287–292.

Wolotira, R.J., Jr., Sample, T.M., and Morin, M., Jr., 1977, 

Demersal fish and shellfish resources of Norton Sound, 

the southeastern Chukchi Sea, and adjacent waters in the 

baseline year 1976: Seattle, Wash., Northwest and Alaska 

Fisheries Center, NMFS, National Oceanic and Atmospheric 

Administration, Processed Report, 69 p.

Wood, G., 2008, Thresholds and criteria for evaluating and 

communicating impact significance in environmental 

studies—‘See no evil, hear no evil, speak no evil’?: 

Environmental Impact Assessment Review, v. 28, no. 1, 

p. 22–38.

Wood, S.N., 2003, Thin plate regression splines: Journal of the 

Royal Statistical Society Series B, Statistical Methodology, 

v., 65, no. 1, p. 95–114.

Wood, S.N., 2006, Low-rank scale-invariant tensor product 

smooths for generalized additive mixed models: Biometrics, 

v. 62, no. 4, p. 1,025–1,036.

Wood, S.N., 2011, Fast stable restricted maximum likelihood 

and marginal likelihood estimation of semiparametric 

generalized linear models: Journal of the Royal Statistical 

Society Series B, v. 73, no. 1, p. 3–36.

Woodgate, R.A., K.M. Stafford, K.M., and Prahl, F.G., 

2015. A synthesis of year-round interdisciplinary mooring 

measurements in the Bering Strait (1990-1994) and the 

RUSALCA years (2004–2011), Oceanography, v. 28, no. 3, 

p. 45–67.

Woodson, D., Dello, K., Flint, L., Hamilton, R., Neilson, R., 

and Winton, J., 2011, Climate change effects in the Klamath 

Basin, chap. 6 of Thorsteinson, L., VanderKooi, S., and 

Duffy, W., eds., Proceedings of the Klamath Basin Science 

Conference, Medford, Oregon, February 1–5, 2010: U.S. 

Geological Survey Open-File Report 2011-1196, p. 123–

149. [Also available at http://pubs.er.usgs.gov/publication/

ofr20111196/.]

Wu, Y., and Fletcher, G.L., 2000, Efficacy of antifreeze protein 

types in protecting liposome membrane integrity depends 

on phospholipid class: Biochimica et Biophysica Acta 

(BBA), General Subjects, v. 1524, no. 1, p. 11–16.

Wu, Y., Kao, M.H., Goddard, S.V., Banoub, J., and Fletcher, 

G.L., 2001, Antifreeze glycoproteins—Relationship 

between molecular weight, thermal hysteresis and the 

inhibition of leakage from liposomes during thermotropic 

phase transition: Comparative Biochemistry and Physiology 

Part B—Biochemistry and Molecular Biology, v. 128, no. 2, 

p. 265–273.
738 Alaska Arctic Marine Fish Ecology Catalog

Wyllie Echeverria, T., 1996, Sea-ice conditions and the 

distribution of walleye pollock (Theragra chalcogramma) 

on the Bering and Chukchi Shelf, in Beamish, R.J., ed., 

Climate change and northern fish populations: Canadian 

Special Publication Fisheries and Aquatic Sciences 121, 

p. 131–136.

Yager, T., Kirchman, D., Sherr, E., Sherr, B., and Lovejoy, C., 

2008, Microbes, in Hopcroft, R., Bluhm, B., and Gradinger, 

R., eds., Arctic Ocean synthesis—Analysis of climate 

change impacts in the Chukchi and Beaufort Seas with 

strategies for future research: Institute of Marine Sciences, 

University of Alaska, Fairbanks, p. 38–44.

Yang, M.-S., 1996, Diets of the important groundfishes in 

the Aleutian Islands in summer 1991: U.S. Department 

of Commerce, National Oceanic and Atmospheric 

Administration Technical Memorandum NMFS-AFSC-60, 

Alaska Fisheries Science Center, 105 p.

Yang, M.-S., 2003, Food habits of the important groundfishes 

in the Aleutian Islands in 1994 and 1997: National Marine 

Fisheries Service, AFSC Processed Report 2003-07, 233 p.

Yang, M.-S., Dodd, K., Hibpshman, R., and Whitehouse, A., 

2006, Food habits of groundfishes in the Gulf of Alaska in 

1999 and 2001: U.S. Department of Commerce, National 

Oceanic and Atmospheric Administration Technical 

Memorandum, NMFS-AFSC-164, 189 p.

Yang, M.-S., and Nelson, M.W., 2000, Food habits of the 

commercially important groundfishes in the Gulf of 

Alaska in 1990, 1993, and 1996: Seattle, Washington, 

U.S. Department of Commerce, National Oceanic 

and Atmospheric Administration, NOAA Technical 

Memorandum NMFS-AFSC-112, 174 p.

Yoshihara, H.T., 1973, Annual progress report for monitoring 

and evaluation of Arctic waters with emphasis on the North 

Slope: Alaska Department of Fish Game, Division of Sport 

Fisheries, 49 p.

Zenger, H.H., Jr., 2004, Data report—2002 Aleutian Islands 

bottom trawl survey: U.S. Department of Commerce, 

National Oceanic and Atmospheric Administration 

Technical Memorandum NMFS-AFSC-143, 247 p.

Zhang, C.I., Wilderbuer, T.K., and Walters, G.E., 1998, 

Biological characteristics and fishery assessment of Alaska 

plaice, Pleuronectes quadritubercultus, in the eastern 

Bering Sea: Marine Fisheries Review, v. 60, no. 4, p. 16–27.

Zimmerman, C.E., Ramsey, A.M., Turner, S.M., Mueter, 

F.J., and Murphy, S.M., 2013, Genetics, recruitment, and 

migration patterns of Arctic cisco (Coregonus autumnalis) 

in the Colville River, Alaska, and Mackenzie River, Canada: 

Polar Biology, v. 36, no. 11, p. 1,543–1,555. 
Appendixes

Appendix A. Ongoing and Recently Completed Fishery Studies Funded by

Bureau of Ocean Energy Management in Arctic Outer Continental Shelf 

Lease Areas

Table A1 provides a comprehensive list with descriptions and access to reports of the following studies can be accessed at 

http://www.boem.gov/Environmental-Stewardship/Environmental- Studies/Alaska/Index.aspx.

Table A1. Ongoing studies in the Arctic Outer Continental Shelf lease areas.

