NOAA Professional Paper 15

Guide to the identifi cation

of larval and early juvenile 

pricklebacks (Perciformes: 

Zoarcoidei: Stichaeidae)

in the northeastern Pacifi c 

Ocean and Bering Sea

Ann C. Matarese 

Deborah M. Blood

Morgan S. Busby

October 2013
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Kathryn Dennis

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NOAA Professional Paper 15

Guide to the identifi cation of 

larval and early juvenile pricklebacks

(Perciformes: Zoarcoidei: Stichaeidae) 

in the northeastern Pacifi c Ocean 

and Bering Sea

Ann C. Matarese

Deborah M. Blood

Morgan S. Busby

October 2013

U.S. Department of Commerce

Seattle, Washington
Suggested reference

Matarese, Ann C., Deborah M. Blood, and Morgan S. Busby. Guide to the identifi -

cation of larval and early juvenile pricklebacks (Perciformes: Zoarcoidei: Stichaeidae) in the northeastern Pacifi c Ocean and Bering Sea. NOAA Professional Paper 

NMFS 15, 96 p. doi:10.7755/PP.15

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publication.
CONTENTS

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

Overall comparisons . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 Family . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 Subfamilies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Subfamily Stichaeinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

 Stichaeus punctatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Subfamily Chirolophinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

 Bryozoichthys lysimus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

 Bryozoichthys marjorius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

 Chirolophis decoratus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

 Chirolophis nugator . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

 Gymnoclinus cristulatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28

Subfamily Lumpeninae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

 Lumpenella longirostris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30

 Lumpenus fabricii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

 Lumpenus maculatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

 Lumpenus medius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

 Lumpenus sagitta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

 Poroclinus rothrocki . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

Subfamily Opisthocentrinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

 Opisthocentrus ocellatus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

 Plectobranchus evides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56

Subfamily Xiphisterinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59

 Alectrias spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59

 Anoplarchus insignis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

 Anoplarchus purpurescens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

 Cebidichthys violaceus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

Other material . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

 Larval characters: relationships within family Stichaeidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

 Larval characters: relationships within suborder Zoarcoidei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

Literature cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

Appendix . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

iii

1

Guide to the identifi cation of larval and early 

juvenile pricklebacks (Perciformes: Zoarcoidei: 

Stichaeidae) in the northeastern Pacifi c Ocean 

and Bering Sea

Ann C. Matarese (contact author)

Deborah M. Blood

Morgan S. Busby

Recruitment Processes Program

Resource Assessment and Conservation Engineering Division

Alaska Fisheries Science Center

National Marine Fisheries Service

National Oceanic and Atmospheric Administration

7600 Sand Point Way NE

Seattle, Washington 98115

Email address for contact author: ann.matarese@noaa.gov

Abstract—Stichaeidae, commonly 

referred to as pricklebacks, are intertidal and subtidal fishes primarily 

of the North Pacific Ocean. Broad 

distribution in relatively inaccessible and undersampled habitats has 

contributed to a general lack of information about this family. In this 

study, descriptions of early life history stages are presented for 25 species 

representing 18 genera of stichaeid 

fishes from the northeastern Pacific 

Ocean, Bering Sea, and Arctic Ocean 

Basin. Six of these species also occur 

in the North Atlantic Ocean. Larval 

stages of 16 species are described for 

the first time. Additional information 

or illustrations intended to augment 

previous descriptions are provided 

for nine species. For most taxa, we 

present adult and larval distributions, descriptions of morphometric, 

meristic, and pigmentation characters, and species comparisons, and 

we provide illustrations for preflexion through postflexion or transformation stages. New counts of meristic features are reported for several 

species.

Introduction

Stichaeidae, commonly referred to as 

pricklebacks, are intertidal and subtidal fi shes primarily of the North Pacifi c Ocean and Bering Sea; a few species also are found in the Arctic and 

North Atlantic oceans (Table 1). They 

are commonly found under rocks and 

among algae in the intertidal zone or 

in shallow bays and reported to occur 

at depths ≥250 m on the outer continental shelf and upper slope (Mecklenburg and Sheiko, 2004). High diversity (76 species in 37 genera) and 

broad distribution in relatively inaccessible and undersampled habitats 

have contributed to a general lack 

of information about this family. Although not commercially important, 

stichaeids are important in ecological 

interactions as forage for commercial 

and noncommercial fishes, marine 

birds, and marine mammals of the 

ecosystems of the North Pacifi c Ocean 

and Bering Sea (Golet et al., 2000). 

Information about stichaeid early life 

history is essential to our understanding of recruitment and survival of intertidal and nearshore fi shes.

Stichaeids have elongate, slightly 

compressed bodies with numerous 

sharp spines in the dorsal fi n. They 

are related most closely to the Pholidae, Zoarcidae, and other zoarcoid 

fi shes but can be distinguished by a 

combination of the following characters: spinous dorsal fi n continuous 

along the entire body or with some 

soft rays posteriorly; anal fi n with 0–5 

anterior spines; pelvic fi n, when present, that has one spine and up to four 

rays; teeth simple; and ribs, intercalar 

bone, and bony actinosts (proximal 

pectoral radials) present (Anderson, 

1984, 1994). Historically, Makushok 

(1958) divided Stichaeidae into eight 

subfamilies. Although Makushok’s 

(1958) comprehensive study has been 

accepted for many years, providing insight into character evolution, 

identification, and classification of 

known stichaeid species, not all taxa 

were examined in that study. The current classifi cation of Stichaeidae with 

six subfamilies results from cladistic 
2 Professional Paper NMFS 15 

Table 1

Distribution of Stichaeidae in the Bering Sea and northeastern Pacifi c Ocean with the current status of early life history 

descriptions based on this study.

 Number

Taxon Distribution1 of species Published Updated New

STICHAEINAE 2

Eumesogrammus praecisus Sea of Okhotsk; Beaufort–Bering Seas; X2

 Arctic–W North Atlantic 

Stichaeus punctatus Okhotsk and Japan Seas; Beaufort Sea– Faber (1976); Farwell et al. X

 North Pacifi c; Arctic–W North Atlantic (1976); Haryu et al. 

 (1985); Fahay (1983, 2007) 

CHIROLOPHINAE 7

Bryozoichthys lysimus Sea of Okhotsk–N Sea of Japan; Shiogaki and Haryu X3

 Bering Sea–N Gulf of Alaska (1988)

Bryozoichthys marjorius Aleutian Is.–S British Columbia X

Chirolophis decoratus E Bering Sea, Aleutian Is.–N Calif. X

Chirolophis nugator Aleutian Is.–S Calif. X

Chirolophis snyderi W Bering Sea–Sea of Okhotsk; Grigor’ev (1992a) X

 Chukchi Sea–W Gulf of Alaska 

Chirolophis tarsodes North Pacifi c and Kodiak Is.–

 N British Columbia

Gymnoclinus cristulatus Hokkaido; Commander Is.– Tokuya and Amaoka X

 Amchitka Island (1980); Shiogaki (1988)

LUMPENINAE 7

Acantholumpenus mackayi Okhotsk and Japan Seas–Hokkaido; X

 Beaufort Sea–NE Gulf of Alaska 

Lumpenella longirostris Sea of Okhotsk–N Japan; Bering Sea– Matarese et al. (1989) X3

 S British Columbia; Greenland 

Lumpenus fabricii Okhotsk and Japan Seas; Beaufort Sea– Grigor’ev (1992b) X3

 North Pacifi c; Arctic–North Atlantic 

Lumpenus maculatus Okhotsk and Japan Seas; Beaufort– Faber (1976); X3

 North Pacifi c (Washington); North Atlantic Grigor’ev (1992b) 

Lumpenus medius Okhotsk and Japan Seas; Beaufort Sea– Grigor’ev (1992b) X3

 North Pacifi c; Arctic–North Atlantic 

Lumpenus sagitta Okhotsk and Japan Seas; Shiogaki and Sasaki (1988); X

 Bering Sea, Aleutian Is.–N Calif. Matarese et al. (1989) 

Poroclinus rothrocki SE Bering Sea, Aleutian Is.–S Calif. Matarese et al. (1989) X3

OPISTHOCENTRINAE 3

Lumpenopsis hypochroma S British Columbia–S Calif. X2

Opisthocentrus ocellatus Okhotsk and Japan Seas–North Korea; Tokuya and Amaoka (1980); X

 Kuril Is.–Commander Is. Shiogaki (1982) 

Plectobranchus evides British Columbia–S Calif. Matarese et al. (1989) X

XIPHISTERINAE 8

Alectridium aurantiacum Commander Is.; Bering Sea, Aleutian Is., X2

 and N Gulf of Alaska 

Alectrias alectrolophus Sea of Okhotsk–N Sea of Japan; X

 Norton Sound (Bering Sea) 

Anoplarchus insignis SE Bering Sea, Aleutian Is.–N Calif. X

Anoplarchus purpurescens SE Bering Sea, Aleutian Is.–S Calif. Wang (1986); X

 Matarese et al. (1989)

Cebidichthys violaceus Oregon–Baja Calif. Wang (1986) X3

Phytichthys chirus Aleutian Is., Gulf of Alaska–S Calif. Matarese et al. (1989) X2

Xiphister atropurpureus W Gulf of Alaska–Baja Calif. Matarese et al. (1989) X2

Xiphister mucosus W Gulf of Alaska–S Calif. Wang (1986); 

 Matarese et al. (1989)

 TOTALS 27 17 9 16

Early life history

1 Mecklenberg et al. (2002).

2 Identifi cation is tentative.

3 Previously published illustration or description incomplete or limited to one early life history stage (prefl exion, fl exion, postfl exion).
3

studies that focus on the Xiphisterinae (Stoddard, 1985; 

Yatsu, 1986). However, more work is required to demonstrate the monophyly of this family and its subfamilies 

and to better defi ne genera and species (Mecklenburg 

and Sheiko, 2004). Recent molecular work has linked 

the stichaeids and other zoarcoids with cottoids (e.g., 

anoplopomatids, hexagrammids, and cottids) (Smith and 

Wheeler, 2004). Identifi cation of a suite of early life history characters may provide additional insight into the 

classifi cation of stichaeids and enable higher-level studies 

of stichaeid and zoarcoid phylogeny.

Early life history stages are known for only 55% of 

stichaeid species that occur in the northeastern Pacifi c 

(NEP). In taxa for which information is available, eggs 

are demersal, spherical, 1.3–2.5 mm in diameter, contain 

one or more oil globules, and form an adhesive egg mass 

that is guarded by the male or female parent (Fujita and 

Uchida, 1959; Shiogaki and Dotsu, 1972; Wourms and 

Evans, 1974a, 1974b; Marliave and DeMartini, 1977; 

Shiogaki, 1981, 1982, 1983; Watson, 1996). Eggs are not 

well known and were not considered in the study presented here. Incomplete developmental series have been 

published for some species (Table 1), but larvae of the 

majority of taxa remain unknown. In this study, we describe the early life history stages of 16 species for the 

fi rst time and provide updated information for nine additional species (Table 1). 

Materials and methods

The study area extends from the Beaufort Sea to the west 

coast of the United States, including the Chukchi Sea, 

Bering Sea, and Gulf of Alaska, as well as the southern 

coast of British Columbia (Fig. 1). Historical ichthyoplankton data were obtained from surveys conducted by 

the Recruitment Processes Program of the Alaska Fisheries Science Center (AFSC), National Marine Fisheries 

Service. Data for cruises prior to 1988 were found in 

Dunn and Rugen,1 and those data from 1989 to 2009 

were found in the AFSC Ichthyoplankton Cruise Database ( http://access.afsc.noaa.gov/icc/index.cfm). Ichthyoplankton data were accessed in ICHBASE, the larval fi sh 

database maintained by and available from the AFSC. 

Most ichthyoplankton were collected with a bongo 

sampler (Posgay and Marak, 1980), with nets that had an 

inside diameter of 0.6 m and mesh of 0.505 mm or 0.333 

mm, similar to that of the Marine Resources Monitoring, 

Assessment, and Prediction Program (MARMAP). Bongo 

nets were lowered at a rate of 40 m/min and retrieved at 

a rate of 20 m/min, sampling obliquely from the surface 

1 Dunn, J. R., and W. C. Rugen. 1989. A catalog of Northwest 

and Alaska Fisheries Center ichthyoplankton cruises 1965–1988. 

Northwest and Alaska Fish.Cent. Natl. Mar. Fish. Serv., NOAA, 

7600 Sand Point Way NE, Seattle WA 98115. NWAFC Proc. Rep. 

89-04, 201 p.

to within 10 m of the seafl oor or to targeted depths in 

deeper waters, following standard MARMAP procedures 

(Smith and Richardson, 1977). Other gear used included 

1-m2 Tucker trawls, sled trawls (modifi ed Tucker trawl 

towed on the seafl oor), Methot nets, Multiple Opening/

Closing Net and Environmental Sensing System (MOCNESS), surface-towed trawls (modifi ed Cantrawls), modifi ed beam trawls, rock dredges, Sameoto neuston nets, 

and dip nets (Tucker, 1951; Sameoto and Jaroszyinski, 

1969; Wiebe et al., 1976; Methot, 1986; Eisner et al., 

2012). Ichthyoplankton samples were preserved in a 5% 

buffered formalin-seawater solution and later sorted for 

eggs and larvae, which were removed and identifi ed to 

the lowest possible taxon at the Plankton Sorting and 

Identifi cation Center in Szczecin, Poland. Egg and larval 

identifi cations were verifi ed at AFSC. Detailed accounts 

of sampling and identifi cation protocols are available in 

Matarese et al. (2003). Other sources of material include

the University of Washington Fish Collection, the Vancouver Public Aquarium, Hokkaido University Museum 

of Zoology, Oregon State University, Canada Department of Fisheries and Oceans, and Chris Ehrler of Tenera 

Environmental, Inc., in San Luis Obispo, California.

Larvae were identifi ed with the serial method. This 

method uses adult characters to identify juveniles and 

progressively links them to smaller specimens through 

a continuous sequence of shared or similar features 

(Powles and Markle, 1984). Characters described by 

Mecklenburg et al. (2002), especially meristic counts, 

were used to progressively link adults and juveniles to larvae through the use of shared characters (Busby, 1998). 

Developmental series were illustrated with a camera lucida attached to a dissecting microscope. Only melanistic 

pigment was described because formalin fails to preserve 

other color pigments. The best representative specimen 

of each taxon at each stage of development (prefl exion, 

fl exion, and postfl exion) was illustrated. Transformation 

or juvenile stages were illustrated if available. Because 

pigmentation among specimens was variable, some pigment described in the taxon accounts may not be visible 

on the illustrations. Some larvae (e.g., High Cockscomb 

[Anoplarchus purpurescens]) used for illustrations were 

reared in captivity (e.g., at Vancouver Public Aquarium 

and NOAA wet labs). Comparisons between reared and 

wild-caught larvae must be done with caution because 

rearing (culture) may affect the morphology, behavior, 

and biochemistry of individuals (Blaxter, 1976; Hunter, 1984). Reared larvae also may be more heavily pigmented than are wild-caught specimens (Watson, 1982). 

Reared larvae are noted on fi gures. 

Overall larval pigmentation for most stichaeids is categorized as follows: 1) lightly pigmented, with pigment 

on the body limited to postanal ventral melanophores 

(PVM); 2) moderately pigmented, with pigment on the 

body consisting of PVM and dorsal or lateral pigment; 

or 3) heavily pigmented, with pigment distributed over 
4 Professional Paper NMFS 15 

Figure 1

Maps of study area showing geographic features described in species accounts for (A) North Pacific Ocean 

off Alaska, and (B) North Pacific Ocean off Canada and the west coast of the United States.
5

the entire body. Descriptions and illustrations of anatomical and morphometric features and other terms used 

herein to describe placement of pigment can be found in 

Matarese et al. (1989) and Moser (1996) with one exception: the area of the body between the anus and posterior edge of the hypural plates is defi ned as the postanal body instead of the tail. 

Measurements were taken on the left side of a fi sh, 

when possible, with a calibrated digital image-analysis 

system. The system consists of a video camera attached 

to a dissecting stereomicroscope or camera lens, a computer with a digital imaging board, software, and a video 

monitor (Blood and Matarese, 2010). Only larvae that 

were not badly damaged during collection or preservation were measured. In some cases, early transformation-stage individuals were included in morphometric 

measurements of postfl exion larvae. The transformation 

stage begins at the fi rst appearance of adult pigmentation and ends at the juvenile stage with the beginning of 

squamation (Kendall et al., 1984); early transformationstage larvae are most similar morphologically to postfl exion larvae. The following measurements were made 

on larvae and early juveniles:

Standard length (SL): Snout tip to notochord tip prior 

to development of caudal fi n (=notochord length), 

then to posterior margin of hypural element. All larval 

body lengths in this study are standard lengths unless 

otherwise noted.

Body depth: Vertical distance from dorsal to ventral body 

margin at the pectoral-fi n origin. Larvae are referred 

to as slender-bodied (body depth <10% SL), having 

a moderate body depth (10–12% SL), or deep-bodied 

(body depth >12% SL).

Snout to anus length: Distance along body midline from 

snout tip to a vertical line through center of anal 

opening.

Head length (HL): Snout tip to posterior edge of opercle.

Snout length: Snout tip to anterior margin of orbit. 

Eye diameter: Greatest distance of orbit.