[Blank cells indicate that the species occurrence has not been confirmed in that sea. Marine waters out to the U.S. Exclusive Economic Zone (200-mile limit) are 

included]

Bureau of Ocean Energy 

Management study Research topic Researcher organization

Genomics of Arctic Cod Genetic structure and diversity U.S. Geological Survey

U.S.-Canada Transboundary fish and 

lower trophic communities

Ecological baselines of marine fish and 

invertebrates in the Beaufort Sea

University of Alaska, Fairbanks, Department 

of Fisheries and Oceans Canada

Trophic links: Forage fish, their prey, and 

Ice Seals in the northeast Chukchi Sea

Food habits and trophic linkages of ice seals University of Alaska, Fairbanks

Dispersal patterns and summer ocean 

distribution of adult Dolly Varden 

from the Wulik River, Alaska, using 

satellite tags

Coastal and ocean habitats of Dolly Varden Alaska Department of Fish and Game

Population assessment of Snow Crab, 

Chionoecetes opilio, in the Chukchi 

and Beaufort Seas (including oil and 

gas lease areas)

Population dynamics of snow crabs University of Alaska, Fairbanks

Distribution and habitat use of fish in the 

nearshore ecosystem of the Beaufort 

and Chukchi Seas (Alaska Coastal 

Ecosystem Survey)

Coastal habitat use by marine fish National Oceanic and Atmospheric 

Administration Alaska Fisheries Science 

Center

Distribution of fish, crab, and lower trophic 

communities in the Chukchi Sea lease 

area (Arctic Integrated Ecosystem Survey)

Ecological baselines of fish and invertebrates in 

the northeastern Chukchi Sea

University of Alaska, Fairbanks, National 

Oceanic and Atmospheric Administration 

Alaska Fisheries Science Center
740 Alaska Arctic Marine Fish Ecology Catalog

Table A2. Recently completed Bureau of Ocean Energy Management studies by year of completion.

Year Bureau of Ocean Energy Management selected reference citations

2014 Bluhm, B., Huettmann, Falk, and Norcross, Brenda, 2014, Ecological analysis of 2008 western Beaufort Sea data: Anchorage, 

Alaska, U.S. Department of the Interior, Minerals Management Service, Alaska OCS Region, BOEM 2014-014.

Hopcroft, R.R., and Clarke, Cheryl, 2013, Retrieval of historical arctic fisheries survey data: Anchorage, Alaska, U.S. 

Department of the Interior, Minerals Management Service, Alaska OCS Region, BOEM 2014-084, 45 p.

Morris, M.C., 2014, Alaska shorezone coastal habitat mapping protocol: Seldovia, Alaska, Contract Report by Nuka Research 

and Planning Group LCC for the Bureau of Ocean Energy and Energy Management (BOEM), Anchorage, Alaska, 164 p. 

[Also available at http://alaskafisheries.noaa.gov/shorezone/chmprotocolO114.pdf].

Talbot, S.L., Sage, G.K., Sonsthagen, S.A., and Fowler, M.C., 2014, Arctic Cod pilot genomics study—Preliminary results 

from analyses of mitochondrial DNA: Anchorage, Alaska, U.S. Department of the Interior, Minerals Management Service, 

Alaska OCS Region, BOEM 2014-050, 29 p.

Zimmerman, C.E., and von Biela, C.E., 2014, Investigation of population of origin and migration of Arctic Cisco found in 

the Colville River, Alaska using molecular and otolith tools: Anchorage, Alaska, U.S. Department of the Interior, Minerals 

Management Service, Alaska OCS Region, BOEM 2014-019, 107 p.

2013 Carothers, C., Cotton, S., and Moerlein, K., 2013, Subsistence use and knowledge of salmon in Barrow and Nuiqsut, Alaska: 

Fairbanks, Alaska, University of Alaska Coastal Marine Institute and U.S. Department of the Interior, BOEM, Alaska 

OCS Region, OCS Study BOEM 2012-0115. [Also available at http:// www.boem.gov/BOEM-Newsroom/Library/

Publications/201 3/BOEM-2013-0015pdf.aspx]

2012 Dunton, K.H., 2012, Chukchi Sea offshore monitoring in drilling area (COMIDA)—Chemical and Benthos (CAB): 

Anchorage, Alaska, prepared by University of Texas for U.S. Department of the Interior, BOEM Alaska OCS Region, Final 

Report, OCS Study BOEM 2012-012. [Also available at http://www.data.boem.gov/P l /PDFimages/ES P1S/5/5182.pdf].

Norcoss, B.L., Holladay, B.A., and Mecklenburg, C.W., 2012, Recent and historical distribution and ecology of demersal 

fishes in the Chukchi Sea planning area: Fairbanks, Alaska, University of Alaska Coastal Marine Institute and U.S. 

Department of the Interior, BOEM, Alaska OCS Region, OCS Study BOEM 2012-073. [Also available at: http://www.

boem.gov/BOEM-Newsroom/Library/Publications/2012/CMI-2012-073_pdf.aspx].

2011 Hardy, S.M., lken, Katrin, Hundertmark, Kris, Albrecht, Greg, 2011, Defining genetic structure in Alaskan populations of the 

snow crab, Chionoecetes opilio: Anchorage, Alaska, U.S. Department of the Interior, Minerals Management Service, Alaska 

OCS Region, BOEMRE 2011-060, 31 p.

Konar, Brenda, 2012, Recovery in a high Arctic kelp community: Anchorage, Alaska, U.S. Department of the Interior, 

Minerals Management Service, Alaska OCS Region, BOEM 2012-0II, 24 p.

Loggerwell, Elizabeth, Rand, Kimberly, Parker-Stetter, Sandra, Horne, John, Weingartner, Tom, and Bluhm, Bodil, 2010, 

Beaufort Sea marine fish monitoring 2008—Pilot survey and test of hypotheses: Anchorage, Alaska, U.S. Department of the 

Interior, Minerals Management Service, Alaska OCS Region, BOEMRE 201 0-048, 262 p.

Pirtle, J.L., and Mueter, F.J., 2011, Beaufort Sea fish and their trophic linkages—Literature search and synthesis: Anchorage, 

Alaska, U.S. Department of the Interior, Minerals Management Service, Alaska OCS Region, BOEMRE 2011-021, 47 p.

2007 Murphy, S.M., Mueter, F.J., and Braund, S.R., 2007, Variation in the abundance of Arctic Cisco in the Colville River—

Analysis of existing data and local knowledge, volumes I and II: Anchorage, Alaska, U.S. Department of the Interior, 

Minerals Management Service, Alaska OCS Region, MMS 2007-042, 240 p.

2004 MBC Applied Environmental Sciences, 2004, Proceedings of a workshop on the variability of Arctic Cisco (Qaaktaq) in the 

Colville River: Anchorage, Alaska, U.S. Department of the Interior, Minerals Management Service, Alaska OCS Region, 

MMS 2004-033, 90 p.

1998 Kline, T., Jr., and Goering, J., 1998, North Slope amphidromy assessment: Anchorage, Alaska, U.S. Department of the Interior, 

Minerals Management Service, Alaska OCS Region, MMS 1998-006, 90 p.
Appendix B. Age-At-Size and Length-Weight Relationships for Arctic 

Marine Fishes

The Bureau of Ocean Energy Management (BOEM) Alaska Outer Continental Shelf (OCS) Region is in the process of 

publishing new age-at-size and length-weight relationships for marine fish species from new studies in the U.S. Chukchi and 

Beaufort Seas. As this report was nearing publication, a summary of the length-weight relationships from BOEM’s U.S.– 

Canada Transboundary study in the Beaufort Sea was provided to the USGS (Brenda L. Norcross and others, University of 

Alaska, Fairbanks, written commun., 2016; and Kelly L. Walker, University of Alaska, Fairbanks, written commun., 2016). 

The transboundary study included field collections in offshore waters of the southeastern Beaufort and western Arctic Canada 

between 2012 and 2014. 

There were 20 species for which sufficient numbers of fish were captured so that length-weight relationships could be 

established (table B1). The species are from nine families: Gadidae–Boreogadus saida, Cottidae–Gymnocanthus tricuspis, Icelus 

bicornis, I. spatula, Triglops nybelini, and Triglops pingelii, Psycholutidae–Cottunculus microps, Agonidae–Aspidophoroides 

olrikii, Cyclopteridae–Eumicretremus derjugini, Liparidae–Careproctus sp. and Liparis fabricii, Zoarcidae–Lycodes adolfi, 

L. polaris, L. sagittarius and L. seminudus, Stichaeidae–Anisarchus medius, Lumpenus fabricii and Sticheaus punctatus, and 

Pleuronectidae–Reinhardtius hippoglossoides. Fish sampling encompassed pelagic and benthic environments of the Beaufort 

Sea and the minimum and maximum lengths reported for each species varied greatly. The weight-at-length regressions fit the 

data closely, with r2 values of 0.90–0.99 and all intercepts (a) were near zero (table B1). The range of slopes (b) was 2.49–3.59. 

A b value close to 3.0 indicates isometric growth, that is, growth of all body parts occurs at the same rate; values outside of that 

range indicate allometric growth, that is, the body changes shape with growth. The b value also indicates body shape; negative 

allometric growth (b < 3) and positive allometric growth (b > 3) indicate decreasing or increasing body thickness or plumpness 

with increasing fish length (Brenda L. Norcross and others, University of Alaska, Fairbanks, written commun., 2016; and 

Kelly L. Walker, University of Alaska, Fairbanks, written commun., 2016).
742 Alaska Arctic Marine Fish Ecology Catalog

Table B1. Summary of length-weight relationships of marine fish collected in the U.S.–Canada transboundary study in the Beaufort 

Sea.

[Source: Brenda L. Norcross and others, University of Alaska, Fairbanks, written commun., 2016; and Kelly L. Walker, University of Alaska, Fairbanks, 

written commun., 2016. a: intercepts. b: range of slopes. r2: coefficient of determination. Abbreviations: g, gram; mm, millimeter]

Species Number Weight range 

(g)

Length range 

(mm) a*105 b r2

Boreogadus saida 2,877 0.03–106.13 15–240 0.587 3.01 0.98

Artediellus scaber 137 0.03–13.63 14–95 1.690 2.98 0.99

Gymnocanthus tricuspia 683 0.08–20.89 19–119 0.315 3.33 0.99

Icelus bicornis 97 0.23–4.45 27–68 0.270 3.37 0.96

Icelus spatula 412 0.09–7.86 24–89 0.488 3.20 0.90

Triglops nybelini 15 4.29–14.67 81–118 0.425 3.14 0.93

Triglops pingelli 234 0.15–14.30 26–130 0.834 2.97 0.98

Cottonculus microps 14 1.27–208.33 45–223 2.770 2.93 0.99

Asidophoroides olrikii 335 0.04–3.69 23–80 0.351 3.17 0.93

Eumicretemus derjugini 8 0.23–14.48 15–64 4.170 3.07 0.99

Careproctus sp. 41 0.72–112.53 47–145 0.071 3.59 0.98

Liparis fabricii 120 0.07–112.53 19–212 0.050 3.58 0.93

Lycodes adolfi 232 0.19–26.62 38–205 0.201 3.09 0.97

Lycodes polaris 64 0.24–26.79 40–164 0.161 3.26 0.99

Lycodes saggittarius 191 0.33–347.60 44–427 0.812 2.88 0.92

Lycodes seminudus 154 0.30–535.99 41–465 1.540 2.82 0.98

Anisarchus medius 65 0.23–5.15 49–134 2.790 2.49 0.93

Lumpenus facricii 157 0.13–5.11 41–124 0.755 2.78 0.97

Sticheaus puncatatus 7 0.17–0.71 29–48 0.122 3.42 0.94

Reinhardtius hippoglossoides 9 400.20 –1,481.23 351–525 0.366 3.15 0.92

As part of this USGS study, the age and growth relationships for 19 marine fishes were reviewed and age-length and 

length-weight regressions are presented from collections in Arctic Alaska and adjacent seas. In certain instances, regressions 

are presented for data obtained from species collections in high-latitude areas far-removed from the Beaufort and Chukchi Seas, 

such as the Barents Sea. These examples highlight the limited availability of data for many species. In every case, the data are 

color-coded by investigator and area of fish collection. Because of the present-day ecological interest in Boreogadus saidi, the 

most current length-weight relationship described for this species from the Beaufort Sea, as indicated in table B1, is included for 

comparison with the historical data.
Appendix B 743

tac14-5222_appendix_b_01

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0 1 2 3 4 5 6 7 8 9 10

Mean fork length, in millimeters

Age, in years

0

100

200

300

400

500

600

0 50 100 150 200 250 300 350

Weight, in grams

Fork length, in millimeters

Wolotira and others (1977), Northeastern Bering Sea - females

Wolotira and others (1977), Northeastern Bering Sea - males

Wolotira and others (1977), Southeastern Chukchi Sea - females

Wolotira and others (1977), Southeastern Chukchi Sea - males

EXPLANATION

Wolotira and others (1977), Northeastern Bering Sea - Southeastern Chukchi Sea

Hopky and Ratynski (1983), Tuktoyaktuk Harbour

EXPLANATION

Pacific Herring

Clupea pallasi

Figure B1. Age-at-length and length-weight relationships for Pacific Herring (Clupea pallasii). Data from Wolotira 

and others (1977).
744 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_appendix_b_02

0

25

50

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100

125

150

0 1 2 3 4 5 6 7 8 9 10

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Age, in years

0

100

200

300

400

500

600

0 50 100 150 200 250 300 350

Weight, in grams

Fork length, in millimeters

DeGraaf (1974), Lake 100, Yukon Territory)