Selected specimens were cleared and differentially 

stained to count meristic structures and follow sequence 

of fi n development (Potthoff, 1984). Meristic counts reported for each species were compiled from this study, 

the AFSC meristic database, and other sources (Tokuya 

and Amaoka, 1980; Watson, 1996; MacDonald, 2001; 

Hatooka, 2002; Mecklenburg et al., 2002; Hastings and 

Walker, 2003). Illustrated specimens were not cleared 

and stained, and, although their meristic counts fall 

within the overall ranges listed for each species (Table 

2), individual counts from illustrations may differ from 

those found in the species-specifi c meristic tables generated from clearing and staining in this study. In descriptions of development of meristic features, unossifi ed 

precursors of fi n and vertebral elements are classifi ed as 

present. The term developed is used to indicate that the 

adult complement of fi n elements, vertebral centra, or 

neural and haemal spines are completely formed but 

have not begun to ossify. Fins and vertebral elements are 

referred to as ossifi ed upon initial uptake of alizarin reds stain. The order of presentation of osteological descriptions is median fi ns, paired fi ns, and vertebrae, by sequence of ossifi cation within each. For some taxa, not all 

stages of development could be cleared and stained since 

the number of specimens was limited. Radiographs of 

adults were used to count meristic features of taxa that 

lack complete published data. 

A brief summary of the life history of each species 

is provided primarily on the basis of previously published literature. Some unpublished AFSC data were used 

to describe adult distributions (RACEBASE, an Oracle 

database developed by and available through the AFSC 

Resource Assessment and Conservation Engineering Division, comprises data from adult groundfi sh assessment, 

hydroacoustic, and foreign surveys conducted by federal 

fi shery scientists from 1948 to the present). Adult lengths 

are reported as SL (standard length) or TL (total length), 

depending on the source of information.

Larval distribution is described on the basis of data 

obtained from this study and from ICHBASE (if available), unless otherwise noted. Larval lengths and months 

when collected pertain to planktonic larvae (i.e., larvae 

collected with dip nets and bongo, Cantrawl, neuston, 

Methot, MOCNESS, and Tucker gear). Previously published information on occurrence of larvae is reported if 

available. 

Maps of larval distribution were produced from data 

collected for this study and display locations of planktonic larvae. Data collected by AFSC include information on abundance. For those taxa accompanied by 

abundance data, a geographic information system (GIS) 

based on ArcInfo software, vers. 8.1.2 (Esri, Inc., Redlands, CA) was used to produce a black-and-white choropleth map to show the general spatial extent of the 

data and the general trend of average larval abundance 

over space. Choropleth maps shade statistical units with 

intensity proportional to the data values. A fourth-root 

transformation was applied to the Slender Cockscomb 

(Anoplarchus insignis) data to normalize the abundance. 

The data for all other species were normally distributed. 

For species lacking abundance data, such as specimens 

obtained from the Hokkaido University Museum of Zoology, Vancouver Public Aquarium, Oregon State University, and Canada Department of Fisheries and Oceans, 

black-and-white maps were produced with point data. 

On these maps, symbols indicate presence only.

Nomenclature generally follows Makushok (1958) 

and common names follow Nelson et al. (2004). Nomenclature of the genus Lumpenus follows MacDonald 

(2001), who cited Makushok (1958) as stating that 

“Lumpenus, Leptoclinus, and Anisarchus form a natural 
6 Professional Paper NMFS 15 Table 2 Meristics of northeastern Pacifi c Ocean and Bering Sea Stichaeidae. Additional meristic information for Eumesogrammus, Chirolophis, Acantholumpenus, Alec- tridium, Anoplarchus, and Cebidichthys obtained from radiographs. Br=branchiostegal rays. Vertebrae Fins Taxon Precaudal Caudal Total Dorsal Anal Pectoral Pelvic Caudal Br STICHAEINAE Eumesogrammus praecisus1 15–17 35–37 50–53 XLVII–L I–II,29–32,II–III 16–19 I,3 3–5,6–7+6–7,4–5 6 Stichaeus punctatus 14–16 36–40 51–55 XLVI–XLIX I–II,32–37 15–16 I,4 5–6,6–7+6–7,3–5 6 CHIROLOPHINAE Bryozoichthys lysimus 16–17 53–58 69–75 LXI–LXVI I,47–53 14–15 I,3 6,6+7–8,3–5 6 Bryozoichthys marjorius 15–17 55–59 71–76 LXVII–LXXI I,51–59 15–16 I,3 5-6,6+7–8,3–4 6 Chirolophis decoratus2 16–17 50–53 65–69 LX–LXIV I–II,44–49 13–16 I,3–4 2–6,6+8–9,1–4 6 Chirolophis nugator3 14–16 42–46 55–61 LIII–LVI I–II,37–42 13–14 I,3–4 3–6,6+7–8,1–4 6 Chirolophis snyderi4 16–17 44–48 61–65 LVII–LXI I–II,43–47 13–16 I,4–5 5,6+7–8,2–4 6 C. tarsodes [holotype radiograph] 15 [15] 47–49 [50] 63–65 [65] LVIII–LX [LIX] I,43–46 [I,46] 14–15 [14?] I,3–4 [I,4?] [4,6+7,2] 6 Gymnoclinus cristulatus 18–19 46–47 65 LVIII–LXIII 40–43 14–16 I,2 5,6+7,2–3 6 LUMPENINAE Acantholumpenus mackayi4 27–30 47–50 76–80 LXVIII–LXXVI II,41–48 13–16 I,3 4,6+7,5 6 Lumpenella longirostris 22–25 47–49 70–75 LXI–LXXI II–V,36–42 12–14 I,2–3 4–5,6+6,4–6 6 Lumpenus fabricii 23–28 43–49 70–75 LX–LXVI I,38–44 13–16 I,3 5–6,6+7,4–6 6 Lumpenus maculatus 25–30 39–43 66–72 LVII–LXIV I–II,34–40 14–16 I,3–4 6-7,7+6–7,5-6 6 Lumpenus medius 21–25 42–50 65–70 LVIII–LXIV I,37–42 13–15 I,3 4–5,6–7+6–7,2–4 6 Lumpenus sagitta 26–29 46–55 75–82 LXIV–LXXII I–II,44–50 13–17 I,3 5–7,6+7–8,4–6 6 Poroclinus rothrocki 18–20 45–48 64–68 LVII–LXVII III,40–44 13–15 I,3 5–6,7+6,4 6 OPISTHOCENTRINAE Lumpenopsis hypochroma 17–18 36–38 54–55 XLIV–XLIX I,30–32 12–13 I,3 6,6+7,6 Opisthocentrus ocellatus 21–23 39–44 61–67 L–LXII I–II,32–39 18–21 absent 5 Plectobranchus evides 20–21 39–40 59–61 LIV–LVII II–III,34–36 14–15 I,3 4,7+6–7,2–4 5–6 XIPHISTERINAE Alectrias alectrolophus 17–20 47–50 64–70 LIX–LXVI I,41–45 09–11 absent 5 Alectridium aurantiacum5 20–21 47 60–68 LII–LXIV I,41–44 11 absent 3–4,6+7,2 5 Anoplarchus insignis6 17–19 42–49 60–69 LVII–LXIV I,38–46 09–10 absent 1–5,6–7+6–7,1–4 5 Anoplarchus purpurescens7 16–19 40–46 58–64 LIV–LX I–II,35–42 09–10 absent 1–4,6–7+6–7,1–4 5 Cebidichthys violaceus8 23–25 40–47 65–71 XXII–XXV, 40–43 I–II,39–42 10–11 absent 4–5,7+7,2–3 5 Phytichthys chirus 24–25 50–52 75–76 LXIX–LXXVIII II–III,40–50 14–15 absent 6+7 6 Xiphister atropurpureus 22–24 51–56 73–80 LXV–LXXII 0–I,49–55 11–12 absent 5–6,6+6–7,3–4 6 Xiphister mucosus 29–31 44–53 73–83 LXXI–LXXVIII 0–I,46–50 12 absent 5,6+6–7,3–4 6 1 Radiographs examined for 11 specimens. 5 Radiographs examined for 2 specimens. 2 Radiographs examined for 5 specimens. 6 Radiographs examined for 32 specimens. 3 Radiographs examined for 25 specimens. 7 Radiographs examined for 42 specimens. 4 Radiographs examined for 6 specimens. 8 Radiographs examined for 10 specimens.
7

group with Leptoclinus and Anisarchus representing 

a more derived condition.” With this taxonomic consideration, Leptoclinus and Anisarchus are considered 

synonyms (subgenera) of Lumpenus, leading to the validity of the species names Lumpenus maculatus and 

L. medius for the Daubed Shanny and Stout Eelblenny. 

The composition of the suborder Zoarcoidei follows 

Nelson (2006), including the families Bathymasteridae, Zoarcidae, Stichaeidae, Cryptacanthodidae, Pholidae, Anarhichadidae, Ptilichthyidae, Zaproridae, and 

Scytalinidae.

Overall comparisons 

Family

Stichaeidae is the most diverse zoarcoid family in terms 

of early life history stages in the NEP. Stichaeidae larvae can be distinguished from other zoarcoid larvae by 

a combination of characters, including myomere count, 

snout to anus length <50% SL, and light pigment patterns. Before fi n rays develop, small larvae are elongate 

and resemble other elongate zoarcoid forms, especially 

bathymasterids and pholids. Larvae in these three families share a relatively similar overall pigment pattern that 

includes PVM and melanophores positioned dorsally 

over the gut and hindgut. Meristic features, not always 

visible in early stage larvae, may serve to distinguish 

stichaeid larvae from other similar zoarcoid larvae. The 

typical I,5 pelvic-fi n count is present in bathymasterids, 

but it is reduced or absent in pholids and stichaeids and 

absent in all other zoarcoid families. Although zoarcoids 

typically hatch at relatively large sizes (e.g., up to 33 

mm), newly hatched stichaeid larvae are small, usually 

hatching at lengths of 6–11 mm (Matarese et al., 1989). 

Larvae do not undergo a dramatic transformation stage, 

but some species have an extended pelagic prejuvenile 

stage (e.g., Lumpenus spp.).

In general, bathymasterids have fewer myomeres (46–

55) than do the other two families, and pholids have a 

comparatively longer gut (snout to anus length >50%) 

and more myomeres (81–101) than do stichaeids (Matarese et al., 1989). Zoarcids lack a pelagic larval stage; after yolk absorption, their larvae resemble small juveniles 

(Kendall et al., 1983). Cryptacanthodids have considerably more pigment than stichaeids. Anarhichadids have 

highly distinctive larvae; Anarhichas larvae have a large 

eye and heavy pigmentation pattern, and Anarrhichthys

larvae are highly elongate with 221–251 myomeres. 

Among the remaining monotypic zoarcoid families (Ptilichthyidae, Scytalinidae, and Zaproridae), larval Ptilichthys are distinctive with an elongate body and 227–240 

myomeres, larval Zaprora have a distinctive heavy pigmentation pattern and large eye, and larval Scytalina are 

unknown (Matarese et al., 1989).

Subfamilies (Table 1)

According to Nelson (1984, 2006), Stoddard (1985), 

Yatsu (1986), Mecklenburg and Sheiko (2004), and others, the family Stichaeidae includes six subfamilies, of 

which fi ve are present in the NEP. Larval Stichaeinae, 

with two species in the NEP, usually have a short gut, 

PVM, hypaxial pigment, and a low myomere count. Larval Chirolophinae, with seven species in the NEP, possess 

more pigment than do other subfamilies except Stichaeinae, with pigment dorsally on the gut, PVM, caudal melanophores, and internal pigment along the notochord. 

Larval Lumpeninae, with seven species in the NEP, generally possess a more elongate body and gut, and they 

usually have pigment dorsally over the gut and PVM but 

lack internal pigment along the notochord. Larval Opisthocentrinae, with three species in the NEP, are individually distinctive and do not form a cohesive group on the 

basis of known larval characters. Larval Xiphisterinae, 

with eight species in the NEP, all lack pelvic fi ns and 

have pigment dorsally on the gut, a row of melanophores 

on the ventral surface of the gut, PVM, and internal pigment along the notochord. 

Subfamily Stichaeinae

Eumesogrammus praecisus, Fourline Snakeblenny

See “Other material” section of this guide.

Stichaeus punctatus, Arctic Shanny

(Figs. 2, 3; Tables 3, 4)

Literature. Faber (1976), Farwell et al. (1976), Eschmeyer et al. (1983), Haryu et al. (1985), Keats et al. 

(1993), Mecklenburg et al. (2002).

Material examined. 80 specimens (6.4–35.0 mm) from 

the Chukchi and Bering seas and Gulf of Alaska (see Appendix for details).

Distribution and life history. Stichaeus punctatus is 

circumarctic on subtidal (0–100 m) rocky to sandy bottom (Mecklenburg et al., 2002), occurring in the seas of 

Okhotsk and Japan, in the Beaufort Sea southward to 

British Columbia in the North Pacifi c Ocean, and in the 

Arctic Ocean south to the Gulf of Maine in the western North Atlantic Ocean. Maximum length is 22 cm TL 

(Eschmeyer et al., 1983) and longevity is 16 years (Keats 

et al., 1993). Information on spawning is restricted to 

the North Atlantic Ocean, where the spawning season is 

February and March, egg diameter is 1.7 mm, and absolute fecundity is about 1600–2500 eggs (Farwell et al., 

1976). Larvae are pelagic. Transformation to the juvenile 

stage occurs at about 25 mm (Farwell et al., 1976).
8 Professional Paper NMFS 15 

Figure 2

Distribution and abundance of larvae of Arctic Shanny (Stichaeus punctatus) collected by bongo net in the Chukchi and 

Bering seas and Gulf of Alaska. Isobath=1000 m.

Larval distribution (Fig. 2). Larvae of S. punctatus are 

found from the western Chukchi Sea, the Bering Sea near 

St. Matthew and the Pribilof Islands, Unimak Pass, nearshore areas along the Alaska Peninsula and Shumagin 

Islands, in Shelikof Strait and around Kodiak Island, to 

nearshore areas along the Kenai Peninsula. Larvae at 

lengths of 6.4–27.0 mm are collected in April–August, 

with highest abundances in May.

Morphology (Table 3). Notochord fl exion in S. punctatus begins at 9.0 mm and is complete by 16.4 mm. Postfl exion larvae are 16.4–28.2 mm. Farwell et al. (1976) 

presented an illustration of a 25.0 mm specimen that 

appears to be in early transformation stage. In our material, transformation was not evident until after 28.2 

mm; the juvenile stage began before 35.0 mm. Larvae 

are initially slender with a mean body depth of 9.0% 

SL in the prefl exion stage, becoming moderately deep 

(10.7% SL) in the fl exion stage and deep-bodied (12.3% 

SL) in the postfl exion stage. Relative head and snout 

lengths increase through development, while eye diameter decreases. Snout to anus length increases slightly with 
9

Figure 3

Larvae of Arctic Shanny (Stichaeus punctatus): (A) preflexion stage, 8.3 mm standard length (SL), University of Washington 

(UW) 144830; (B) flexion stage, 12.4 mm SL, UW 144858; and (C) postflexion stage, 16.8 mm SL, UW 144862. Illustrations by R. Cartwright.

development from 37.4% SL in the prefl exion stage to 

40.2% SL in the postfl exion stage.

Pigmentation (Fig. 3). Larvae of S. punctatus are 

moderately pigmented (dorsal and lateral pigment in 

addition to PVM). Although the earliest prefl exionstage larvae lack head pigment, it is present on the 

head throughout most of larval development. Nape pigment is present in prefl exion and later stages. Prefl exion larvae have several melanophores anteriorly along 

the dorsal surface of the gut, and a continuous line 

of pigment along the anterior ventral midline of the 

gut persists throughout development. Melanophores 

are present along the hindgut nearly to the anus. A 

row of PVM, spaced at about one melanophore per 

myomere, begins about fi ve myomeres posterior to the 

anus. Several melanophores are on the dorsal midline 

of the caudal peduncle, and internal pigment is present 

along the dorsal surface of the notochord. Postanal pigment along the hypaxial myomeres, characteristic for 

larvae of S. punctatus, is present in the earliest prefl exion larvae. 
10 Professional Paper NMFS 15 

Flexion-stage larvae have pigment dorsally on the 

head and nape. Melanophores extend along the entire 

dorsal surface of the gut, and pigment on the postanal 

body increases signifi cantly. Several melanophores appear along each side of the dorsal midline, continue 

to increase anteriorly and posteriorly, and appear as a 

double row when viewed dorsally. The series of PVM 

begins about 4–6 myomeres posterior to the anus and 

hypaxial pigment increases to appear as oblique pigment 

lines. The internal pigment along the dorsal surface of 

the notochord is visible only in the area of the caudal peduncle. A small patch of pigment develops at the base of 

the caudal-fi n rays along the margin of the lower hypural 

plate in late fl exion.

Head pigment increases in postfl exion larvae; melanophores are added dorsally on the head and posterior to the eye in the opercular region. Nape pigment 

was visible during early postfl exion in some specimens 

but was less visible in later larvae as the melanophores 

became embedded. The dorsal pigment on the gut becomes internal. Several melanophores are visible on the 

hindgut. The double row of dorsal pigment extends anteriorly onto the preanal myomeres. PVM (>30) begin 

just posterior to the anus. In the caudal peduncle area, 

internal pigment is present above and below the notochord. Pigment at the base of the caudal-fi n rays is present along the margins of the upper and lower hypural 

plates.

Table 3

Body proportions of Arctic Shanny (Stichaeus punctatus) larvae. Except for standard length (SL), values 

given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± standard 

deviation, and range (in parentheses). 