EXPLANATION

Chereshnev and others (1999), Sea of Okhotsk, Russia

DeGraaf and Machniak (1977), Mackenzie Delta

EXPLANATION

DeGraaf (1986), Lake 100, Yukon Territory

Pond Smelt

Hypomesus olidus

Figure B2. Age-at-length and length-weight relationships for Pond Smelt (Hypomesus olidus).
Appendix B 745

tac14-5222_appendix_b_03

70

80

90

100

110

120

130

140

150

160

170

0 1 2 3 4 5 6

Mean fork length, in millimeters

Age, in years

0

10

20

30

40

50

0 50 100 150 200

Weight, in grams

Fork length, in millimeters

Pahlke (1985), Southeastern Bering Sea and Kodiak Island

Fechhelm and others (1984), U.S. Chukchi Sea

Brown (2002), Gulf of Alaska

Winters (1970), Newfoundland - females

Winters (1970), Newfoundland - males

EXPLANATION

Naumenko (1996), U.S. Bering Sea - females

Naumenko (1996), U.S. Bering Sea - males

Fechhelm and others (1984), U.S. Chukchi Sea - females

Fechhelm and others (1984) U.S. Chukchi Sea - males

EXPLANATION

Capelin

Mallotus catervarius

Figure B3. Age-at-length and length-weight relationships for Capelin (Mallotus catervarius).
746 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_appendix_b_04

0

0

50

100

150

200

250

300

2 4 6 8 10 12 14

Mean fork length, in millimeters

Age, in years

0

100

200

300

400

500

600

0 50 100 150 200 250 300 350

Weight, in grams

Fork length, in millimeters

Fechhelm and others (1984), Northeastern Chukchi Sea

Wolotira and others (1977), Southeastern Chukchi Sea and Norton Sound

Haldorson and Craig (1984), Alaska Beaufort Sea

Bond and Erickson (1989), Phillips Bay, Yukon Territory

Bond and Erickson (1987), Phillips Bay, Yukon Territory - females

Bond and Erickson (1987), Phillips Bay, Yukon Territory - males

Karpenko and Vasilets (1996), Southwestern Bering Sea

Craig and Schmidt (1985), Point Lay, Chukchi Sea

Haldorson and Craig (1984), Alaska Beaufort Sea

Lawrence and others (1984), Southeastern Beaufort Sea

Bond and Erickson (1989), Phillips Bay, Yukon Territory

EXPLANATION

EXPLANATION

Arctic Smelt

Osmerus mordax

Figure B4. Age-at-length and length-weight relationships for Arctic Smelt (Osmerus dentex)
Appendix B 747

tac14-5222_appendix_b_05

0

50

100

150

200

250

300

350

400

450

500

0 2 4 6 8 10 12 14 16 18 20

Mean fork length, in millimeters

Age, in years

0

0

200

400

600

800

1,000

1,200

1,400

 50 100 150 200 250 300 350 400 450

Weight, in grams

Fork length, in millimeters

Schmidt and others (1991), Harrison Bay/Colville River

Schmidt and others (1991), Prudhoe Bay

Schmidt and others (1991), Phillips Bay

Schmidt and others (1991), McKinley Bay

Craig and Haldorson (1981), Simpson Lagoon

Palmer and Dugan (1990), Arctic National Wildlife Refuge

VanGerwen-Toyne and others (2008), Peel River

Bond (1982), Tuktoyaktuk Harbour - females

Bond (1982), Tuktoyaktuk Harbour - males

EXPLANATION

EXPLANATION

Arctic Cisco

Coregonus autumnalis

Figure B5. Age-at-length and length-weight relationships for Arctic Cisco (Coregonus autumnalis). Data from 

Schmidt and others (1991).
748 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_appendix_b_06

200

250

300

350

400

450

2 3 4 5 6 7 8 9 10

Mean fork length, in millimeters

Age, in years

0

200

400

600

800

1,000

0 50 100 150 200 250 300 350 400

Weight, in grams

Fork length, in millimeters

Craig and Haldorson (1981), Simpson Lagoon

Alt (1973b), Port Clarence - Grantley Harbor

Moulton (MJM Research, written commun., 2010), Colville River Region

Alt (1973b), Hess Creek

EXPLANATION

EXPLANATION

Bering Cisco

Coregonus laurettae

Figure B6. Age-at-length and length-weight relationships for Bering Cisco (Coregonus laurettae).
Appendix B 749

tac14-5222_appendix_b_07

0

 100

 200

 300

 400

 500

 600

0 5 10 15 20 25 30 35 40

Fork length, in millimeters

Age, in years

0

200

400

600

800

1,000

1,200

1,400

1,600

0 50 100 150 200 250 300 350 400 450

Weight, in grams

Fork length, in millimeters

Craig and Haldorson (1981), Simpson Lagoon

VanGerwen-Toyne and others (2008), Peel River

Percy (1975), Outer MacKenzie River

Muth (1969), Coppermine River

Philo and others (1993a), Dease Inlet Area

Philo and others (1993a), Colville River

Tallman and others (2002), Arctic Red River

Tallman and others (2002), Tuktoyaktuk Harbour

EXPLANATION

EXPLANATION

Broad Whitefish

Coregonus nasus

Figure B7. Age-at-length and length-weight relationships for Broad Whitefish (Coregonus nasus). 
750 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_appendix_b_08

0

 100

 200

 300

 400

 500

0 4 8 12 16 20 24 28

Mean fork length, in millimeters

Age, in years

0

400

800

1,200

1,600

2,000

2,400

0 100 200 300 400 500 600

Weight, in grams

Fork length, in millimeters

Philo and others (1993a), Dease Inlet/Admiralty Bay, Alaska

Craig and Haldorson (1981), Simpson Lagoon, Alaska

Brown (2004), Selawik River

Philo and others (1993a), Dease Inlet Area

Percy (1975), Outer MacKenzie River Delta

EXPLANATION

EXPLANATION

Humpback Whitefish

Coregonus pidschian

Figure B8. Age-at-length and length-weight relationships for Humpback Whitefish (Coregonus pidschian)
Appendix B 751

tac14-5222_appendix_b_09

100

200

300

400

0 5 10 15 20 25 30

Mean fork length, in millimeters

Age, in years

0

200

400

600

800

1,000

 50 100 150 200 250 300 350 400 450

Weight, in grams

Fork length, in millimeters

Philo and others (1993a), Teshekpuk Lake - females

Philo and others (1993a), Teshekpuk Lake - males

Craig and Haldorson (1981), Simpson Lagoon

Percy (1975), Outer MacKenzie River Delta

Brown (2004), Selawik River Delta

Philo and others (1996b), Dease inlet

Philo and others (1996a), Teshekpuk Lake

Philo and others (1996b), Colville River

Percy (1975), Outer MacKenzie River Delta

EXPLANATION

EXPLANATION

0

Least Cisco

Coregonus sardinella

Figure B9. Age-at-length and length-weight relationships for Least Cisco (Coregonus sardinella).
752 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_appendix_b_10