 Prefl exion Flexion Postfl exion

Sample size 16 28 30

Standard length (mm) 7.8±0.8 (6.4–9.0) 11.1±1.7 (9.0–15.5) 21.5±3.3 (16.4–28.2)

Head length/SL 12.8±1.3 (10.8–15.2) 13.7±1.5 (9.6–16.5) 18.6±2.1 (14.2–23.2)1

Snout length/HL 16.6±6.2 (7.9–27.0)2 23.6±5.2 (14.7–32.1) 26.3±6.1 (15.8–33.6)3

Eye diameter/HL 48.1±4.7 (41.3–57.4) 43.1±6.2 (30.7–59.2) 29.0±3.9 (20.9–34.9)3

Snout to anus length/SL 37.4±2.3 (34.4–43.6)2 38.8±2.0 (32.9–43.1) 40.2±3.1 (31.4–44.9)

Body depth/SL 9.0±1.0 (7.9–11.2) 10.7±0.9 (9.1–12.0) 12.3±1.6 (8.9–15.5)2

1 n=29 2 n=15 3 n=28

Table 4

Meristic counts of cleared and stained Arctic Shanny (Stichaeus punctatus) larvae and juveniles. Counts are of ossifi ed elements only. Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 6.2

 7.5

 9.1

 11.2

 13.1 15 32 47

 14.8 4 15 301 45 31 15 38 53 7+7,1

 16.2 6 14 371 51 37 14 38 52 3,7+7,1

 18.5 XLVI II,34 15 I,2 6 14 381 52 38 14 39 53 5,7+7,4

 20.5 XLVII II,34 15 I,4 6 14 391 53 38 14 40 54 6,6+6,5

 24.5 XLVIII II,35 15 I,4 6 14 391 53 391 14 39 53 5,7+6,5

 28.0 XLIX II,35 15 I,4 6 15 381 53 38 15 39 54 6,7+6,5

 35.0 XLVII II,33 15 I,4 6 15 371 52 37 15 38 53 5,7+7,3

1 Two neural and haemal spines on second preural centra.

Neural spines Centra

Spines, rays
11

Meristic features (Table 4). Principal caudal-fi n rays are 

present by 9.1 mm, developed by 13.1 mm, and ossifi ed 

by 14.8 mm. Procurrent caudal-fi n rays are present by 

14.8 mm, and ossifi cation begins before all elements are 

developed. All procurrent caudal-fi n rays are ossifi ed by 

18.5 mm. Anal-fi n elements are present by 14.8 mm and 

dorsal-fi n spines by 16.2 mm. Ossifi cation begins before 

all elements in both fi ns are developed, with the dorsal 

spines ossifi ed by 18.5 mm and the anal-fi n elements ossifi ed by 20.5 mm. Pectoral-fi n rays are present by 11.2 

mm, developed by 16.2 mm, and ossifi ed by 18.5 mm. A 

pelvic-fi n bud is present at 16.2 mm. Pelvic-fi n elements 

are present and begin to ossify by 18.5 mm; development 

and ossifi cation are complete by 20.5 mm. Vertebral centra are present by 6.2 mm; development and ossifi cation 

are complete by 14.8 mm. Neural and haemal spines are 

present and partially ossifi ed by 14.8 mm and are developed and ossifi ed by 16.2 mm. Ranges of adult meristic 

counts are given in Table 2.

Species comparisons. Larvae of S. punctatus are unique 

among stichaeids because they have slash-like postanal 

pigment along the hypaxial myomeres and the lowest 

total vertebral counts (51–55) in the family, except for 

Eumesogrammus praecisus (50–53). In addition, S. punctatus is deeper bodied (mean 9.0–12.3% SL) compared 

to most other larval stichaeids, particularly during the 

fl exion and postfl exion stages. 

Larval S. punctatus are most likely to be confused 

with the bathymasterid Northern Ronquil (Ronquilus 

jordani). However, S. punctatus can be distinguished in 

the prefl exion stage by the postanal hypaxial pigment, 

which is absent in R. jordani. Ronquilus jordani also 

has a row of melanophores along the dorsal midline and 

some pigment spots on the ventral caudal fi nfold that 

are absent in prefl exion-stage S. punctatus. In later-stage 

larvae, slash-like postanal pigment along the hypaxial 

myomeres forms in R. jordani but is restricted to the 

posterior one-fourth of the body. Ronquilus jordani is 

deeper bodied throughout development, has a shorter 

snout to anus length, has fewer vertebrae (49–50 versus 

51–55), and completes its fl exion and postfl exion stages 

at smaller sizes than does S. punctatus.

Subfamily Chirolophinae

Bryozoichthys lysimus, Nutcracker Prickleback 

(Figs. 4–6; Tables 5, 6)

Literature. Haryu et al. (1985), Shiogaki and Haryu 

(1988), Mecklenburg et al. (2002), Tokranov and Orlov 

(2004).

Material examined. 82 specimens (6.9–49.0 mm) from 

the Bering Sea and Gulf of Alaska (see Appendix for 

details).

Distribution and life history. Bryozoichthys lysimus 

has been collected in the eastern Bering Sea northwest 

of St. Matthew Island, along the entire Aleutian Island 

Archipelago (RACEBASE), and northern Gulf of Alaska. 

It is also found in the western Bering Sea, Sea of Okhotsk, Kuril Islands, and northern Sea of Japan (Mecklenburg et al., 2002). Collected at depths of 45–600 m 

(most commonly at 100–200 m), Nutcracker Prickleback 

reach a maximum length of 270 mm TL (Tokranov and 

Orlov, 2004). Little is known of their life history. Eggs 

are probably demersal and spawned in an adhesive mass; 

larvae are pelagic. 

Larval distribution (Figs. 4, 5). Larvae of B. lysimus 

(7.4–33.5 mm, mean 19.8 mm) are collected in February–July in bongo tows, with highest abundance in 

April, over the Bering Sea shelf and slope, west of and 

around Unimak Pass, in the northern Gulf of Alaska 

on the shelf south of the Alaska Peninsula, in Shelikof 

Strait, and north and south of Kodiak Island (Fig. 4). The 

spatial distribution of larvae collected in neuston tows 

is similar, but neustonic larvae are more abundant and 

those larvae in the Bering Sea are found farther west in 

the Aleutian Islands and into Bristol Bay (Fig. 5). Neustonic larvae are larger, on average, than are larvae collected in bongo tows (14.1–35.6 mm, mean 24.3 mm) 

and occur in March–July, with greatest numbers collected in May. 

Morphology (Table 5). Notochord fl exion in B. lysimus 

begins before 12.0 mm and is complete by about 22.0 

mm. Postfl exion larvae are 21.6–35.6 mm. Transformation begins after 36.0 mm and the juvenile stage begins 

before 49.0 mm. Larvae are slender-bodied with a mean 

body depth of 6.4% SL in the prefl exion stage (n=2), 

increasing to 8.4% SL in the postfl exion stage. Relative 

head and snout lengths increase throughout larval development, while relative eye diameter decreases. Relative 

snout to anus length increases from the prefl exion stage 

to the fl exion stage, then decreases during the postfl exion 

stage. 

Pigmentation (Fig. 6). Larvae of B. lysimus are moderately pigmented. Head pigment in the prefl exion stage 

is restricted to several melanophores on the dorsal surface over the midbrain. The gut is pigmented dorsally 

and ventrally. A double row of internal melanophores, 

spaced one melanophore per myomere, extends along the 

dorsal surface of the gut. About four closely spaced melanophores occur along the dorsal surface of the hindgut. About four melanophores are positioned ventrally 

at midgut and may appear as a solid line in some individuals. A double row of melanophores on either side of 

the dorsal midline begins just anterior to midbody and 

extends to the caudal peduncle. Pigment is irregularly 

spaced anteriorly and becomes more regular posteriorly, 
12 Professional Paper NMFS 15 

Figure 4

Distribution and abundance of larvae of Nutcracker Prickleback (Bryozoichthys lysimus) collected by bongo net in the Bering Sea and Gulf of Alaska. Isobath=1000 m.

about one melanophore per myomere. Internal pigment 

(one melanophore per myomere) dorsally along the notochord begins as a single row over the hindbrain and 

extends to the caudal area. A series of about 46–48 PVM 

begins about 1–3 myomeres posterior to the anus and 

consists of a melanophore on each myo septum along 

the entire length of the postanal body to the caudal 

peduncle. The number, spacing, and anterior and posterior extent of the PVM are similar at all stages of development. Caudal pigment is restricted to small melanophores around the tip of the notochord and two short 

streaks of pigment extending at oblique angles dorsad 

and ventrad from the notochord tip.

In the early fl exion stage, larvae develop 2–4 additional melanophores over the midbrain and occasionally develop light pigment over the hindbrain. Internal 

pigment is visible in the otic area by 17 mm and on the 

midbrain by 21 mm. Light pigment on the jaw angle 

is visible by 21 mm. Pigment on the isthmus begins as 

one pair of melanophores at 17 mm and increases to 

2–3 pairs by 20 mm. Pigment on the ventral surface of 

the gut expands during early fl exion to extend along 

the anterior one-third to one-half of the gut in a continuous line, and 1–3 melanophores are present on the 

lateral surface of the gut behind the pectoral fi n. Internal pigment along the notochord became increasingly 
13

Figure 5

Distribution and abundance of larvae of Nutcracker Prickleback (Bryozoichthys lysimus) collected by neuston gear in the 

Bering Sea and Gulf of Alaska. Isobath=1000 m.

diffi cult to see during fl exion and was visible only at 

the nape and caudal peduncle in some specimens. Additional PVM that are more closely spaced than the 

rest of the series are visible in the area of the caudal 

peduncle. A row of closely spaced small melanophores 

is present on the lateral surface of the caudal peduncle 

and upper hypural plate near the notochord in latefl exion larvae. Light caudal-fi n pigment was visible in 

some specimens by 17 mm, but it commonly is not apparent until about 22 mm and is restricted in fl exion 

larvae to a streak that extends from the tip of the notochord toward the dorsoposterior margin of the caudal fi n. 

Pigment changes in postfl exion larvae occur mainly on the head and tail. Additional pigment is present over the midbrain, on the opercle, and, in some 

specimens, was seen on the snout. Internal pigment on 

the midbrain increases by 25 mm. In addition to the 

streak that extends from the tip of the notochord, pigment on the caudal fi n is concentrated into a narrow 

vertical band at the midpoint of the fi n rays at about 

26 mm; some specimens had an additional band twothirds of the distance to the posterior margin of the 

caudal fi n. More pigment appears on the caudal fi n by 

about 28 mm, but it is more concentrated on the distal half.
14 Professional Paper NMFS 15 

Figure 6

Larvae of Nutcracker Prickleback (Bryozoichthys lysimus): (A) preflexion stage, 10.5 mm standard length (SL), University 

of Washington (UW) 33190; (B) flexion stage, 16.5 mm SL, UW 144927, previously published as Bryozoichthys-Chirolophis

(Matarese et al., 1989); and (C) postflexion stage, 29.0 mm SL, UW 144936, previously published as Bryozoichthys-Chirolophis (Matarese et al., 1989). Illustrations by B. Vinter.

Table 5

Body proportions of Nutcracker Prickleback (Bryozoichthys lysimus) larvae. Except for standard length (SL), 

values given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± standard deviation, and range (in parentheses). 

 Prefl exion Flexion Postfl exion

Sample size 2 33 36

Standard length (mm) 8.5±2.3 (6.9–10.2) 17.6±3.4 (12.0–22.2) 27.5±3.9 (21.6–35.6)

Head length/SL 11.3±3.1 (9.1–13.5) 13.1±1.9 (9.5–19.0) 14.2±1.6 (11.6–18.1)

Snout length/HL 9.71 18.6±3.7 (8.3–26.0) 22.6±3.6 (13.9–29.9)

Eye diameter/HL 44.1±11.7 (35.8–52.4) 33.1±5.5 (24.2–46.1)2 25.0±3.2 (18.2–32.7)

Snout to anus length/SL 23.2±17.5 (10.8–35.6) 35.2±1.9 (31.4–40.2) 33.9±1.1 (31.2–36.5)

Body depth/SL 6.4±2.2 (4.8–8.0) 5.7±0.8 (4.3–8.1) 8.4±1.5 (5.3–11.9)

1 n=1 (6.9-mm specimen) 2 n=32
15

Meristic features (Table 6). Principal caudal-fi n rays 

are present at 14.5 mm, are developed and begin to ossify by 19.0 mm, and complete ossifi cation by 21.5 mm. 

Procurrent caudal-fi n rays are present at 19.0 mm, developed by 25.0 mm, and ossifi ed by 32.0 mm. Dorsalfi n and anal-fi n elements are present at 21.5 mm, developed by 25.0 mm, and ossifi ed by 32.0 mm; the count 

of I,53 anal-fi n elements, observed in the 32.0-mm specimen, is greater than the count previously reported for 

this species (I,47–50; Mecklenburg et al., 2002). Pectoral-fi n rays are present at 17.0 mm, developed by 21.5 

mm, and ossifi ed by 25.0 mm. Pelvic-fi n elements are 

developed by 32.0 mm and ossifi ed by 49.0 mm. Vertebral centra are present at 12.0 mm and are developed 

by 21.5 mm. Ossifi cation of the vertebral centra begins 

at 19.0 mm, before development is complete, and progresses anteriorly and posteriorly from midbody. Ossifi cation of all vertebral centra is complete by 21.5 mm; 

the count of 58 caudal centra, as recorded for the 32.0-

mm specimen, is greater than the count previously reported for this species (53–54; Matarese et al., 1989). 

All neural and haemal spines are ossifi ed by 25.0 mm. 

Ranges of adult meristic counts are given in Table 2.

Species comparisons. Larvae of B. lysimus resemble 

species of Chirolophis, particularly during the prefl exion 

and fl exion stages, but they can be distinguished by the 

continuous row of evenly spaced internal melanophores 

dorsally on the notochord from the nape to the hypural 

region, in contrast to species of Chirolophis in which 

this pigmentation is absent on the preanal body. Melanophores also are present around the notochord tip and on 

the caudal fi n and fi nfold in B. lysimus at all stages of development, but they are absent in species of Chirolophis. 

In addition, B. lysimus has more vertebrae (myomeres; 

69–75) than all Chirolophis species except the Decorated 

Warbonnet (C. decoratus) (65–69).

Larval B. lysimus are most likely to be confused with 

its congener, the Pearly Prickleback (B. marjorius), but it 

can be distinguished by the internal notochord series that 

begins near the nape as a single rather than double row 

of melanophores. In addition, ventral melanophores on 

the gut are restricted to the anterior half rather than the 

entire length as in B. marjorius. Bryozoichthys lysimus

larvae usually have fewer caudal vertebrae (myomeres; 

53–58) than do B. marjorius larvae (55–59). 

Bryozoichthys marjorius, Pearly Prickleback 

(Figs. 7–9; Tables 7, 8)

Literature. Haryu et al. (1985), Mecklenburg et al. 

(2002).

Material examined. 90 specimens (7.7–53.0 mm) from 

the Bering Sea and Gulf of Alaska (see Appendix for 

details).

Distribution and life history. Bryozoichthys marjorius 

has been collected in the Bering Sea (RACEBASE), along 

the entire Aleutian Island Archipelago, in the northern 

Gulf of Alaska south of the Kenai Peninsula, and from 

Southeast Alaska to southern British Columbia, Canada. 

Collected at depths of 80–450 m (most commonly at 

100–250 m) (RACEBASE), Pearly Prickleback reach a 

maximum length of 305 mm TL (Mecklenburg et al., 

2002). Little is known of their life history. Eggs are probably demersal and spawned in an adhesive mass; larvae 

are pelagic. 

Larval distribution (Figs. 7, 8). Larvae of B. marjorius

(8.8–35.0 mm, mean 21.1 mm) are collected in bongo 

Table 6

Meristic counts of cleared and stained Nutcracker Prickleback (Bryozoichthys lysimus) larvae and juveniles. Counts are of 

ossifi ed elements only. Specimens between dashed lines (-----) were undergoing notochord fl exion. 

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 8.0 

 10.4 

 12.0 

 14.5 

 17.0 

 19.0 13 50 63 5+7

 21.5 16 53 69 6+7

 25.0 14 6 16 53 69 53 16 54 70 6+6

 32.0 LXIV I,53 15 6 16 57 73 57 16 58 74 6,6+7,5

 49.0 LXIV I,50 14 I,3 6 16 52 68 52 16 53 69 6,6+8,3

Neural spines Centra

Spines, rays
16 Professional Paper NMFS 15 

tows over the slope in the southern Bering Sea, Unimak 

Pass, Shelikof Strait in the northern Gulf of Alaska, and 

over the slope south of Kodiak Island (Fig. 7) in February–June, with most larvae collected in April–May. 

Larvae collected in neuston tows are larger, more widely 

distributed, and more abundant than larvae collected 

in bongo tows. Neustonic larvae (15.0–38.5 mm, mean 

26.0 mm) are found as far north in the Bering Sea as 59° 

N, typically over the slope and outer shelf, farther west 

in the Bering Sea north of the central Aleutian Islands, 

and farther east near Kayak Island in Southeast Alaska 

in March–July (Fig. 8), with the greatest numbers collected in April–May. 

Morphology (Table 7). Notochord flexion in B. marjorius begins after 11.0 mm and is complete by 24.0 

mm. Postflexion larvae are 21.4–40.0 mm. Transformation begins after 40.0 mm, and the juvenile 

stage begins before 53.0 mm. Larvae are slenderbodied with a mean body depth of 3.5% SL during 

the preflexion stage (n=2), increasing to 6.3% SL 

in the postflexion stage. Relative head length decreases from the preflexion stage to the flexion stage, 

then increases during the postflexion stage. Relative 

snout length increases during larval development, 

but relative eye diameter and snout to anus length 

decrease.

Figure 7

Distribution and abundance of larvae of Pearly Prickleback (Bryozoichthys marjorius) collected by bongo net in the Bering 

Sea and Gulf of Alaska. Isobath=1000 m.
17

Pigmentation (Fig. 9). Larvae of B. marjorius are moderately pigmented. Head pigment is absent in prefl exion 

larvae. There is one melanophore ventral to the base of 

the pectoral fi n on each side of the body. A single row of 

closely spaced melanophores extends along the ventral 

midline of the gut, becoming more widely spaced from 

about midgut to the anus. A double row of internal melanophores, spaced one per myomere, extends along the 

dorsal surface of the gut. There are 2–3 melanophores on 

the lateral surface of the gut near the pectoral fi n and an 

equal number on the dorsal surface of the hindgut. Melanophores form a double row on either side of the dorsal 

midline, starting just anterior to midbody, and are evenly 

spaced on every other myomere until about the last 15 

myomeres, where they become closer together.