0

100

200

300

400

500

600

0 2 4 6 8 10 12 14

Mean fork length, in millimeters

Age, in years

0

1,000

2,000

3,000

4,000

5,000

0 100 200 300 400 500 600 700 800

Weight, in grams

Fork length, in millimeters

DeCicco (1996), Wulik River. Alaska - females

DeCicco (1996), Wulik River, Alaska- males

Wiswar (1994), Sadlerochit Spring, Alaska

Harwood and others (2009), Rat River, Northwest Territories - females

Underwood and others (1995), Arctic National Wildlife Refuge

EXPLANATION

EXPLANATION

Dolly Varden

Salvelinus malma

Figure B10. Age-at-length and length-weight relationships for Dolly Varden (Salvelinus malma). Age-length 

relationship not described from U.S. Beaufort and Chukchi Seas.
Appendix B 753

tac14-5222_appendix_b_11

0

25

50

75

100

125

150

0

10 20 30 40

Fork length (cm)

Age, in years

Inconnu (Stenodus leucichthys)

0

4

8

12

16

40 50 60 70 80 90 100

Weight, in kilograms

Standard length, in centimeters

Alt (1969), Selawik River - females

Alt (1969), Selawik River - males

Alt (1969), Kobuk River - females

Alt (1969), Kobuk River - males

Alt (1973b), Kuskokwim River

Alt (1973b), Lower - Yukon River

Alt (1973b), Upper - Yukon River

Alt (1973b), Minto Flats

Alt (1973b), Kobuk - Selowik

Howland and others (2004), MacKenzie River

EXPLANATION

EXPLANATION

Figure B11. Age-at-length and length-weight relationships for Inconnu (Stenodus leucichthys).
754 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_appendix_b_12

0

50

100

150

200

250

300

0 1 2 3 4 5 6 7 8 9 10

Mean fork length, in millimeters

Age, in years

0

25

50

75

100

125

150

0 25 50 75 100 125 150 175 200 225 250

Weight, in grams

Fork length, in millimeters

Gillispie and others (1997), Northeastern Chukchi Sea

Frost and Lowry (1981b), Northeastern Chukchi and Western Beaufort Seas

Craig and Haldorson (1981), Simpson Lagoon

Fruge and others (1989), Arctic National Wildlife Refuge

Palmer and Dugan (1990), Arctic National Wildlife Refuge

Kelly L. Walker, University of Alaska, Fairbanks, written commun., 2016

Falk-Petersen and others (1986), Spitsbergen

Hognestad (1968), Barents Sea

Craig and others (1982), Northeastern Chukchi Sea

Gillispie and others (1997), Northeastern Chukchi Sea

Palmer and Dugan (1990), Arctic National Wildlife Refuge

Hop and others (1997), Canadian High Arctic

EXPLANATION

EXPLANATION

Arctic Cod

Boreogadus saida

Figure B12. Age-at-length and length-weight relationships for Arctic Cod (Boreogadus saida).
Appendix B 755

tac14-5222_appendix_b_13

0

 50

 100

 150

 200

 250

 300

 350

 400

0 2 4 6 8 10

Mean fork length, in millimeters

Age, in years

0

100

200

300

400

0 50 100 150 200 250 300 350

Weight, in grams

Fork length, in millimeters

Wolotira and others (1977), U.S.Bering Sea

Fechhelm and others (1984), Northeastern Chukchi Sea

Hopky and Ratynski (1983), Tuktoyaktuk Harbour, Northwest Territories

Chen and Sakurai (1993), Hokkaido - Pacific Waters

Wolotira and others (1977), Northern Bering Sea - females

Wolotira and others (1977), Northern Bering Sea - males

Wolotira and others (1977), Southeastern Chukchi Sea - females

Wolotira and others (1977), Southeastern Chukchi Sea - males

EXPLANATION

EXPLANATION

Saffron Cod

Eleginus gracilis

Figure B13. Age-at-length and length-weight relationships for Saffron Cod (Eleginus gracilis).
756 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_appendix_b_14

0

100

200

300

400

500

600

700

800

0 50 100 150 200 250 300

Weight, in grams

Standard length, in millimeters

 25

 50

 75

 100

 125

 150

0

0 1 2 3 4 5 6 7 8 9 10 11 12 13

Standard length, in millimeters

Age, in years

Smith and others (1997a), Northern Chukchi Sea

Smith and others (1997a), Northern Chukchi Sea - females

Smith and others (1997a), Northern Chukchi Sea - males

EXPLANATION

EXPLANATION

Arctic Staghorn Sculpin

Gymnocanthus tricuspis

Figure B14. Age-at-length and length-weight relationships for Arctic Staghorn Sculpin (Gymnocanthus tricuspis).
Appendix B 757

tac14-5222_appendix_b_15

0

50

100

150

200

250

300

350

400

450

0 50 100 150 200 250 300

Weight, in grams

Total length, in millimeters

54

56

58

60

2 3 4 5 6

Mean total length, in millimeters

Age, in years

Frost and Lowry (1983), U.S. Beaufort Sea

Frost and Lowry (1983), U.S. Beaufort Sea

EXPLANATION

EXPLANATION

Twohorn Sculpin

Icelus bicornis

Figure B15. Age-at-length and length-weight relationships for Twohorn Sculpin (Icelus bicornis).
758 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_appendix_b_16

0

100

200

300

400

500

600

0 50 100 150 200 250 300 350

Weight, in grams

Total length, in millimeters

0

50

100

150

200

250

300

0 2 4 6 8 10 12 14 16

Mean total length, in millimeters

Age, in years

Fechhelm and others (1984), Northeastern Chukchi Sea

Craig and Haldorson (1981), Simpson Lagoon

Griffiths and others (1975), Nunaluk Lagoon, Yukon Territory

Bond and Erickson (1987), Phillips Bay, Yukon Territory - females

Bond and Erickson (1987), Phillips Bay, Yukon Territory - males

Craig and Haldorson (1981), Simpson Lagoon

Underwood and others (1995), Arctic National Wildlife Refuge

Griffiths and others (1975), Nunaluk Lagoon, Yukon Territory - females

Griffiths and others (1975), Nunaluk Lagoon, Yukon Territory - males

Jones and Den Beste (1977), Tuktoyaktuk Peninsula

EXPLANATION

EXPLANATION

Fourhorn Sculpin

Myoxocephalus quadricornis

Figure B16. Age-at-length and length-weight relationships for Fourhorn Sculpin (Myoxocephalus quadricornis).
Appendix B 759