Internally, there are two melanophores on each side 

of the nape that converge to a single row of internal 

melanophores along the dorsal midline beginning at the 

fi rst myomere. This series, consisting of one melanophore 

per myomere, extends the full length of the body internally along the dorsal midline to the area of the caudal peduncle. A row of PVM (about 46 melanophores) 

originates about four myomeres posterior to the anus 

and extends to the last myomere. A single melanophore 

is associated with each myomere until, at about 75% 

SL, spacing increases to one melanophore every 2–3 

Figure 8

Distribution and abundance of larvae of Pearly Prickleback (Bryozoichthys marjorius) collected by neuston gear in the Bering Sea and Gulf of Alaska. Isobath=1000 m.
18 Professional Paper NMFS 15 

myomeres. Posteriorly, the series ends in the area of the 

caudal peduncle with four closely spaced melanophores. 

In the caudal area, pigment extends along the dorsal 

margin of the notochord, around its tip, and along the 

posterior half of the ventral margin of the notochord. 

Light pigment is scattered on the dorsal and ventral caudal fi nfold.

Bryozoichthys marjorius develops head pigment during the early fl exion stage. Two melanophores appear 

over the hindbrain by about 14 mm. By the midfl exion 

stage (18 mm), these melanophores are embedded and 

additional external pigment appears over the midbrain. 

Late fl exion-stage larvae develop light pigment on the 

jaw angle and internal pigment in the otic area. Along 

the anterior margin of the gut, 2–3 pairs of melanophores appear by about 15 mm and form a curved line 

with pigment near the base of the pectoral fi n on the lateral surface of the gut. By the late fl exion stage (21 mm), 

2–4 melanophores are visible on the isthmus. Melanophores in the double row along the dorsal midline are on 

almost every myomere by about 14 mm; those melanophores in the posterior one-fourth of the series are closer 

Figure 9

Larvae of Pearly Prickleback (Bryozoichthys marjorius): (A) preflexion stage, 11.4 mm standard length (SL), University of 

Washington (UW) 144942; (B) early flexion stage, 14.0 mm SL, UW 60255; (C) late flexion stage, 20.5 mm SL, UW 82725; 

and (D) postflexion stage, 30.5 mm SL, UW 36366. Illustrations by B. Vinter.
19

together. Internal pigment along the notochord was diffi cult to see in some specimens except near the nape and 

on the posterior one-fourth of the body. PVM increase 

to 2–3 per myomere for the anterior three-fourths of the 

length of the series in larvae in the early fl exion stage. 

Posteriorly, the spacing becomes irregular and then resumes a spacing of 2–3 per myomere in the area of the 

caudal peduncle; the number of PVM is about 78. Pigment becomes embedded along most of the length of the 

series at about 16 mm and appears as a continuous row 

in the area of the caudal peduncle by about 19 mm. By 

the late fl exion stage (22 mm) about 53 PVM are spaced 

at one melanophore per myomere for most of the series, 

with pigment aligned along developing anal-fi n rays. 

Caudal pigment around the notochord and on the fi nfold remains unchanged in fl exion-stage larvae; additional pigment appears in the hypural area by 16 mm and 

above the notochord in the area of the caudal peduncle 

by about 22 mm.

Changes in head pigment in postfl exion larvae included the addition of a few melanophores on the ventroposterior edge of the opercle on some specimens by 

about 28 mm and of light pigment on the snout by about 

38 mm. The tip of the lower jaw is lightly pigmented 

by 31 mm. Pigment on the jaw angle was not visible in 

some specimens. Pigment along the ventral midline of 

Table 7

Body proportions of Pearly Prickleback (Bryozoichthys marjorius) larvae. Except for standard length (SL), 

values given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± 

standard deviation, and range (in parentheses). 

 Prefl exion Flexion Postfl exion

Sample size 2 39 44

Standard length (mm) 11.2±0.3 (10.9–11.4) 18.5±2.6 (13.8–23.6) 29.9±5.0 (21.4–40.0)

Head length/SL 11.9±1.2 (11.1–12.8) 11.2±1.4 (7.8–14.1) 13.0±1.6 (9.4–16.7)

Snout length/HL 17.6±6.6 (12.9–22.3) 19.9±7.2 (11.4–42.2)1 23.2±4.7 (15.6–37.5)

Eye diameter/HL 42.4±2.1 (40.9–43.9) 36.7±6.4 (26.8–51.7)1 26.5±4.6 (19.6–37.4)

Snout to anus length/SL 34.2±0.3 (34.0–34.4) 32.9±1.4 (30.3–35.6) 32.0±1.4 (29.3–36.7)

Body depth/SL 3.5±1.2 (2.7–4.4) 4.8±0.7 (3.6–6.6) 6.3±1.1 (4.1–9.2)

1 n=38

Table 8

Meristic counts of cleared and stained Pearly Prickleback (Bryozoichthys marjorius) larvae and a juvenile. Counts are of 

ossifi ed elements only. Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

7.7 

 11.2 

 13.7 

 14.8 

 16.1 16 56 72 56 16 57 73

 22.5 16 56 74 56 16 57 73

 25.0 XXX 5 16 58 74 58 16 58 74

 28.0 L I,35 6 16 55 71 55 16 56 72 4,6+8,3

 31.5 LXVI I,52 15 6 17 54 71 54 17 55 72 6,6+8,4

 33.0 LXIX I,54 15 I,3 6 16 57 73 57 16 58 74 5,6+8,3

 38.5 LXX I,54 * * 6 16 58 74 58 16 59 75 6,6+7,3

 39.5 LXIX I,55 15 I,3 6 16 58 74 58 16 59 75 6,6+8,4

 53.0 LXXI I,56 15 I,3 6 17 58 75 58 17 59 76 6,6+8,4

* Character damaged or missing.

Neural spines Centra

Spines, rays
20 Professional Paper NMFS 15 

the gut appears as a continuous line for three-fourths 

of its length by 22 mm. Melanophores along the margin 

of the isthmus increase to 3–4 pairs by 23 mm and to 5 

pairs by 31 mm. Caudal-fi n pigment is light in larvae in 

the early postfl exion stage (about 22 mm), except for a 

streak of darker pigment that extends from the tip of the 

notochord onto the caudal-fi n membrane between the 

fi n rays; this pigment was present in all specimens. More 

pigment develops on the rest of the caudal fi n membrane 

by about 28 mm; by about 34 mm, the posterior onethird of the caudal fi n is more heavily pigmented than 

the remainder. Ventral notochord pigment persisted in all 

specimens.

Meristic features (Table 8). Principal caudal-fi n rays are 

present before 16.1 mm and are developed and ossifi ed 

by 28.0 mm. Dorsal-fi n spines are present at 22.5 mm 

and ossifi cation appears to progress nearly synchronously 

with development, beginning at 25.0 mm and ending by 

33.0 mm. Anal-fi n elements are present at 25.0 mm. Ossifi cation begins at 28.0 mm before all elements are developed; all elements are developed and ossifi ed at 31.5 mm. 

Procurrent caudal-fi n rays are present by 22.5 mm and 

are ossifi ed at 28.0 mm before all are developed; all are 

developed and ossifi ed by 31.5 mm. Pectoral-fi n rays are 

present by 16.1 mm, developed by 25.0 mm, and ossifi ed 

by 31.5 mm. Pelvic-fi n elements are present at 31.5 mm 

and developed and ossifi ed by 33.0 mm. Vertebral centra 

are present at 14.8 mm, and all are developed at 16.1 

mm. Ossifi cation of the vertebral centra progresses from 

anterior to posterior and occurs by 16.1 mm. The count 

of 55 caudal centra, observed in the 31.5-mm specimen, 

is lower than the count previously reported for this species (56–59; Matarese et al., 1989), and the total count of 

76 vertebral centra observed in the 53.0-mm specimen is 

greater than the total count previously reported for this 

species (71–73; Mecklenberg et al., 2002). All neural and 

haemal spines are ossifi ed at 16.1 mm. Ranges of adult 

meristic counts are given in Table 2.

Species comparisons. See Bryozoichthys lysimus species-comparison section to distinguish larvae from Chirolophis species: similar pigment differences apply to 

B. marjorius. Also similarly, B. marjorius has more vertebrae (myomeres; 71–76) than all Chirolophis species 

(57–69). 

Larval B. marjorius are most likely to be confused 

with B. lysimus. However, B. marjorius can be distinguished by the internal series of dorsal melanophores 

that begins near the nape as a double row rather than a 

single row. In addition, pigment on the ventral surface of 

the gut extends along nearly its entire length rather than 

just the anterior half, and B. marjorius has more caudal 

pigment at similar stages of development. Larvae of B. 

marjorius usually have more caudal myomeres (55–59) 

than do B. lysimus larvae (53–58). 

Chirolophis decoratus, Decorated Warbonnet 

(Figs. 10–12; Tables 9, 10)

Literature. Eschmeyer et al. (1983), Shiogaki (1983), 

Mecklenburg et al. (2002).

Material examined. 70 specimens (8.7–42.5 mm) from 

the Bering Sea, Gulf of Alaska, and the coasts of British 

Columbia, Canada, and Washington (see Appendix for 

details).

Distribution and life history. Chirolophis decoratus occurs among rocky bottoms and reef crevices from subtidal areas to depths of 300 m (Eschmeyer et al., 1983) 

in the eastern Bering Sea shelf region as far north as 

Norton Sound, along the Aleutian Islands east of Adak 

Island, throughout the Gulf of Alaska shelf, and south 

to northern California. Adults reach a maximum length 

of 420 mm TL (Mecklenburg et al., 2002). Little is 

known of their life history, but a congener, Chirolophis 

japonicus, spawns in Mutsu Bay, Japan, in late November–December. In the aquarium, C. japonicus females 

guarded egg masses laid in a tube-like hollow of a concrete block. Eggs were 2.5 mm in diameter, had a milky 

white chorion and light yellow oil globule, and were adhesive to each other but not to the substrate. Time to 

hatch was about 2 months at 3.5–10.0°C; newly hatched 

larvae were 12.2–13.5 mm TL (Shiogaki, 1983).

Larval distribution (Figs. 10, 11). Larvae of C. decoratus (10.0–32.5 mm, mean 18.4 mm) are collected in 

bongo tows in January–May, with the greatest numbers 

collected in April and May, over the shelf in the Gulf of 

Alaska from south of Unimak Island to Shelikof Strait 

and east of Kodiak Island, southward off the central Oregon coast to the Oregon–California border (Fig. 10). 

Larvae collected in neuston tows are more widely distributed in the northern part of their range: on the southeastern Bering Sea shelf and over the shelf in the Gulf of 

Alaska to just west of Prince William Sound (Fig. 11). 

Neustonic larvae (12.0–34.0 mm, mean 25.1 mm) are 

larger than larvae collected in bongo tows and are found 

in February–June, with the greatest abundances observed 

in May. 

Morphology (Tables 9, 10). Notochord fl exion in C. 

decoratus begins after 12.5 mm and is completed by 19.5 

mm. Postfl exion larvae are 19.5–31.6 mm. Transformation occurs after 32.0 mm; length at the beginning of 

the juvenile stage is unknown. Larvae are slender-bodied 

with a mean body depth of 5.4% SL in the prefl exion 

stage, increasing to 8.9% SL in the postfl exion stage. 

Relative head and snout lengths increase, and eye diameter and snout to anus length decrease during larval development (Table 9). 
21

Pigmentation (Fig. 12). Larvae of C. decoratus are 

moderately pigmented. Head pigment is light and variable in the prefl exion stage: the nape region has several 

spots early in this stage, and some larvae have one or 

two melanophores at the posterior margin of the hindbrain. The gut is pigmented internally along its dorsal 

surface with 10 or more closely spaced melanophores. 

Several melanophores occur dorsally on the hindgut. 

Ventral pigment on the gut, when present, is restricted 

to a few melanophores anterior to midgut. A PVM series (>40) is present with a melanophore each at most 

myosepta along the entire length of the postanal body. A 

series of internal melanophores is present dorsally above 

the notochord, and a short series of external dorsal midline melanophores (about 8–10) is present posteriorly. 

Few changes occur during the fl exion and postfl exion 

stages. Two melanophores persist over the hindbrain but 

may be diffi cult to see as they become embedded in late 

postfl exion larvae. Melanophores on the dorsal surface 

of the gut decrease to about six and move dorsad to the 

lower margin of the body wall over the gut. The spots on 

the hindgut coalesce into one large spot. A few external 

hypaxial spots appear on the body in the area of the caudal peduncle. 

Figure 10

Distribution and abundance of larvae of Decorated Warbonnet (Chirolophis decoratus) collected by bongo net in the Bering 

Sea, Gulf of Alaska, and North Pacific Ocean off the Oregon coast. Isobath=1000 m.
22 Professional Paper NMFS 15 

Meristic features (Table 10). Principal caudal-fi n rays 

in C. decoratus are present by 10.2 mm and begin to 

ossify by 13.7 mm before all are developed; development and ossifi cation are complete by 16.8 mm. Procurrent caudal-fi n rays are present by 16.8 mm, developed 

and partially ossifi ed by 17.9 mm, and fully ossifi ed by 

25.5 mm. Dorsal-fi n and anal-fi n elements are present 

by 17.9 mm. The dorsal-fi n spines are ossifi ed posteriorly by 21.0 mm, and all dorsal-fi n and anal-fi n elements 

are developed and ossifi ed at 25.5 mm. The count of LX 

dorsal-fi n spines observed on radiographs is lower than 

the count previously reported for this species, and the 

count of LXIV dorsal-fi n spines observed in the 42.5-mm 

specimen is greater than the count previously reported 

for this species (LXI–LXIII; Matarese et al., 1989). Pectoral-fi n rays are present by 13.7 mm, developed and 

partially ossifi ed by 17.9 mm, and fully ossifi ed by 21.5 

mm. The count of 16 pectoral-fi n rays observed on radiographs is greater than the count previously reported 

for this species (13–15; Mecklenburg et al., 2002). Pelvic-fi n elements are present by 25.5 mm and are developed and ossifi ed by 28.5 mm. Vertebral centra are present at 9.1 mm. Ossifi cation of the vertebral centra begins 

by 16.8 mm before all are developed; all vertebral centra 

are developed and ossifi ed from 21.0 to 22.5 mm. Neural and haemal spines begin to ossify by 16.8 mm and 

Figure 11

Distribution and abundance of larvae of Decorated Warbonnet (Chirolophis decoratus) collected by neuston gear in the 

Bering Sea and Gulf of Alaska. Isobath=1000 m.
23

Figure 12

Larvae of Decorated Warbonnet (Chirolophis decoratus): (A) yolk-sac stage, 8.7 mm standard length (SL), University 

of Washington (UW) 145169; (B) late preflexion stage, 9.5 mm SL, UW 145169; (C) flexion stage, 13.0 mm SL, UW 

69771; (D) early postflexion stage, 21.5 mm SL, composite of 21.5- and 22.0-mm SL specimens, UW 145168 and UW 

145161; and (E) late postflexion stage, 32.0 mm SL, UW 145160. Illustrations (A–D) by B. Vinter (A–B reared) and (E) 

by A. Overdick.
24 Professional Paper NMFS 15 

Table 9

Body proportions of Decorated Warbonnet (Chirolophis decoratus) larvae. Except for standard length (SL), 

values given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± standard deviation, and range (in parentheses).

 Prefl exion Flexion Postfl exion

Sample size 21 8 25

Standard length (mm) 10.3±1.0 (8.9–11.9) 15.6±1.7 (13.1–17.6) 27.1±3.6 (20.9–31.6)

Head length/SL 13.8±1.6 (11.0–17.9) 13.9±0.9 (12.8–15.3) 15.9±1.3 (13.3–19.7)

Snout length/HL 14.8±4.6 (7.1–25.7)1 17.5±3.3 (12.8–22.8) 25.1±2.6 (18.7–30.9)

Eye diameter/HL 39.6±5.6 (28.1–49.4) 31.1±2.4 (28.3–34.2) 24.3±2.7 (16.4–29.3)

Snout to anus length/SL 37.8±1.0 (34.7–39.0) 34.3±2.5 (28.3–35.9) 35.4±1.8 (31.8–41.3)

Body depth/SL 5.4±0.7 (4.6–7.4) 6.2±0.8 (4.8–7.0) 8.9±0.8 (7.5–10.9)

1 n=20

Table 10

Meristic counts of cleared and stained Decorated Warbonnet (Chirolophis decoratus) larvae. Counts are of ossifi ed elements 

only. Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

9.1 

 10.2 

 11.4 

 12.5 

 13.7 4+5

 14.7 6+7

 16.8 4 513 55 51 49 49 6+8

 17.9 131 16 373 53 37 16 49 65 3,6+8,2

 19.5 17 463 63 46 17 462 63 3,6+8,1

 21.0 15 5 17 513 68 52 17 52 69 4,6+8,2

 21.0 X 15 6 17 513 68 52 17 52 69 4,6+8,2

 21.5 14 4 16 473 50 47 16 472 63 2,6+8,1

 22.5 15 6 16 523 68 51 16 52 68 4,6+8,2

 23.0 14 6 16 523 67 52 16 52 68 2,6+8,1

 25.5 XXXIII I,45 15 6 16 503 66 50 16 51 67 5,6+8,3

 28.5 LXIII I,48 15 I,3 6 16 513 67 51 16 52 68 6,6+8,4

 31.5 LXIII I,48 15 I,3 6 16 523 68 52 16 52 68 6,6+8,4

 42.5 LXIV I,49 15 I,3 * 16 523 68 52 16 53 69 ***

1 Dorsal-fi n and ventralmost pectoral-fi n rays not completely formed or ossifi ed. 2 Posteriormost 4–5 vertebral centra, neural and haemal spines unossifi ed. Ural centra ossifi ed in both specimens. 3 Two neural spines on preural centra.