tac14-5222_appendix_b_17

0

100

200

300

400

500

600

0 50 100 150 200 250 300

Weight, in grams

Standard length, in millimeters

0

 50

 100

 150

 200

0 1 2 3 4 5 6 7 8 9 10 11 12 13

Standard length, in millimeters

Age, in years

Smith and others (1997), Northeastern Chukchi Sea

Smith and others (1997), Northeastern Chukchi Sea - females

Smith and others (1997), Northeastern Chukchi Sea - males

EXPLANATION

EXPLANATION

Bering Flounder

Hippoglossoides robustus

Figure B17. Age-at-length and length-weight relationships for Bering Flounder (Hippoglossoides robustus).
760 Alaska Arctic Marine Fish Ecology Catalog

tac14-5222_appendix_b_18

0

500

1,000

1,500

2,000

2,500

3,000

0

50 100 150 200 250 300 350 400 450 500 550

Weight, in grams

Total length, in millimeters

 150

 200

 250

 300

 350

 400

 450

 500

 550

0 4 8 12 16 20 24 28

Mean total length, in millimeters

Age, in years

Kolpakov (2005a), Sea of Japan - females

Kolpakov (2005a), Sea of Japan - males

Hopky and Ratynski (1983), Tuktoyaktuk Harbour, Northwest Territories

Wildermuth (1983), Puget Sound)

Kolpakov (2005a), Sea of Japan - females

Kolpakov (2005a), Sea of Japan - males

Lawrence and others (1984), Southeastern Beaufort Sea

Orcutt (1950), Central California - females

Orcutt (1950), Central California - males

EXPLANATION

EXPLANATION

Starry Flounder

Platichthy stellatus

Figure B18. Age-at-length and length-weight relationships for Starry Flounder (Platichthy stellatus).
Appendix B 761

tac14-5222_appendix_b_19a

0

50

100

150

200

250

300

350

0 5 10 15 20 25 30

Mean total length, in millimeters

Age, in years

EXPLANATION

Kobelev (1989), Barents Sea - females

Kobelev (1989), Barents Sea - males

Fechhelm and others (1984), Northeast Chukchi Sea - females

Palmer and Dugan (1990), Arctic National Wildlife Refuge

Bond and Erickson (1989), Yukon Coast

Percy (1975), MacKenzie River Delta

Lawrence and others (1984), Tuktoyaktuk Harbour

Arctic Flounder

Lioposetta glacialis

Figure B19. Age-at-length and length-weight relationships for Arctic Flounder (Lioposetta glacialis) from the 

Chukchi, Beaufort, and Barents Seas.

0

100

200

300

400

500

600

0 50 100 150 200 250 300 350 400

Weight, in grams

Total length, in millimeters

EXPLANATION

Kobelev (1989), Barents Sea - females

Kobelev (1989), Barents Sea - males

Fruge and others (1989), Arctic National Wildlife Refuge

Palmer and Dugan (1990), Arctic National Wildlife Refuge

Griffiths and others (1975), Yukon Coast

Arctic Flounder

Lioposetta glacialis
762 Alaska Arctic Marine Fish Ecology Catalog

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Appendix C. Models Evaluated for Simulating Effects of Climate Change on the

Distributions and Abundances of Arctic Cod and Saffron Cod in the Eastern 

Bering Sea

Table C1. Candidate models evaluated in search of a model to simulate the distribution (presence/absence) of Arctic Cod in the eastern 

Bering Sea using a smooth (s) GAM model.

[Abbreviations: REML, restricted maximum likelihood; r2, coefficient of determination; n, number]

Arctic Cod (Boreogadus saida) distribution models

gam(Boreogadus saida) family=binomial,method=”REML” smooth (s) models

Model

Akaike 

Information 

Criterion

Deviance

(percent)

REML 

score

r2 n FALSE TRUE

s(GEAR_TEMPERATURE,BOTTOM_

DEPTH,SURFACE_TEMPERATURE)

2,781.838 54.70 1,425.3 0.524 9,423 FALSE 8,349 339

TRUE 173 562

s(GEAR_TEMPERATURE,BOTTOM_DEPTH) 2,941.793 51.30 1,504.3 0.484 9,423 FALSE 8,343 377

TRUE 179 524

s(GEAR_TEMPERATURE,SURFACE_

TEMPERATURE)

2,976.595 50.80 1,515.9 0.493 9,423 FALSE 8,356 388

TRUE 166 513

s(GEAR_TEMPERATURE) + s(BOTTOM_

DEPTH) + s(SURFACE_TEMPERATURE)

2,797.761 53.50 1,419.1 0.511 9,423 FALSE 8,357 365

TRUE 165 536

s(GEAR_TEMPERATURE) + s(BOTTOM_

DEPTH)

2,946.398 50.90 1,488.9 0.482 9,423 FALSE 8,355 388

TRUE 167 513

s(GEAR_TEMPERATURE) + s(SURFACE_

TEMPERATURE)

3,129.55 47.90 1,584.6 0.466 9,432 FALSE 8,366 419

TRUE 156 482

s(BOTTOM_DEPTH) + s(SURFACE_

TEMPERATURE)

5,484.76 8.15 2,759.9 0.052 9,432 FALSE 8,521 899

TRUE 1 2
764 Alaska Arctic Marine Fish Ecology Catalog

Table C2. Candidate models evaluated in search of a model to predict the distribution of Arctic Cod in the eastern Bering Sea using a 

tensor smooth and (or) interaction smooth GAM models.

[Abbreviations: REML, restricted maximum likelihood; r2, coefficient of determination; n, number]

Arctic Cod (Boreogadus saida) distribution models

Tensor product smooths (te) and Tensor product Interaction (ti) models 

gam(Boreogadus saida) family=binomial,method=”REML”

Model

Akaike 

Information 

Criterion

Deviance

(percent)

REML 

score

r2 n FALSE TRUE

te(GEAR_TEMPERATURE, BOTTOM_

DEPTH, SURFACE_TEMPERATURE)1

2,663.922 56.5 1,338 0.545 9,423 FALSE 8,367 314

TRUE 155 587

te(GEAR_TEMPERATURE, BOTTOM_

DEPTH) + s(SURFACE_TEMPERATURE) 

2,782.59 53.8 1,402.3 0.515 9,423 FALSE 8,349 351

TRUE 173 550

te(GEAR_TEMPERATURE, BOTTOM_

DEPTH)

2,915.461 51.4 1,463.2 0.487 9,423 FALSE 8,344 370

TRUE 178 531

te(GEAR_TEMPERATURE, SURFACE_

TEMPERATURE) + s(BOTTOM_DEPTH) 