* Character damaged or missing. 

Neural spines Centra

Spines, rays
25

are developed and ossifi ed by 21.0 mm. Ranges of adult 

meristic counts are given in Table 2.

Species comparisons. Larvae of C. decoratus resemble 

species of Bryozoichthys, particularly during the prefl exion and fl exion stages, but they can be distinguished by 

the absence of a continuous row of evenly spaced, internal melanophores that extend from the nape to the 

hypural region on the dorsal surface of the notochord. 

Species of Chirolophis have a row of internal dorsolateral pigment similar to species of Bryozoichthys, but in 

Chirolophis species it is present only on the postanal 

body. Melanophores around the notochord tip, caudal 

fi nfold, and caudal fi n in Bryozoichthys species are absent in C. decoratus at all stages of development. Chirolophis decoratus usually has fewer vertebrae (myomeres; 

65–69) than do Bryozoichthys lysimus (69–75) and B. 

marjorius (71–76).

Larval C. decoratus are most likely to be confused 

with Mosshead Warbonnet (C. nugator) but can be distinguished in the prefl exion and fl exion stages by the 

presence of a row of several evenly spaced melanophores 

along the dorsal midline on the caudal peduncle; this 

row is absent in C. nugator. Pigment may be present or 

absent on the ventral surface of the gut in C. decoratus, 

but it is always present in C. nugator as a row of several melanophores or a continuous line throughout development. Also, prefl exion C. decoratus have >40 PVM, 

whereas C. nugator have <40 PVM during the prefl exion 

stage. Larvae of C. decoratus are more slender-bodied 

than are larvae of C. nugator throughout development 

(mean body depth: 5.4–8.9% SL vs. 6.9–9.9% SL) and 

have a shorter mean snout length during the prefl exion 

and fl exion stages (14.8–17.5% HL vs. 20.0–22.9% 

HL). In addition, C. decoratus has more vertebrae (myomeres; 65–69) than does C. nugator (55–61) and more 

dorsal-fi n and anal-fi n elements (LX–LXIV; I–II,44–49 

vs. LIII–LVI; I–II,37–42). 

Chirolophis nugator, Mosshead Warbonnet 

(Figs. 13, 14; Tables 11, 12)

Literature. Hart (1973), Eschmeyer et al. (1983), Mecklenburg et al. (2002).

Material examined. 76 specimens (8.7–25.0 mm) 

from the Bering Sea; Gulf of Alaska; the coast of British Columbia, Canada; Straits of Georgia, Canada; Puget 

Sound, Washington; and the coasts of Oregon and California (see Appendix for details).

Distribution and life history. Chirolophis nugator ranges throughout the Aleutian Islands and along the Alaska 

Peninsula to southern California in the North Pacifi c 

Ocean in intertidal and subtidal areas to a depth of 80 m 

(Eschmeyer et al., 1983), although it is usually found in 

water less than 20 m deep, sometimes in holes or inside 

empty shells. Adults reach 150 mm TL (Mecklenburg et 

al., 2002). Eggs are 2 mm in diameter, and spawning may 

occur in January (Hart, 1973); larvae are pelagic.

Larval distribution (Fig. 13). Larvae of C. nugator have 

been collected over the shelf in the Bering Sea, in Unimak Pass, and eastward into the Gulf of Alaska to Kodiak Island, as well as off the coast of British Columbia, 

Canada, to northern California. Larvae (11.0–26.0 mm, 

mean 16.4 mm) are collected in the plankton in January–June with the highest abundances observed in May.

Morphology (Tables 11, 12). Notochord fl exion in C. 

nugator begins by 12.3 mm and is complete around 17.0 

mm. Postfl exion larvae are 17.1–25.0 mm. Transformation begins after 25.0 mm; length at the beginning of 

the juvenile stage is unknown. Larvae are slender-bodied with mean body depth of 9.2% SL in the prefl exion 

stage, decreasing to 6.9% SL in the fl exion stage and increasing to 9.9% SL in the postfl exion stage. Relative 

head length increases slightly, and snout length increases 

more, during larval development. Relative eye diameter 

decreases during development while snout to anus length 

remains relatively constant.

Pigmentation (Fig. 14). Larvae of C. nugator are moderately pigmented. A large spot on the hindbrain persists 

throughout development. Initially, the nape region has 

several spots. About six internal, stellate melanophores 

occur on the dorsal surface of the gut, and several external melanophores occur dorsally on the hindgut. Internal 

pigment occurs in the area of the cleithrum. Pigment on 

the ventral surface of the gut is restricted to a series or 

continuous line of melanophores at midgut. A series of 

about 33 PVM begins about fi ve myomeres posterior to 

the anus with a melanophore on almost every myoseptum along the entire length of the postanal body. Internal 

melanophores are present dorsally above the notochord. 

Several pigmentation changes occur during the fl exion 

and postfl exion stages. The large spots on the head become more prominent, and pigment on the nape becomes 

internal. One or more additional spots are present in the 

opercular area in postfl exion larvae. The melanophores on 

the dorsal surface of the gut become spaced farther apart 

as the larvae develop. The cleithral pigment becomes more 

prominent during the fl exion stage, and then it is no longer visible in the postfl exion stage. The ventral midline pigment on the gut is highly variable: some larvae have discrete 

melanophores at midgut, and others have some discrete 

melanophores together with a continuous line of pigment. 

PVM remain <40 (35–37), and internal pigment above the 

notochord remains visible posteriorly. Large melanophores 

appear along the dorsal midline, restricted mostly to the 

posterior region but occasionally extend anteriorly. A few 

small spots may appear along the hypural edge. 
26 Professional Paper NMFS 15 

Meristic features (Table 12). Principal caudal-fi n rays 

are present in C. nugator by 10.1 mm and are all developed and weakly ossifi ed at 14.3 mm. The count of 8 inferior principal caudal-fi n rays observed in the 17.1-mm 

specimen is greater than the count previously reported 

for this species (7; Watson, 1996). Procurrent caudal-fi n 

rays are present by 16.1 mm, begin to ossify by 17.1 

mm, and are developed and ossifi ed by 23.0 mm. The 

counts of 3 superior and 1 inferior procurrent caudal-fi n 

rays observed on radiographs are lower than the counts 

previously reported for this species (5–6, 3–4; Watson, 

1996). By 17.1 mm, the dorsal-fi n and anal-fi n elements 

are present. Development and ossifi cation of the anal-fi n 

elements can be complete as early as 18.5 mm. The count 

of II anal-fi n spines observed on radiographs is greater 

than the count previously reported for this species (I; 

Mecklenburg et al., 2002). The dorsal-fi n spines are developed by 18.5 mm and ossifi ed by 23.0 mm. The count 

of LVI dorsal-fi n spines observed on radiographs is greater than the count previously reported for this species 

(LIII–LV; Mecklenburg et al., 2002). Pectoral-fi n rays are 

present by 10.1 mm, developed by 16.1 mm, and weakly 

ossifi ed by 17.1 mm. The pelvic-fi n buds are fi rst discernable at 17.1 mm. Individual elements are present by 18.5 

Figure 13

Distribution and abundance of larvae of Mosshead Warbonnet (Chirolophis nugator) collected in the Bering Sea; Gulf 

of Alaska; North Pacific Ocean off the coasts of British Columbia, Canada, and Oregon; and Puget Sound, Washington. 

Squares indicate collections made by bongo net; black circles indicate additional nonquantitative collections (presence only) 

and may represent more than one occurrence. Isobath=1000 m.
27

Figure 14

Larvae of Mosshead Warbonnet (Chirolophis nugator): (A) preflexion stage, 11.9 mm standard length (SL), University of 

Washington (UW) 145053; (B) flexion stage, 15.7 mm SL, UW 145053; and (C) postflexion 22.5 mm SL, UW 145053. Illustrations by B. Vinter.

mm and developed and ossifi ed by 23.0 mm. Vertebral 

centra are present at 12.0 mm and developed and ossifi ed by 17.1 mm. The count of 16 precaudal centra, 

the range of 42–46 caudal centra, and the total count of 

61 vertebral centra observed on radiographs are greater 

than the counts previously reported for this species (14–

15+43–44=55–59; Watson, 1996). Neural and haemal 

spines are present by 10.1 mm and ossifi ed by 12.0 mm. 

Ranges of adult meristic counts are given in Table 2.

Species comparisons. See the species comparison section for Chirolophis decoratus to distinguish larvae from 

species of Bryozoichthys; similar pigment differences 

apply to C. nugator. Also similar to C. decoratus, C. 

nugator has fewer vertebrae (myomeres: 55–61) than 

both Bryozoichthys species (69–76).

Larvae of C. nugator are most similar to C. decoratus, 

but they can be distinguished in the prefl exion and fl exion stages by the absence of a row of several evenly 

spaced melanophores along the dorsal midline of the 

caudal peduncle; this row is present in C. decoratus. Ventral pigmentation on the gut is always present as a row 

of several melanophores or a continuous line throughout 

development, but in C. decoratus it may be present or 

absent. In the prefl exion stage, C. nugator has <40 PVM, 

whereas C. decoratus has >40. Larvae of C. nugator are 

deeper bodied than are larvae of C. decoratus throughout development (mean body depth: 9.2–9.9% SL vs. 
28 Professional Paper NMFS 15 

5.4–8.9% SL) and have a longer mean snout length during the prefl exion and fl exion stages (20.0–22.9% HL vs. 

14.8–17.5% HL). In addition, C. nugator has fewer vertebrae (myomeres; 55–61 vs. 65–69) and fewer dorsal-fi n 

and anal-fi n elements (LIII–LVI; I–II,37–42 vs. LX–LXIV; 

I–II,44–49) than does C. decoratus.

Chirolophis snyderi, Bearded Warbonnet

See “Other material” section of this guide.

Gymnoclinus cristulatus, Trident Prickleback 

(Fig. 15; Tables 13, 14)

Literature. Shiogaki (1988; reported by Tokuya and 

Amaoka (1980) as Chirolophis otohime), Mecklenburg 

et al. (2002).

Material examined. Eight specimens (18.8–26.8 mm) 

from the Sea of Japan (see Appendix for details).

Table 11

Body proportions of Mosshead Warbonnet (Chirolophis nugator) larvae. Except for standard length (SL), 

values given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± 

standard deviation, and range (in parentheses).

 Prefl exion Flexion Postfl exion

Sample size 19 17 29

Standard length (mm) 10.1±0.8 (8.7–11.4) 13.9±1.2 (12.3–16.8) 20.9±2.0 (17.1–24.8)

Head length/SL 15.1±1.5 (13.2–18.1)1 15.7±1.1 (14.5–18.4) 16.4±1.6 (13.9–22.9)

Snout length/HL 20.0±3.6 (11.6–26.5)2 22.9±2.8 (16.9–27.1) 25.2±3.1 (18.8–30.8)

Eye diameter/HL 38.5±5.2 (29.6–46.2)1 34.9±3.4 (27.8–36.9) 30.8±2.8 (24.7–35.8)

Snout to anus length/SL 36.6±1.4 (33.8–39.5) 36.8±1.0 (35.2–39.5) 37.1±1.0 (35.9–41.3)

Body depth/SL 9.2±1.9 (6.3–12.2)2 6.9±0.7 (5.7–8.1)3 9.9±1.4 (7.8–12.1)

1 n=18 2 n=17 3 n=16

Table 12

Meristic counts of cleared and stained Mosshead Warbonnet (Chirolophis nugator) larvae. Counts are of ossifi ed elements 

only. Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 10.1 

 10.9 

 12.0 16 421 58 42 

 13.2 15 431 58 43 

 14.3 15 431 58 42 6+7

 15.5 *

 16.1 6+7

 17.1 14 6 15 441 59 43 15 44 59 5,6+8,2

 18.5 XXV I,40 13 6 15 441 59 43 15 44 59 4,6+7,2

 19.0 6 14 441 59 43 15 44 59 4,6+7,2

 20.5 62 14 441 58 43 6+7

 23.0 LIV I,40 14 I,3 6 15 431 58 43 15 44 59 5,6+7,4

 25.0 LIV I,40 14 I,3 6 15 431 58 43 15 44 59 6,6+7,5

1 Two neural spines on preural centra. 

2 Right side of specimen counted.

*Character missing.

Neural spines Centra

Spines, rays
29

Distribution and life history. Gymnoclinus cristulatus is found in intertidal areas to a depth of about 40 

m (mainly <20 m) in the Sea of Japan near Hokkaido, 

eastern Kamchatka, Kuril and Commander Islands, and 

Amchitka Island in the western Aleutian Island Archipelago; it has not been collected during AFSC adult surveys. 

Adults can grow to 116 mm TL (Mecklenburg et al., 

2002). Little is known of its life history. Eggs are probably demersal and spawned in an adhesive mass; larvae 

are pelagic.

Larval distribution. Larvae of Trident Prickleback have 

not been collected from our study area. Eight larvae 

(18.8–26.8 mm) collected in April at one location off 

Usujiri, southwest Hokkaido, Japan, were in the postfl exion, transformation, and juvenile stages; specimens in 

the prefl exion and fl exion stages were not available. 

Morphology (Table 13). Length at the end of notochord 

fl exion in G. cristulatus is unknown; transformation begins by 19.0 mm. Juvenile stage begins after 23.2 mm 

but before 25.0 mm. Postfl exion larvae are deep-bodied 

(mean body depth 16.0% SL) with a relatively large head 

(head length 21.2% SL, snout length 19.7% HL), large 

eyes (eye diameter 30.2% HL), and long gut (snout to 

anus length 47.2% SL).

Pigmentation (Fig. 15). Pigment description is based in 

part on Tokuya and Amaoka (1980). Larvae of G. cristulatus are heavily pigmented. The single postfl exion-stage 

larva examined (18.8 mm SL) was already developing 

some initial juvenile pigmentation. The head has a prominent dorsal patch and spots along the snout, lower jaw, 

and opercular area. The nape region is lightly pigmented 

with small melanophores. The gut is pigmented with small 

melanophores along the dorsal and lateral surfaces, in addition to one large internal dorsal spot at midgut. The ventral surface of the gut is unpigmented. The dorsal fi nfold 

is lightly pigmented with small, scattered melanophores. 

Laterally, anterior pigment consists of large punctate melanophores above the gut; postanal lateral pigment is fi ne 

and scattered over most of the body. A line of pigment 

extends along the dorsal margin of the body from midgut 

to the caudal peduncle. A similar line extends along the 

ventral margin from the anus to about 80% SL. Light pigment is visible on the anal and caudal-fi n membranes.

Figure 15

Larvae of Trident Prickleback (Gymnoclinus cristulatus): (A) postflexion stage, 18.8 mm standard length (SL), Hokkaido 

University Museum of Zoology (HUMZ-L) 03113; and (B) transformation stage, 24.9 mm SL, HUMZ-L 02569. Illustrations by A. Overdick.
30 Professional Paper NMFS 15 

Table 13

Body proportions of Trident Prickleback (Gymnoclinus cristulatus) larvae. Except for standard length (SL), 

values given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± 

standard deviation, and range (in parentheses).

 Prefl exion Flexion Postfl exion

Sample size 0 0 7

Standard length (mm) 21.9±1.9 (18.8–24.9)

Head length/SL 21.2±1.5 (19.0–23.1)

Snout length/HL 19.7±2.0 (17.1–22.7)

Eye diameter/HL 30.2±2.3 (26.9–32.9)

Snout to anus length/SL 47.2±1.8 (44.5–48.9)

Body depth/SL 16.0±1.1 (14.1–17.2)

Table 14

Meristic counts of cleared and stained Trident Prickleback (Gymnoclinus cristulatus) postfl exion larva (21.0 mm) and juvenile (26.8 mm). Counts are of ossifi ed elements only.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 21.0 LX 42 142 6 18 471 65 471 18 47 65 5,6+7,3

 26.8 LXI 42 152 I3 7 19 461 65 471 19 47 66 5,6+7,2

1 Two neural or haemal spines on second preural centra.

2 Ventralmost pectoral-fi n ray (16) not ossifi ed. 3 Two pelvic-fi n rays present but not ossifi ed.

Neural spines Centra

Spines, rays

Pigmentation increased in early transformation specimens. The juvenile pigment pattern on the head, with 

large dark melanophores on the crown and a streak of 

pigment that extends posterioventrally from the eye, is 

visible by about 20 mm SL. Additional melanophores are 

present along the pectoral-fi n base. The postanal body 

is almost entirely pigmented with tiny spots. Along the 

body midline, up to 10 circular spots are unpigmented. 

Patches of pigment occur throughout the dorsal-fi n rays; 

similar pigment patches are just beginning to extend 

onto the anal-fi n rays. 

Meristic features (Table 14). Vertebral centra, neural and haemal spines, and all fi n elements except the 

pelvic fi ns are developed and ossifi ed by 21.0 mm. 

The count of LVIII dorsal-fi n spines observed in the 

24.9-mm illustrated transformation-stage specimen 

is lower than the count previously recorded for this 

species (LXI–LXIII: Shiogaki, 1988; LXI: Mecklenburg et al., 2002). Two pelvic-fi n rays are present and 

the single pelvic-fi n spine is developed and ossifi ed by 

26.8 mm. Ranges of adult meristic counts are given in 

Table 2. 

Species comparisons. Prefl exion and fl exion larvae of 

G. cristulatus are unknown, but postfl exion larvae are 

distinguished from all presently known stichaeid larvae 

by melanophores scattered over the entire dorsal region 

of the gut and lateral body. They also have the greatest 

body depth during postfl exion (16.0% SL vs. 6.3–12.3% 

SL). The absence of an anal-fi n spine is useful in identifi -

cation of transformation-stage G. cristulatus.

Subfamily Lumpeninae

Acantholumpenus mackayi, Blackline Prickleback

See “Other material” section of this guide.

Lumpenella longirostris, Longsnout Prickleback 

(Figs. 16, 17; Tables 15, 16)

Literature. Amaoka et al. (1983), Mecklenburg et al. 

(2002).

Material examined. 126 specimens (8.0–64.0 mm) 

from the Bering Sea and Gulf of Alaska (see Appendix 

for details).
31

Distribution and life history. Lumpenella longirostris 

adults occur in the Sea of Okhotsk to northern Japan, 

the Bering Sea, Aleutian Islands to southern British Columbia, and in the North Atlantic Ocean near Greenland. 

Although found most commonly at depths of 100–200 

m (RACEBASE), Longsnout Prickleback have been collected at depths of 22–1140 m. Maximum length is 411 

mm TL (Mecklenburg et al., 2002). Little is known of its 

life history. Eggs are probably demersal and spawned in 

an adhesive mass; larvae are pelagic.

Larval distribution (Fig. 16). Larvae of the fourthmost abundant stichaeid in our study area, Longsnout 

Prickleback (8.4–41.0 mm), are found during April–June, 

with the highest abundance observed in May, on the 

shelf in the Gulf of Alaska south of the Alaska Peninsula, 

in Shelikof Strait, and along the shelf east of Kodiak Island to the Kenai Peninsula. Although adults occur in the 

central and southeastern Bering Sea, larvae are collected 

infrequently in these regions. 

Morphology (Table 15). The smallest prefl exion larval 

L. longirostris examined (8.0 mm) had a small yolk sac. 

Notochord fl exion begins at about 13.0 mm and is complete by 25.0 mm. Postfl exion larvae are 24.9–40.4 mm. 

Transformation begins after 40.4 mm, and the juvenile 

Figure 16

Distribution and abundance of larvae of Longsnout Prickleback (Lumpenella longirostris) collected by bongo net in the 

Bering Sea and Gulf of Alaska. Isobath=1000 m.
32 Professional Paper NMFS 15 

Figure 17

Larvae of Longsnout Prickleback (Lumpenella longirostris): (A) preflexion stage, 9.6 mm standard length (SL), University 

of Washington (UW) 67120; (B) flexion stage, 11.8 mm SL, UW 94969; (C) flexion stage, 17.5 mm SL, UW 145094; (D) 

postflexion stage, 27.5 mm SL, UW 68867; and (E) late postflexion stage, 40.0 mm SL, UW 145093. Illustrations (A, C–E) 

by B. Vinter and (B) by A. Overdick. 
33

stage begins by 50.0 mm. Larvae are slender-bodied with 

mean body depth of 7.6% SL in the prefl exion stage, 

decreasing to 5.7% SL during the fl exion stage and 

increasing slightly to 6.5% SL in the postfl exion stage. 

Both relative head length and snout length decrease from 

the prefl exion stage to the fl exion stage and increase 

again in the postfl exion stage. Relative eye diameter 

decreases throughout larval development, and relative 

snout to anus length is nearly constant, decreasing only 

slightly. 

Pigmentation (Fig. 17). Larvae of L. longirostris are 

lightly pigmented (pigment on postanal body is limited 

to PVM). The head is unpigmented in prefl exion larvae. 

The gut is pigmented on its lateral surface with about 

5–6 melanophores. One anterior spot is initially located 

on the ventral body margin. Two melanophores occur on

the dorsal surface of the hindgut. A single melanophore

appears ventral to the pectoral fi n base. A series of about

50 PVM, consisting of a melanophore at each myoseptum, 

begins just after the anus and extends to the caudal 

Table 15

Body proportions of Longsnout Prickleback (Lumpenella longirostris) larvae and juveniles. Except for standard length (SL), values given for each body proportion are expressed as percentage of SL or head length 

(HL): mean, ± standard deviation, and range (in parentheses).

 Prefl exion Flexion Postfl exion

Sample size 40 51 14

Standard length (mm) 10.3±1.3 8.0–12.9) 18.2±4.0 (13.1–24.9) 29.9±5.4 (24.9–40.4)

Head length/SL 14.0±2.1 (10.0–19.3) 12.7±1.2 (9.2–15.8) 13.1±1.1 (11.0–15.6)

Snout length/HL 26.1±6.9 (13.0–42.2)1 22.5±4.9 (9.3–35.0) 28.4±2.8 (23.6–33.0)

Eye diameter/HL 33.4±5.5 (24.2–46.7) 25.0±4.5 (16.9–34.7)2 19.5±1.6 (16.9–22.2)

Snout to anus length/SL 44.8±3.4 (32.2–50.7) 42.8±1.5 (38.1–45.8) 42.6±1.0 (40.7–44.8)

Body depth/SL 7.6±1.9 (3.9–9.9) 5.7±0.7 (4.0–7.0) 6.5±0.4 (5.8–7.0)

1 n=37 2 n=50

Table 16

Meristic counts of cleared and stained Longsnout Prickleback (Lumpenella longirostris) larvae and juveniles. Counts are of 

ossifi ed elements only. Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 9.0 

 9.4 

 9.7 

 10.1 

 12.0 

 12.9 

 13.3 

 15.9 

 17.6 

 19.8 

 20.0 23 20 43 

 22.5 22 48 70 3,6+6,1

 26.5 23 48 71 6+6,1

 32.5 XL IV,42 6 22 47 69 47 22 48 70 4,6+6,4

 38.0 LXIV IV,42 13 I,3 6 22 47 69 47 22 48 70 5,6+6,5

 49.0 LXVI IV,41 12 I,3 6 23 49 72 49 23 49 73 5,6+6,4

 64.0 LXVI IV,41 13 I,3 6 23 48 71 48 23 49 72 5,6+6,6

Neural spines Centra

Spines, rays
34 Professional Paper NMFS 15 

peduncle. Caudal pigment is restricted to small melanophores around the notochord tip and in the hypural area.

In fl exion larvae, the melanophore ventral to the pectoral 

fi n is larger and more prominent. The number of melanophores along the lateral surface of the gut behind the pectoral fi n increases up to 10, including 1–2 on the hindgut. The 

number of PVM decreases to <50. Light caudal-fi n pigment 

was visible in some specimens by 17 mm. 

Pigment changes in postfl exion larvae occur mainly 

on the head and tail. Pigment forms in the otic area. Additional pigment is visible on the caudal fi n by about 28 

mm, but it is more concentrated on the distal half. Larger specimens had essentially the same pigment pattern, 

with a series of melanophores on the isthmus as well as a 

melanophore at the pectoral-fi n base. About six melanophores remain on the lateral surface of the gut, with another two spots that extend onto the hindgut. Although 

PVM (42–44) become embedded, they are still visible to 

about 40 mm. Pigment remains visible around the notochord tip and continues to increase on the caudal fi n.

Meristic features (Table 16). Principal caudal-fi n rays 

are present between 9.7 and 10.1 mm, developed by 

20.0 mm, and ossifi ed by 22.5 mm. Procurrent caudalfi n rays are present by 20.0 mm, and ossifi cation begins 

at 22.5 mm before all are developed; development and 

ossifi cation are complete by 32.5 mm. Dorsal-fi n spines 

and anal-fi n rays are present by 22.5 mm. Anal-fi n elements are developed by 26.5 mm and ossifi ed by 32.5 

mm. Ossifi cation of dorsal fi n spines begins at 32.5 mm, 

progresses from posterior to anterior, and is complete 

at 38.0 mm. Pectoral-fi n rays are present by 20.0 mm, 

developed by 22.5 mm, and ossifi ed by 38.0 mm. Pelvicfi n elements are present at 32.5 mm. The pelvic-fi n spine 

is developed and ossifi ed by 38.0 mm, and the pelvic-fi n 

rays are developed and ossifi ed by 49.0 mm. Vertebral 

centra are present at 15.9 mm. All centra and neural and 

haemal spines are developed by 19.8 mm. Ossifi cation 

of vertebral centra begins at 20.0 mm and progresses 

from anterior to posterior; all vertebral centra are ossifi ed by 22.5 mm. The count of 22 precaudal centra observed in the 32.5- and 38.0-mm specimens is lower than 

the count previously reported for this species (23–25; 

Amaoka et al., 1983). The total count of 70 vertebral 

centra observed in the 32.5- and 38.0-mm specimens is 

lower than the total count previously reported for this 

species (71–75; Mecklenburg et al., 2002). All neural and 

haemal spines are ossifi ed by 32.5 mm. Ranges of adult 

meristic counts are given in Table 2. 

Species comparisons. Larvae in the subfamily 

Lumpeninae (Lumpenella, Lumpenus, and Poroclinus) 

can be distinguished from other stichaeids by a combination of characters that include typically slender bodies at 

all stages of development (mean body depth 5.7–9.5% 

SL; Lumpenus medius has a mean body depth during 

the fl exion stage of 10.0% SL), long guts (snout to anus 

length 42.5–50.4% SL), and sparse pigmentation typically restricted to the lateral or dorsal surfaces of the gut, 

PVM, and caudal fi n region.

Larval Lumpenella longirostris are similar to larvae of 

Lumpenus species and Whitebarred Prickleback (Poroclinus rothrocki). Prefl exion- and fl exion-stage L. longirostris can be distinguished by the presence of fi ne melanophores around the notochord tip; postfl exion larvae have 

pigment along the dorsal edge of the hypural. Species 

of Lumpenus and P. rothrocki have no pigment around 

the notochord tip at any stage and no pigment along 

the dorsal edge of the hypural. Poroclinus rothrocki has 

an irregularly spaced row of embedded melanophores, 

both above and below the notochord on the caudal peduncle; this row is absent in L. longirostris. Larvae of 

L. longirostris are more slender-bodied than are larvae 

of P. rothrocki at all stages of development (mean body 

depth: 7.6% SL, 5.7% SL, 6.5% SL vs. 9.5% SL, 6.3% 

SL, 8.5% SL). Lumpenella longirostris has more total 

vertebrae (70–75) than does P. rothrocki (64–68), usually 

has more vertebrae than does Lumpenus medius (65–

70), and usually has fewer vertebrae than does Snake 

Prickleback (Lumpenus sagitta; 75–82). It is worth noting that among late-postfl exion-through-transformationstage stichaeid larvae only L. longirostris has 4–5 analfi n spines.

Lumpenus fabricii, Slender Eelblenny 

(Figs. 18, 19; Tables 17, 18)

Literature. Grigor’ev (1992b), MacDonald (2001), 

Mecklenburg et al. (2002), Mecklenburg and Sheiko 

(2004).

Material examined. 22 specimens (12.1–57.0 mm) 

from the Chukchi, Beaufort, and Bering seas and Gulf of 

Alaska (see Appendix for details). 

Distribution and life history. Lumpenus fabricii is 

a circumarctic species. It is found in the seas of Japan 

and Okhotsk; Beaufort, Chukchi, and Bering seas; in 

the Aleutian Islands as far west as Unimak Island; along 

shelf areas of the Gulf of Alaska to southeastern Alaska; 

and in the Arctic and North Atlantic oceans. Slender Eelblenny have been collected over sandy to rocky bottoms 

to depths of 175 m, but this species is usually found on 

the inner shelf <50 m deep, often in seagrass or algae 

(Mecklenburg et al., 2002; Mecklenburg and Sheiko, 

2004). Adults can grow to 365 mm TL (Mecklenburg 

et al., 2002). Little is known of their life history. Eggs 

are probably demersal and spawned in an adhesive mass; 

larvae are pelagic.

Larval distribution (Fig. 18). Larvae of Slender Eelblenny

(11.9–46.0 mm) have been collected infrequently during 
35

May–July in the Chukchi Sea near Wrangel Island and 

the Chukchi Peninsula; Beaufort Sea off the coast of 

Northwest Territories, Canada; on the Bering Sea shelf; 

along the shelf in the northern Gulf of Alaska east of 

Unimak Pass; and close to shore along the southeast side 

of Kodiak Island. Juveniles (55.0–57.0 mm) have been 

collected in the Chukchi Sea in September. Larvae also 

have been collected in June–July in the Sea of Okhotsk 

near the Kamchatka Peninsula (Grigor’ev, 1992b).

Morphology (Table 17). Notochord fl exion in L. 

fabricii begins at 13.0 mm and is complete by 21.0 

mm. Postfl exion larvae are 20.0–46.0 mm. Transformation begins after 45.0 mm, and the juvenile stage 

begins after 55.0 mm. Larvae are slender-bodied with 

a mean body depth of 8.3% SL and 6.5% SL for the 

prefl exion (n=1) and fl exion stages, respectively, increasing slightly to 6.9% SL in the postfl exion stage. 

Relative head length changes little through the fl exion stage, and then it increases in postfl exion larvae. 

Relative snout length increases from the fl exion stage 

to the postfl exion stage, and relative eye diameter 

and snout to anus length decrease throughout larval 

development.

Figure 18

Distribution of larvae of Slender Eelblenny (Lumpenus fabricii) collected by bongo and Tucker gear in the Chukchi and 

Bering seas and Gulf of Alaska. Each symbol may indicate more than one occurrence. Isobath=1000 m.
36 Professional Paper NMFS 15 

Pigmentation (Fig. 19). Larvae are lightly pigmented. In 

the only prefl exion larva available for examination, the 

head was unpigmented, a single melanophore appeared 

ventral to each pectoral fi n, the gut had a row of melanophores along its dorsolateral and lateral surface that 

included the hindgut (about 6–7 + 1–2 hindgut melanophores spaced irregularly throughout), and a PVM series 

(about 40–45) that consisted of one melanophore per 

myoseptum extended from near the anus to the caudal 

peduncle. Caudal pigment is restricted to several small 

melanophores in the hypural area. 

In the fl exion and postfl exion stages, the melanophore 

below the pectoral-fin base becomes larger and more 

prominent, and up to 12 melanophores appear along the 

dorsolateral and lateral surface of the gut posterior to the 

pectoral fi n, including several spots on the hindgut. PVM 

decrease in number to 33–37. A band of pigment is present 

along the base of the developing caudal-fi n rays. 

Transformation-stage larvae develop melanophores 

on the upper jaw tip and cheek. Preanal pigment consists of a series of 3–4 ventral dashes on the isthmus, 1–2 

spots near the base of the pelvic fi ns, and lateral spots 

along the gut (4–8 + 2 on hindgut); the spot below the 

pectoral-fi n base that was prominent in fl exion and postfl exion larvae becomes deeply embedded and diffi cult to 

see. The number of PVM increases to 37–39, beginning 

several myomeres posterior to the anus; melanophores 

are embedded in tissue at the base of the anal-fi n elements. Clusters of pigment are present along the lateral 

line and scattered in 2–3 areas above and below it on 

Figure 19

Larvae of Slender Eelblenny (Lumpenus fabricii): (A) postflexion stage, 20.5 mm standard length (SL), University of Washington (UW) 145063; (B) early transformation stage, 46.0 mm SL, composite of 41.0 and 46.0 mm SL specimens, UW 

145896 and UW 94975; and (C) juvenile, 55.0 mm SL, UW 145069. Illustrations by A. Overdick.
37

the posterior one-third of the body; scattered melanophores along the dorsal midline are present at 40% and 

80–90% SL. Pigment at the hypural edge extends onto 

the caudal-fi n rays.

In juveniles, irregularly shaped, diffuse patches and 

streaks of melanophores are present on the snout, crown, 

and lateral surfaces of the head and opercle, but there is 

little pigment ventrally on the head. A single melanophore 

is present on the pectoral-fi n base near the insertion of 

the medial rays. Irregularly spaced blotches and patches 

of pigment along the dorsal midline extend into the dorsal fi n around many of the spines. The body is covered 

dorsolaterally with irregularly shaped patches of pigment 

and with uniform, oval-shaped patches spaced regularly 

along the lateral line. Five or more evenly spaced clusters 

of melanophores are present along the postanal ventral 

midline. Additional patches of pigmentation at the hypural edge extend onto the caudal-fi n rays.

Meristic features (Table 18). Few larvae of L. fabricii

were available to clear and stain. All median and paired 

fi n elements, vertebral centra, and neural and haemal 

spines are developed and ossifi ed by 41.0 mm during 

postfl exion. The count of LX dorsal-fi n spines observed 

in the 55.0-mm illustration specimen is lower than the 

count previously reported for this species (LXI–LXVI; 

Mecklenburg et al., 2002). The count of 38 anal-fi n rays 

obtained from MacDonald (2001) is lower than the 

count otherwise reported for this species (40–44; Mecklenburg et al., 2002). The count of 23 precaudal centra obtained from MacDonald (2001) is lower than the 

count previously reported for this species (26–28; Matarese et al., 1989). The range of 43–49 caudal centra obtained from MacDonald (2001) is greater than the range 

previously reported for this species (44–48; Matarese et 

al., 1989). By 57.0 mm, scales begin to form. Ranges of 

adult meristic counts are given in Table 2.

Table 17

Body proportions of Slender Eelblenny (Lumpenus fabricii) larvae. Except for standard length (SL), values 

given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± standard 

deviation, and range (in parentheses). 