2,740.1 54.6 1,386.5 0.528 9,423 FALSE 8,342 326

TRUE 180 575

te(GEAR_TEMPERATURE, SURFACE_

TEMPERATURE) + te(GEAR_

TEMPERATURE, BOTTOM_DEPTH)

2,708 55.3 1,377 0.534 9,423 FALSE 8,355 328

TRUE 167 573

ti(GEAR_TEMPERATURE, BOTTOM_DEPTH, 

SURFACE_TEMPERATURE)

4,473.568 26.2 2,340.7 0.307 9,423 FALSE 8,428 625

TRUE 94 276

ti(GEAR_TEMPERATURE, BOTTOM_

DEPTH) +te(SURFACE_TEMPERATURE)

3,452 42.50 1,766.4 0.447 9,423 FALSE 8,376 426

TRUE 146 475

1

Selected as the best model based on model fit, low Akaike Information Criterion (AIC) score and the percent of deviance explained by the model.
Appendix C 765

Table C3. Candidate models evaluated in search of a model to predict abundance of Arctic Cod in the eastern Bering Sea using a 

smooth (s) GAM model.

[Abbreviations: REML, restricted maximum likelihood; r2, coefficient of determination; n, number]

Arctic Cod (Boreogadus saida) abundance models

gam(ln(abundance +1)) method=”REML” smooth (s) models

Model

Akaike 

Information 

Criterion

Deviance

(percent)

REML 

score

r2 n

s(GEAR_TEMPERATURE,BOTTOM_DEPTH, SURFACE_TEMPERATURE) 20,072.9 51 10,190 0.505 9,423

s(GEAR_TEMPERATURE) + s(BOTTOM_DEPTH) + s(SURFACE_TEMPERATURE) 19,500.69 53.25 9,794.7 0.531 9,423

s(GEAR_TEMPERATURE) + s(BOTTOM_DEPTH) 19,618 52.50 9,846.2 0.524 9,423

s(GEAR_TEMPERATURE,BOTTOM_DEPTH) 23,235.07 30.50 11,671 0.303 9,423

s(GEAR_TEMPERATURE) + s(SURFACE_TEMPERATURE) 19,741.89 51.90 9,904.7 0.518 9,423

s(GEAR_TEMPERATURE,SURFACE_TEMPERATURE) 19,530.79 53.10 9,828.3 0.529 9,423

s(BOTTOM_DEPTH) + s(SURFACE_TEMPERATURE) 26,186.95 4.64 13,120 0.0448 9,423

Table C4. Candidate models evaluated in search of a model to predict abundance of Arctic Cod in the eastern Bering Sea using a tensor 

smooth and (or) interaction smooth GAM model.

[Abbreviations: REML, restricted maximum likelihood; r2, coefficient of determination; n, number]

Arctic Cod (Boreogadus saida) abundance models

Tensor product smooths (te) and Tensor product Interaction (ti) models

gam(ln(abundance +1)) method=”REML”

Model

Akaike 

Information 

Criterion

Deviance

(percent)

REML 

score

r

2 n

te(GEAR_TEMPERATURE, BOTTOM_DEPTH, SURFACE_TEMPERATURE)1 18,772.98 57.10 9,501.1 0.568 9,423

ti(GEAR_TEMPERATURE, BOTTOM_DEPTH, SURFACE_TEMPERATURE) 24,648.37 19.50 12,413 0.191 9,423

ti(GEAR_TEMPERATURE, BOTTOM_DEPTH)+ te(SURFACE_TEMPERATURE) 21,108.77 44.40 10,600 0.443 9,423

ti(GEAR_TEMPERATURE, SURFACE_TEMPERATURE)+ te(BOTTOM_DEPTH) 25,968.32 6.91 13,047 0.0672 9,432

te(GEAR_TEMPERATURE, SURFACE_TEMPERATURE)+ te(BOTTOM_DEPTH) 20,081.09 50.20 10,096 0.501 9,432

te(GEAR_TEMPERATURE) 20,833.78 45.80 10,428 0.458 9,432

te(SURFACE_TEMPERATURE) 26,546.83 0.68 13,281 0.00643 9,432

te(BOTTOM_DEPTH) 26,450.3 1.69 13,229 0.0165 9,423

1

Selected as the best model based on model fit, low Akaike Information Criterion (AIC) score and the percent of deviance explained by the model.
766 Alaska Arctic Marine Fish Ecology Catalog

Table C5. Candidate models evaluated in search of a model to predict distribution of Saffron Cod in the eastern Bering Sea using a 

smooth (s) GAM model.

[Abbreviations: REML, restricted maximum likelihood; r2, coefficient of determination; n, number]

Saffron Cod (Eleginus gracilis) distribution models

Models gam(Eleginus gracilis) family=binomial,method=”REML” smooth(s) models

Model

Akaike 

Information 

Criterion

Deviance

(percent)

REML 

score

r2 n FALSE TRUE

s(GEAR_TEMPERATURE,BOTTOM_DEPTH, 

SURFACE_TEMPERATURE)1

2,026.113 51.60 1,022.6 0.422 9,423 FALSE 8,794 298

TRUE 98 233

s(GEAR_TEMPERATURE,BOTTOM_DEPTH) 2,103.427 49.20 1,063.4 0.388 9,423 FALSE 8,798 316

TRUE 94 215

s(GEAR_TEMPERATURE,SURFACE_

TEMPERATURE)

2,628.688 36.70 1,329.5 0.325 9,423 FALSE 8,823 351

TRUE 69 180

s(GEAR_TEMPERATURE) + s(BOTTOM_

DEPTH) + s(SURFACE_TEMPERATURE)2

2,048.516 50.60 1,035.1 0.417 9,423 FALSE 8,796 286

TRUE 96 245

s(GEAR_TEMPERATURE) + 

s(BOTTOM_DEPTH)

2,134.97 48.10 1,072.6 0.38 9,423 FALSE 8,804 322

TRUE 88 209

s(GEAR_TEMPERATURE) + 

s(SURFACE_TEMPERATURE)

2,662.401 35.50 1,346.5 0.312 9,423 FALSE 8,829 362

TRUE 63 169

s(BOTTOM_DEPTH) + 

s(SURFACE_TEMPERATURE)

2,061.879 50.00 1,039.2 0.408 9,423 FALSE 8,780 285

TRUE 112 246

s(BOTTOM_DEPTH, SURFACE_

TEMPERATURE)

2,069.542 50.00 1,044.4 0.398 9,423 FALSE 8,795 309

TRUE 97 222

1

Identified as a potential best model.

2

Selected as the best model after reviewing the environmental response curves for the other models even though the Akaike Information Criterion (AIC) is 

higher and the deviance explained was lower.
Appendix C 767

Table C6. Candidate models evaluated in search of a model to predict distribution of Saffron Cod in the eastern Bering Sea using a 

tensor smooth and (or) interaction smooth GAM model.