 Prefl exion Flexion Postfl exion

Sample size 1 3 11

Standard length (mm) 12.1 16.2±4.3 (13.0–21.0) 33.5±8.9 (20.0–46.0)

Head length/SL 11.6 11.4±0.9 (10.8–12.4) 15.3±3.1 (10.4–19.1)

Snout length/HL – 1 23.2±2.5 (21.4–24.9)2 27.2±4.3 (21.6–32.7)3

Eye diameter/HL 35.7 31.2±10.7 (19.0–38.9) 21.3±5.2 (16.2–32.7)

Snout to anus length/SL 50.4 46.1±2.0 (43.9–47.6) 43.1±4.0 (33.7–49.8)

Body depth/SL 08.3 6.5±1.0 (5.6–7.7) 6.9±1.2 (5.4–9.3)3

1 n=0 2 n=2 3 n=9

Table 18

Meristic counts of cleared and stained Slender Eelblenny (Lumpenus fabricii) larvae and juveniles. Counts are of ossifi ed 

elements only. Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 12.5 

 13.3 

 19.5* 

 22.5 

 41.0 LXIII I,42 14 I,3 6 24 45 69 45 24 46 70 5,6+7,4

 57.0 LXIII I,42 16 I,3 6 26 45 71 45 26 46 72 6,6+7,6

* Damaged caudal fi n.

Neural spines Centra

Spines, rays
38 Professional Paper NMFS 15 

Species comparisons. Pigmentation patterns are not 

useful for species-level identifi cation of larvae of the genus Lumpenus; species are distinguished primarily by a 

combination of morphological and meristic differences 

that are present in older, more developed larvae. Larval 

L. fabricii are most similar to other species of Lumpenus, 

particularly L. medius. Although this measurement was 

based on one specimen, mean body depth appeared greater in prefl exion L. fabricii (8.3% SL) than in prefl exion 

L. medius and L. sagitta (7.0% SL, 7.7% SL). During 

the fl exion and postfl exion stages, larvae of L. fabricii

have a shorter gut (mean snout to anus length: 46.1% 

SL, 43.1% SL) than do larvae of L. maculatus (49.4% 

SL, 50.3% SL) and are more slender-bodied (mean body 

depth: 6.5% SL, 6.9% SL) than are larvae of L. maculatus (mean body depth: 7.9% SL, 7.5% SL) and L. medius (mean body depth: 10.0% SL, 7.9% SL). Transformation in L. fabricii begins at a larger size (>45 mm) than 

it does in L. medius (>35 mm). In transformation-stage 

and early juvenile L. fabricii, the dorsal and anal fi ns are 

not connected to the caudal fi n as they are in L. medius

(Mecklenburg et al., 2002). Counts of precaudal, caudal, 

and total myomeres or vertebrae (23–28+43–49=70–75) 

distinguish larvae and juveniles of L. fabricii from other 

species of Lumpenus. Lumpenus fabricii typically has 

more precaudal vertebrae (23–28) than does L. medius

(21–25), typically fewer precaudal vertebrae than does 

L. sagitta (26–29), more caudal vertebrae (43–49) than 

does L. maculatus (39–43), and usually fewer total vertebrae (70–75) than does L. sagitta (75–82).

Lumpenus maculatus, Daubed Shanny 

(Figs. 20, 21; Tables 19, 20)

Literature. Faber (1976), Eschmeyer et al. (1983), 

Grigor’ev (1992b), MacDonald (2001), Mecklenburg et 

al. (2002).

Material examined. 80 specimens (8.1–72.1 mm) from 

the Bering Sea and Gulf of Alaska (see Appendix for 

details). 

Distribution and life history. Lumpenus maculatus is 

found over mud, sand, stone, or pebble bottoms at depths 

of 2–475 m (almost always <170 m) (Mecklenberg et al., 

2002) in the seas of Okhotsk and Japan, Beaufort and 

Bering seas, in the Aleutian Islands as far west as Unalaska Island, throughout the Gulf of Alaska, and south 

to Puget Sound, Washington. It also has been reported in 

the North Atlantic Ocean to Spitsbergen Island. Adults 

can grow to 180 mm TL (Eschmeyer et al., 1983) in the 

North Pacifi c Ocean and to 200 mm TL near Iceland 

(Mecklenburg et al., 2002). Little is known of its life 

history. Eggs are probably demersal and spawned in an 

adhesive mass; larvae are pelagic.

Larval distribution (Fig. 20). Larvae of Daubed Shanny 

have the highest frequency of occurrence of any stichaeid 

in our study area. Daubed Shanny at lengths of 8.0–64.0 

mm (mostly 10–25 mm) are collected in March–October, with the highest abundance observed in May, from 

the Chukchi Sea near Wrangel Island, the Beaufort Sea, 

Bering Strait and on the Bering Sea shelf, throughout 

the shelf in the northern Gulf of Alaska east of Unimak 

Pass, around Kodiak Island, and eastward to the mouth 

of Cook Inlet. Larvae also have been collected in June in 

the Sea of Okhotsk near the Kamchatka Peninsula and in 

the Gulf of Anadyr (Grigor’ev, 1992b).

Morphology (Tables 19, 20). Size of larvae at the beginning and end of developmental stages was more variable 

in L. maculatus than in other species examined in this 

study. Notochord fl exion begins at 13.0–15.0 mm and 

can be complete at 20.0–25.0 mm. Postfl exion larvae are 

20.0–66.2 mm. Transformation begins at about 50 mm, 

and the juvenile stage begins by 72.1 mm. Larvae are 

slender-bodied and become more so with development, 

with mean body depth decreasing from 8.3% SL in the 

prefl exion stage to 7.5% SL in the postfl exion stage. Relative head length is constant through the fl exion stage, 

and then it increases in postfl exion larvae. Relative snout 

length increases and relative eye diameter decreases 

throughout larval development. Snout to anus length is 

long (49.4–50.4% SL) and remains relatively constant 

throughout development. 

Pigmentation (Fig. 21). Larvae of L. maculatus are 

lightly pigmented. The head is unpigmented through the 

fl exion stage. Prefl exion larvae have a row of melanophores (about 5–9 spots) along the lateral surface of the 

gut, 2–3 melanophores on the protruding hindgut, a row 

of 34–41 PVM beginning just posterior to the anus, a 

single melanophore ventral to the base of each pectoral 

fi n, and a patch of melanophores in the caudal region 

ventral to the notochord. 

As development continues, body pigmentation increases gradually. By the early fl exion stage, a patch of 

melanophores is present at the base of the caudal rays. 

Flexion larvae also develop 2–3 small melanophores on 

the isthmus on each side of the ventral midline. 

Most pigmentation develops during the postfl exion 

and transformation stages. By 20 mm, the melanophores 

on the isthmus are more prominent. As the gut becomes 

enclosed by the body wall (about 24 mm), the number 

of melanophores on its dorsal surface increases dramatically to 15–20 by 28 mm. The number of melanophores 

on the isthmus also increases to four, and the hypural 

edge and caudal fi n rays are heavily pigmented. A series of groups of 1–5 pigment spots begins to develop 

posteriorly along the lateral midline of the body; within 

these groups, melanophores are sometimes arranged in 

a line. With development, more of these groups appear, 
39

advancing anteriorly as far forward as the posterior edge 

of the pectoral fi n. By 41 mm, larvae have developed a 

patch of melanophores at the anterior tips of the upper 

and lower jaws, and multiple short groups of melanophores are present along the entire lateral midline of the 

body; pigment is added along the margins of both jaws 

with further development. By 58 mm, a row of fi ve melanophores has developed across the lateral surface of the 

cheek and operculum, and a single melanophore forms 

posterior to the base of the pectoral fi n. By the end of the 

transformation stage (about 70 mm), the adult pigment 

pattern is present, although residual larval gut melanophores and PVM are still visible.

Meristic features (Table 20). Principal and inferior procurrent caudal-fi n rays begin ossifying at 19.4 mm before 

they are fully developed; superior procurrent caudal-fi n 

rays begin ossifying at 25.2 mm before they are fully 

developed. Principal caudal-fi n rays are developed and 

ossifi ed by 25.2 mm; procurrent caudal-fi n rays are developed and ossifi ed by 44.0 mm. With the exception of 

the pelvic fi ns, all fi n elements are developed and ossifi ed by 25.2 mm (postfl exion stage). Pelvic-fi n buds are 

present by 31.0 mm, and all elements are developed and 

ossifi ed by 54.0 mm. By 25.2 mm, all vertebral centra are 

developed with the exception of the three posterior caudal centra; however, all of them, including the urostyle, 

Figure 20

Distribution and abundance of larvae of Daubed Shanny (Lumpenus maculatus) collected by bongo net in the Chukchi and 

Bering seas and Gulf of Alaska. Isobath=1000 m.
40 Professional Paper NMFS 15 

Figure 21

Larvae of Daubed Shanny (Lumpenus maculatus): (A) preflexion stage, 11.0 mm standard length (SL), University of Washington (UW) 95036; (B) flexion stage, 17.7 mm SL, composite of 17.7- and 16.0-mm SL specimens, UW 145099 and UW 

145098; (C) postflexion stage, 20.8 mm SL, UW 145101; and (D) transformation stage, 52.8 mm SL, UW 145102. Illustrations by R. Cartwright.
41

Table 19

Body proportions of Daubed Shanny (Lumpenus maculatus) larvae. Except for standard length (SL), values 

given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± standard 

deviation, and range (in parentheses).

 Prefl exion Flexion Postfl exion

Sample size 10 13 27

Standard length (mm) 10.8±0.9 (9.5–12.1) 15.5±2.2 (13.0–18.8) 41.5±12.4 (20.0–66.2)

Head length/SL 14.1±1.3 (11.5–16.3) 14.1±0.7 (12.8–15.4) 17.7±1.8 (14.5–21.3)

Snout length/HL 16.3±2.9 (12.8–20.8)1 21.0±2.5 (16.7–24.5)2 26.2±4.4 (17.6–41.7)3

Eye diameter/HL 35.7±4.2 (30.8–41.7) 30.3±2.6 (26.1–33.3) 20.5±3.1 (15.5–29.3)

Snout to anus length/SL 50.4±1.2 (48.9–53.0) 49.4±1.7 (46.8–52.2) 50.3±1.3 (48.8–54.2)

Body depth/SL 8.3±0.6 (7.3–9.4) 7.9±0.6 (6.9–8.8) 7.5±0.7 (6.5–9.5)3

1 n=8 2 n=12 3 n=23

Table 20

Meristic counts of cleared and stained Daubed Shanny (Lumpenus maculatus) larvae and juveniles. Counts are of ossifi ed 

elements only. Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 8.1 

 8.2 

 10.5 

 11.8 

 13.5 

 13.9 

 15.5 

 17.0 

 19.4 6+6,2

 22.4 

 25.2 XVI I,35 15 5 26 40 66 4,7+6,4

 31.0 

 42.7 LIX I,36 

 44.0 LX I,37 15 6 26 39 65 39 26 40 66 7,7+7,5

 44.01 27 41 68 41 27 42 69 *

 54.0 LXI I,35 15 I,3 6 27 40 67 40 27 41 68 6,7+6,5

 72.1 LIX I,35 15 I,3 6 27 38 65 38 27 39 66 7,7+7,6

1 Paired and median-fi n elements completely developed but unossifi ed (counts XII; I,37;15; I,3). Elements of vertebral column very 

weakly ossifi ed. 

* Character damaged.

Neural spines Centra

Spines, rays
42 Professional Paper NMFS 15 

are at least partially ossifi ed. The count of 25 precaudal 

centra obtained from MacDonald (2001) is lower than 

the count previously reported for this species (26–30; 

Matarese et al., 1989). Neural and haemal spines are developed by 25.2 mm and ossifi ed by 44.0 mm. By 55.0 

mm, scale pockets begin to form. Ranges of adult meristic counts are given in Table 2. 

Species comparisons. Pigmentation patterns are not 

useful for species-level identifi cation of larvae of the genus Lumpenus; species are distinguished primarily by a 

combination of morphological and meristic differences 

that are present in older, more developed larvae. Larvae of L. maculatus have a longer gut (mean snout to 

anus length of about 50% SL) than do larvae of other 

Lumpenus species (42–47% SL). Counts of precaudal, 

caudal, and total myomeres or vertebrae (25–30+39–

43=66–72) distinguish larvae and juveniles of L. maculatus from other species of Lumpenus in most cases. 

Lumpenus maculatus has more precaudal vertebrae (25–

30) than does L. medius (21–25), fewer total vertebrae 

(66–72) than does L. sagitta (75–82), and usually fewer 

vertebrae than does L. fabricii (70–75). The number of 

caudal vertebrae is usually lower in L. maculatus (39–

43) than in all other species of Lumpenus (42–55).

Lumpenus medius, Stout Eelblenny 

(Figs. 22, 23; Tables 21, 22)

Literature. Makushok (1986), Grigor’ev (1992b), 

Mecklenburg et al. (2002), Novikov et al. (2002).

Material examined. 78 specimens (9.6–46.5 mm) from 

the Arctic Ocean, Beaufort Sea, and Gulf of Alaska (see 

Appendix for details). 

Distribution and life history. Lumpenus medius is circumarctic over muddy bottom from nearshore waters to 

a depth of 150 m (Mecklenberg et al., 2002) in the seas 

of Okhotsk and Japan; Beaufort, Chukchi, and Bering 

seas; in the Aleutian Islands as far west as Unimak Island; along shelf areas of the Gulf of Alaska to southeastern Alaska; and in the Arctic and North Atlantic oceans. 

It is typically found at depths <100 m (Mecklenburg et 

al., 2002), and in the Northeast Atlantic is commonly 

found in areas where temperatures are <0°C (Makushok, 

1986). Adults can grow to 180 mm TL (Mecklenburg 

et al., 2002), although there is one record of an adult 

that was 300 mm TL (Novikov et al., 2002). Little is 

known of the life history of this species. Eggs are probably demersal and spawned in an adhesive mass; larvae 

are pelagic.

Larval distribution (Fig. 22). Only two larvae of Stout 

Eelblenny have been found during standard AFSC ichthyoplankton surveys; most larvae (72) were collected in 

1978–79 during special studies in which sampling was 

conducted inshore and in bays in the Gulf of Alaska. 

Larvae 9.6–46.5 mm in length are found in March–August at depths of 30–130 m in the Gulf of Alaska near 

Unimak Island and close to shore along the southeast 

side of Kodiak Island. Larvae 32.0–40.0 mm in length 

are found in August–September at depths of 4–57 m in 

the Arctic Ocean and the Beaufort Sea off the coast of 

Canada. No larvae have been collected in the Bering Sea 

in Alaskan waters. Larvae have been collected in June in 

the Sea of Okhotsk near the Kamchatka Peninsula and in 

the Bering Sea near the Russian coast (Grigor’ev, 1992b). 

Morphology (Table 21). Notochord fl exion in L. medius begins at 10.7–12.1 mm. Our largest fl exion specimen was 14.4 mm, and the smallest postfl exion larva 

was 31.0 mm, so it is unknown when notochord fl exion 

is complete. Postfl exion larvae from our material were 

31.0–46.5 mm, and transformation begins after 35.0 

mm. Our largest specimen (46.5 mm) was at transformation stage, and it is unknown when the juvenile stage 

begins. Larvae are slender-bodied during the prefl exion 

and postfl exion stages (mean body depth: 7.0% SL, 7.9% 

SL), but they have a moderate mean body depth during the fl exion stage (10% SL). Relative head and snout 

length increase throughout larval development. Relative 

eye diameter and snout to anus length initially increase, 

and then they decrease in postfl exion larvae. 

Pigmentation (Fig. 23). Larvae of L. medius are lightly pigmented. The head is unpigmented throughout the 

early stages of development. Prefl exion larvae have a row 

of melanophores (about fi ve spots) along the dorsolateral surface of the gut, one melanophore on the hindgut, 

a melanophore near the anterior ventral midline of the 

gut, 27–35 PVM beginning two myomeres posterior to 

the anus, and a single melanophore ventral to the base 

of each pectoral fi n. There is a patch of melanophores 

in the caudal region ventral to the notochord that, with 

development, is positioned at the base of the developing 

caudal-fi n rays in late prefl exion larvae. 

Body pigmentation increases little during the fl exion 

stage. A few melanophores form along the margin of the 

lower jaw. Melanophores on the dorsal surface of the gut 

are smaller. The caudal-fi n pigment extends along the margin of both developing hypural plates, with a few melanophores on the base of the caudal-fi n rays. 

Postfl exion larvae have a single melanophore on the 

opercle and a short row of pigment on the isthmus. Preanal pigment is limited to the melanophore below the 

pectoral-fi n base and lateral spots along the gut; 1–2 

spots are on the hindgut. The PVM (now 29–38) originate several myomeres posterior to the anus and are associated with the anal-fi n pterygiophores. Pigment at the 

hypural edge extends onto the caudal-fi n rays. 
43

Diffuse patches of melanophores are present on the 

snout, crown, area between the maxilla and eye, ventrolateral surfaces of the head, and medial area of the opercle during the juvenile stage; very little ventral pigment is 

present on the head. A single melanophore is present on 

the pectoral-fi n base, and another diffuse patch is present 

above the insertion of the pelvic fi n. Approximately eight 

regularly spaced melanophore patches are present along 

the dorsal midline, with some that extend slightly onto 

the dorsal fi n. The lateral body is covered with widely 

spaced, irregularly shaped pigment patches of variable 

size. A row of fi ve or more regularly spaced embedded 

melanophores is present above the notochord on the caudal peduncle. Small patches of pigment are present on 

the caudal fi n near its base. 