[Abbreviations: REML, restricted maximum likelihood; r2, coefficient of determination; n, number]

Saffron Cod (Eleginus gracilis) distribution models

gam(Eleginus gracilis) family=binomial,method=”REML” tensor product smooths (te) and Tensor product Interaction (ti) models

Model

Akaike 

Information 

Criterion

Deviance

(percent)

REML 

score

r2 n FALSE TRUE

te(GEAR_TEMPERATURE, BOTTOM_DEPTH, SURFACE_

TEMPERATURE)1

1,953.825 53.80 967.42 0.45 9,423 FALSE 8,814 286

TRUE 78 245

te(BOTTOM_DEPTH, SURFACE_TEMPERATURE) 2,041.313 50.60 1,022.7 0.414 9,423 FALSE 8,780 287

TRUE 112 244

te(BOTTOM_DEPTH) + te(SURFACE_TEMPERATURE) 2,074.152 49.60 1,040.5 0.402 9,423 FALSE 8,784 305

TRUE 108 226

te(GEAR_TEMPERATURE, BOTTOM_DEPTH) + 

s(SURFACE_TEMPERATURE)

2,030.963 51.10 1,017.7 0.422 9,423 FALSE 8,805 295

TRUE 87 236

te(GEAR_TEMPERATURE, BOTTOM_DEPTH) 2,109.372 48.90 1,051.7 0.387 9,423 FALSE 8,809 329

TRUE 83 202

te(GEAR_TEMPERATURE, SURFACE_TEMPERATURE) + 

s(BOTTOM_DEPTH)

1,990.718 52.20 1,003.1 0.44 9,423 FALSE 8,806 281

TRUE 86 250

te(GEAR_TEMPERATURE, SURFACE_TEMPERATURE) + 

te(GEAR_TEMPERATURE, BOTTOM_DEPTH)

1,964.257 53.10 998.51 0.446 9,423 FALSE 8,813 288

TRUE 79 243

ti(GEAR_TEMPERATURE, BOTTOM_DEPTH, SURFACE_

TEMPERATURE)

2,564.903 38.70 1,316.4 0.381 9,423 FALSE 8,827 320

TRUE 65 211

ti(GEAR_TEMPERATURE, BOTTOM_DEPTH) + 

te(SURFACE_TEMPERATURE)

2,365.397 42.90 1,203.3 0.375 9,423 FALSE 8,810 326

TRUE 82 205

ti(BOTTOM_DEPTH, SURFACE_TEMPERATURE) + 

s(BOTTOM_DEPTH)

2,028.882 50.90 1,021.3 0.416 9,423 FALSE 8,772 281

TRUE 120 250

1

Identified as a potential best model.
768 Alaska Arctic Marine Fish Ecology Catalog

Table C7. Candidate models evaluated in search of a model to predict abundance of Saffron Cod in the eastern Bering Sea using a 

smooth (s) GAM model.

[Abbreviations: REML, restricted maximum likelihood; r2, coefficient of determination; n, number]

Saffron Cod (Eleginus gracilis) abundance models

gam(ln(abundance +1)) method=”REML” smooth (s) models

Model

Akaike 

Information 

Criterion

Deviance

(percent)

REML 

score

r2 n

s(GEAR_TEMPERATURE,BOTTOM_DEPTH, SURFACE_TEMPERATURE) 18,888.36 56 9,597.5 0.55 9,423

s(GEAR_TEMPERATURE) + s(BOTTOM_DEPTH) + s(SURFACE_TEMPERATURE) 19,871.85 49.90 9,997.6 0.497 9,423

s(GEAR_TEMPERATURE) + s(BOTTOM_DEPTH) 20,658.18 45.40 10,370 0.453 9,423

s(GEAR_TEMPERATURE,BOTTOM_DEPTH) 20,606.26 45.80 10,367 0.457 9,423

s(GEAR_TEMPERATURE) + s(SURFACE_TEMPERATURE) 21,309.22 41.50 10,699 0.414 9,423

s(GEAR_TEMPERATURE,SURFACE_TEMPERATURE) 20,304.42 47.50 10,219 0.474 9,423

s(BOTTOM_DEPTH) + s(SURFACE_TEMPERATURE) 20,783.68 44.70 10,434 0.446 9,423

Table C8. Candidate models evaluated in search of a model to predict abundance of Saffron Cod in the eastern Bering Sea using a 

tensor smooth and (or) interaction smooth GAM models.

[Abbreviations: REML, restricted maximum likelihood; r2, coefficient of determination; n, number]

Saffron Cod (Eleginus gracilis) abundance models

gam(ln(abundance + 1)) method=”REML” tensor product smooths (te) and Tensor product Interaction (ti) models

Model

Akaike 

Information 

Criterion

Deviance

(percent)

REML 

score

r2 n

te(GEAR_TEMPERATURE, BOTTOM_DEPTH, SURFACE_TEMPERATURE)1 17,389.91 61.80 8,795.9 0.615 9,423

ti(GEAR_TEMPERATURE, BOTTOM_DEPTH, SURFACE_TEMPERATURE) 18,566.81 56.50 9,408.1 0.563 9,423

ti(GEAR_TEMPERATURE, BOTTOM_DEPTH)+ te(SURFACE_TEMPERATURE) 20,012.76 49.00 10,046 0.489 9,423

ti(GEAR_TEMPERATURE, SURFACE_TEMPERATURE)+ te(BOTTOM_DEPTH) 20,115.11 48.50 10,135 0.484 9,423

te(GEAR_TEMPERATURE, SURFACE_TEMPERATURE)+ te(BOTTOM_DEPTH) 19,389.26 52.40 9,755 0.522 9,423

te(GEAR_TEMPERATURE) 22,310.26 34.70 11,165 0.347 9,423

te(SURFACE_TEMPERATURE) 25,199.78 11.30 12,612 0.113 9,423

te(BOTTOM_DEPTH) 22,614.04 32.60 11,319 0.326 9,423

1

Selected as the best model based on model fit, low Akaike Information Criterion (AIC) score and the percent of deviance explained by the model.
Publishing support provided by the U.S. Geological Survey

Science Publishing Network, Tacoma Publishing Service Center

For more information concerning the research in this report, contact the

 Regional Director, Alaska

U.S. Geological Survey

4210 University Drive

Anchorage, Alaska 99508-4560

https://www.usgs.gov/science/regions/alaska-region
 Thorsteinson and Love

—Alaska Arctic Marine Fish Ecology Catalog

—Scientific Investigations Report 2016–5038

ISSN 2328-0328 (online)

http://dx.doi.org/10.3133/sir20165038