Meristic features (Table 22). Nearly all of the larvae of 

L. medius available to clear and stain were in the postfl exion stage. All fi n elements, vertebral centra, and neural and haemal spines are developed and ossifi ed by 33.0 

mm. The count of LXIV dorsal-fi n spines observed in one 

of the 40.0-mm specimens is greater than the count previously reported for this species (LVIII–LXIII; Matarese 

et al., 1989). The counts of 21 precaudal centra and 42 

Figure 22

Distribution of larvae of Stout Eelblenny (Lumpenus medius) collected by bongo, Tucker, and sled gear in the Arctic Ocean, 

Beaufort Sea, and Gulf of Alaska. Each symbol may indicate more than one occurrence. Isobath=1000 m.
44 Professional Paper NMFS 15 

Figure 23

Larvae of Stout Eelblenny (Lumpenus medius): (A) preflexion stage, 12.0 mm standard length (SL), University of Washington (UW) 33080; (B) flexion stage, 13.1 mm SL, UW 33044; (C) postflexion stage, 31.0 mm SL, UW 145885; and (D) 

transformation stage, 40.0 mm SL, UW 33064. Illustrations by A. Overdick.

caudal centra obtained from MacDonald (2001) are lower than the counts previously reported for this species 

(23–25+43–50; Matarese et al., 1989). Small, early scale 

pockets were present on a 44.0-mm specimen that was 

not cleared and stained. Ranges of adult meristic counts 

are given in Table 2.

Species comparisons. Pigmentation patterns are not 

useful for species-level identifi cation of larvae of the 

genus Lumpenus; species are distinguished primarily 

by a combination of morphological and meristic differences that are present in older, more developed larvae. 

Prefl exion-stage larvae of L. medius, compared to larvae of L. fabricii and L. maculatus, have a shorter gut 

(mean snout to anus length: 43.6% SL vs. 50.4% SL 

for both L. fabricii and L. maculatus) and more slender 

body (mean body depth: 7.0% SL vs. 8.3% SL for both 

of these other species). Flexion-stage larvae of L. medius
45

have a longer gut (mean snout to anus length 47.2% SL) 

than do larvae of L. sagitta (42.5% SL), and postfl exionstage larvae of L. medius have a shorter gut (mean snout 

to anus length 44.0% SL) than do larvae of L. maculatus

(50.3% SL). During the fl exion and postfl exion stages, L. 

medius is deeper bodied (mean body depth: 10.0% SL, 

7.9% SL) than all other species of Lumpenus. Counts 

of precaudal, caudal, and total myomeres or vertebrae 

(21–25+42–50=65–70) distinguish larvae and juveniles 

of L. medius from other species of Lumpenus. Lumpenus

medius usually has fewer precaudal vertebrae (21–25) 

than do L. maculatus (25–30), L. fabricii (23–28), and 

L. sagitta (26–29) and usually has fewer total vertebrae 

(65–70) than do L. fabricii (70–75) and L. sagitta (75–

82). Transformation in L. medius begins at a smaller 

size (>35 mm) than it does in L. fabricii (>45 mm). In 

transformation-stage and early juvenile L. medius, the 

dorsal and anal fi ns are connected to the caudal fi n, but 

this connection is not observed in L. fabricii (Mecklenburg et al., 2002).

Lumpenus sagitta, Snake Prickleback 

(Figs. 24, 25; Tables 23, 24)

Literature. Blackburn (1973), Hart (1973), Shiogaki 

and Sasaki (1988), Mecklenburg et al. (2002).

Table 21

Body proportions of Stout Eelblenny (Lumpenus medius) larvae. Except for standard length (SL), values 

given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± standard 

deviation, and range (in parentheses).

 Prefl exion Flexion Postfl exion

Sample size 24 13 33

Standard length (mm) 11.1±0.7 (9.6–12.1) 12.3±1.1 (10.7–14.4) 38.6±4.3 (31.0–46.5)

Head length/SL 12.6±1.7 (10.7–18.8) 13.0±1.0 (11.9–15.8) 17.2±1.6 (13.6–19.9)

Snout length/HL 20.4±4.2 (15.2–29.6)1 20.9±5.1 (15.9–27.9)2 27.3±3.9 (18.9–37.9)3

Eye diameter/HL 32.7±4.6 (26.1–42.5)4 35.7±5.1 (26.8–41.7) 18.9±1.6 (16.9–23.7)

Snout to anus length/SL 43.6±1.8 (40.6–46.9) 47.2±2.1 (42.1–49.2) 44.0±1.0 (42.4–46.6)

Body depth/SL 7.0±0.8 (5.7–8.7)5 10.0±2.3 (6.2–12.7)6 7.9±0.9 (6.6–9.8)7

1 n=18 5 n=22 2 n=4 6 n=9 3 n=26 7 n=25 4 n=23

Table 22

Meristic counts of cleared and stained Stout Eelblenny (Lumpenus medius) larvae. Counts are of ossifi ed elements only. 

Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 11.2

 11.8

 33.0 LVIII I,421 13 I,3 6 22 42 64 42 22 43 65 5,6+7,4

 34.0 LX I,401 13 I,3 6 23 42 65 41 23 43 66 4,6+7,3

 35.0 LXII I,401 14 I,3 6 24 44 68 44 24 45 69 4,6+7,2

 40.0 LXIV I,421 15 I,3 6 23 44 67 44 23 45 68 5,7+7,3

 40.0 LXII I,411 14 I,3 6 23 44 67 44 23 45 68 4,7+6,2

1 Two rays on posteriormost anal-fi n pterygiophore.

Neural spines Centra

Spines, rays
46 Professional Paper NMFS 15 

Material examined. 93 specimens (5.8–53.0 mm) from 

the Bering Sea, Gulf of Alaska, and Puget Sound, Washington (see Appendix for details). 

Distribution and life history. Lumpenus sagitta lives 

in demersal habitats ranging from shallow bays to offshore waters 435 m deep (but almost always <200 m) 

in the seas of Okhotsk and Japan, over the shelf from 

Norton Sound in the Bering Sea and along the Aleutian 

Islands, to Humboldt Bay in northern California. Adults 

can reach 510 mm TL but are usually less than 250 mm 

TL (Mecklenburg et al., 2002). Little is known of its life 

history. Eggs are probably demersal and spawned in an 

adhesive mass; larvae are pelagic. Larvae of Snake Prickleback have been collected in February–April in Puget 

Sound, Washington (Blackburn, 1973); larvae 5–52 mm 

in length are abundant near the surface in April and May 

off the outlet of the Fraser River in British Columbia 

(Hart, 1973). 

Larval distribution (Fig. 24). Larvae of Snake Prickleback of lengths 6.7–56.1 mm (mostly 17–21 mm) are 

collected in March–June, with the highest abundances 

observed in April–May, in the Bering Sea along the shelf 

Figure 24

Distribution and abundance of larvae of Snake Prickleback (Lumpenus sagitta) collected in the Bering Sea, Gulf of Alaska, 

and Puget Sound, Washington. Squares indicate collections made by bongo net; black circles indicate additional nonquantitative collections (presence only) and may represent more than one occurrence. Isobath=1000 m.
47

of the Alaska Peninsula, in Unimak Pass, on the shelf in 

the northern Gulf of Alaska, Shelikof Strait, and around 

Kodiak Island to just east of Prince William Sound. 

Larvae are frequently collected in and around Shelikof 

Strait. Neither adults nor larvae have been collected during AFSC surveys off the west coast of the United States, 

but larvae have been collected in February–April in Puget 

Sound, Washington (this study; Blackburn, 1973). Larvae 

(5–52 mm) are abundant near the surface in April and 

May off the outlet of the Fraser river in British Columbia 

(Hart, 1973). 

Morphology (Table 23). Notochord fl exion in L. sagitta begins at 15.5 mm and is complete by 25.5 mm. Postfl exion larvae are 26.0–53.0 mm. Length at the beginning of transformation is 42–44 mm; the juvenile stage 

begins at >53.0 mm. Larvae are slender-bodied and become more so with development, with mean body depth 

Figure 25

Larvae of Snake Prickleback (Lumpenus sagitta): (A) yolk-sac stage, 13.0 mm standard length (SL), University of Washington (UW) 95112; (B) flexion stage, 17.3 mm SL, UW 37176; (C) postflexion stage, 35.1 mm SL, UW 70841; and (D) 

transformation stage, 42.6 mm total length (TL). Illustrations (A–C) by B. Vinter and (D) from Shiogaki and Sasaki (1988). 
48 Professional Paper NMFS 15 

Table 23

Body proportions of Snake Prickleback (Lumpenus sagitta) larvae. Except for standard length (SL), values 

given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± standard 

deviation, and range (in parentheses). 

 Prefl exion Flexion Postfl exion

Sample size 17 21 32

Standard length (mm) 13.3±1.7 (9.2–15.4) 19.0±3.4 (15.5–25.5) 37.5±8.5 (26.0–53.0)

Head length/SL 12.4±1.6 (9.8–16.2) 11.9±1.0 (9.9–13.4) 15.1±1.7 (11.7–17.5)

Snout length/HL 20.0±4.9 (13.0–32.3)1 20.1±4.6 (10.2–26.8)2 24.7±2.6 (17.1–29.4)3

Eye diameter/HL 35.5±5.8 (27.3–47.7) 36.1±5.2 (28.1–43.8)4 21.2±4.7 (15.0–30.3)

Snout to anus length/SL 44.4±3.2 (39.9–49.1) 42.5±1.6 (40.5–45.8) 43.8±1.4 (41.5–46.7)

Body depth/SL 7.7±0.7 (6.4–9.3) 6.4±0.5 (5.1–7.1)5 6.4±0.4 (5.7–7.0)6

1 n=15 4 n=20 2 n=16 5 n=18 3 n=28 6 n=29

Table 24

Meristic counts of cleared and stained Snake Prickleback (Lumpenus sagitta) larvae. Counts are of ossifi ed elements only. 

Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 5.8 

 9.1 

 11.7 

 12.1 

 12.9 2 

 14.9 2 

 15.3 4 

 15.5 

 15.9 

 16.1 

 17.9 3 

 18.7 13 5+6

 19.0 

 19.5 271 151 42 6+7

 22.0 261 551 81 6+8

 25.0 16 4 271 531 80 6+8,2

 25.5 20 16 4 281 521 80 2,6+8,1

 27.0 

 29.5 27 51 78 51 271 521 78 3,6+8,4

 31.2 LX2 I,49 16 I,3 6 27 52 79 52 271 531 80 6,6+8,5

 34.3 LXIV2 I,47 16 I,3 6 27 50 76 50 5,6+8,4

 40.0 XLI2 I,51 17 I,3 6 27 54 81 54 271 551 82 6,6+8,5

 42.5 LXXI I,48 16 I,3 6 28 51 79 51 281 521 80 7,6+8,5

 46.0 LXX I,50 16 I,3 6 26 52 78 52 261 531 79 7,6+8,4

1 Count estimated, fi rst haemal spine not distinguishable. 2 Dorsal-fi n spines not completely developed or ossifi ed.

Neural spines Centra

Spines, rays
49

declining from 7.7% SL during the prefl exion stage to 

6.4% SL in later stages. Relative head length initially 

decreases, and then it increases in postfl exion larvae. 

Relative snout length is constant through the fl exion 

stage but increases in the postfl exion stage. Relative eye 

diameter increases slightly from the prefl exion stage to 

the fl exion stage, and then it decreases in postfl exion 

larvae. Relative snout to anus length decreases from the 

prefl exion stage to the fl exion stage and then increases 

slightly in postfl exion larvae.

Pigmentation (Fig. 25). Larvae of L. sagitta are lightly 

pigmented. The head is unpigmented in larvae prior to 

transformation. A single melanophore appears ventral to 

the pectoral-fi n base in the prefl exion stage. About 5–6 

melanophores are spaced irregularly over the dorsolateral surface of the gut and hindgut. A PVM series (≤40), 

sometimes irregularly spaced, extends from about the 

anus to the caudal peduncle. Caudal pigment is restricted 

to several small melanophores in the hypural area. 

The melanophore below the pectoral-fi n base becomes 

larger and more prominent in the fl exion stage. Up to 10 

(usually 5–8 + 2 on hindgut) melanophores appear along 

the dorsolateral surface of the gut. The number of PVM 

increases to 44–53, with about one melanophore per 

myoseptum in late fl exion. Several spots appear in the 

hypural and caudal fi nfold areas and at the base of the 

developing caudal-fi n rays. 

Pigmentation changes are minimal in postfl exion larvae. Preanal pigment is limited to the large melanophore 

below the pectoral-fi n base and the dorsolateral spots 

on the gut. The PVM series begins several myomeres 

posterior to the anus. Pigment remains at the hypural 

edge and on the base of the caudal-fi n rays.

During transformation, melanophores are present 

on the snout, the area posterior to the orbit, and the 

medial area of the opercle. A single, large melanophore 

is present at the edge of the gill cover, and another is 

present above the insertion of the pelvic fi n. Approximately six regularly spaced clusters of melanophores 

are present along the dorsal midline. The mediolateral 

body is covered with regularly spaced, dash-like clusters 

of melanophores above the gut, with patches of variable size positioned posteriorly along the lateral line. 

Additional small, regularly spaced clusters or series of 

melanophores are present between the lateral and dorsal 

midline. A loose aggregation or cluster of melanophores 

is present at the caudal-fi n base and hypural region with 

a row of melanophores that radiate onto the ventral 

caudal fi n. 

Meristic features (Table 24). Principal caudal-fi n rays 

are present by 12.1 mm and developed by 20.0 mm. Ossifi cation begins before all principal caudal-fi n rays are 

developed; some of these rays are weakly ossifi ed before 

19.0 mm. The count of 8 inferior principal caudal-fi n 

rays found in most of the cleared and stained specimens 

is greater than the count previously reported for this 

species (7; Watson, 1996). Procurrent caudal-fi n rays are 

present by 19.0 mm and are developed and ossifi ed by 

31.2 mm. The count of 5 superior procurrent caudalfin rays observed in the 34.3-mm specimen is lower 

than the count previously reported for this species (6–7; 

Watson, 1996). Anal-fi n elements are present after 22.0 

mm, begin to ossify by 25.5 mm, and are developed and 

ossifi ed by 31.2 mm. Dorsal-fi n spines are present by 

25.0 mm and begin to ossify by 31.2 mm; development 

and ossifi cation are complete by 40.0 mm. The anteriormost dorsal-fi n spines were only partially developed 

and weakly ossifi ed in the largest larva examined (46.0 

mm). Pectoral-fi n rays are present by 18.0 mm and can 

begin to ossify by 19.0 mm, but they more typically are 

developed and ossifi ed by 25.0 mm. The pelvic-fi n elements are fi rst present as fl eshy buds at 27.0 mm and 

are developed and ossifi ed by 31.2 mm. Vertebral centra 

are present by 19.0 mm and begin to ossify at around 

19.5 mm before development is complete. By 22.0 mm, 

all vertebral centra were developed and ossifi ed in most 

specimens. The count of 55 caudal centra observed 

in the 22.0- and 40.0-mm specimens is greater than 

the count previously reported for this species (46–54; 

Watson, 1996). Neural and haemal spines are ossifi ed 

by 29.5 mm. Ranges of adult meristic counts are given 

in Table 2.

Species comparisons. Species of Lumpenus can be distinguished primarily by a combination of morphological 

and meristic differences that are present in older, more 

developed larvae. Mean snout to anus length is shorter 

for larval L. sagitta (44.4% SL, 42.5% SL, and 43.8% 

SL at prefl exion, fl exion, and postfl exion stages) than 

for L. fabricii during prefl exion stage (50.4% SL) and 

for L. maculatus at all stages (50.4% SL, 49.4% SL, 

and 50.3% SL at prefl exion, fl exion, and postfl exion 

stages). Larval L. sagitta are more slender (mean body 

depth 7.7% SL) than are larvae of L. fabricii (8.3% SL) 

during the prefl exion stage, L. medius during the fl exion 

and postfl exion stages (mean body depth: 6.4% SL vs. 

10.0–7.9% SL), and L. maculatus at all stages (mean 

body depth: 7.7–6.4% SL vs. 8.3–7.5% SL). Counts 

of precaudal, caudal, and total myomeres or vertebrae 

(26–29+46–55=75–82) can in most cases distinguish 

larvae and juveniles of L. sagitta from other species of 

Lumpenus. Larvae of L. sagitta have more precaudal 

vertebrae (26–29) than do larvae of L. medius (21–25), 

more total vertebrae (75–82) than do larvae of L. maculatus (66–72) and L. medius (65–70), and usually more 

total vertebrae than do larvae of L. fabricii (70–75). 

In addition, larvae of L. sagitta have more PVM during postfl exion (44–53, usually ≥50) than do larvae of 

other species of Lumpenus (L. fabricii 33–37, L. medius

29–38, L. maculatus 34–41). 
50 Professional Paper NMFS 15 

Figure 26

Distribution and abundance of larvae of Whitebarred Prickleback (Poroclinus rothrocki) collected in the Bering Sea, Gulf 

of Alaska, and North Pacific Ocean off the coasts of British Columbia, Canada, and Washington and Oregon. Squares indicate collections made by bongo net; black circle indicates additional nonquantitative collection (presence only) and may 

represent more than one occurrence. Isobath=1000 m.

Poroclinus rothrocki, Whitebarred Prickleback 

(Figs. 26, 27; Tables 25, 26)

Literature. Miller and Lea (1972), Eschmeyer et al. 

(1983), Mecklenburg et al. (2002).

Material examined. 70 specimens (5.5–48.0 mm) from 

the Bering Sea, Gulf of Alaska, the coast of British Columbia, Canada, and the Washington and Oregon coasts 

(see Appendix for details). 

Distribution and life history. Poroclinus rothrocki is 

commonly found at depths of 46–128 m (Mecklenberg 

et al., 2002), but it has been found as deep as 420 m 

(Miller and Lea, 1972) in the southeastern Bering Sea 

and Aleutian Islands to southern California. Maximum 

length is 250 mm TL (Eschmeyer et al., 1983). Little is 

known of its life history. Eggs are probably demersal and 

spawned in an adhesive mass; larvae are pelagic.

Larval distribution (Fig. 26). Larvae of Whitebarred 

Prickleback are the most abundant of any stichaeid in 
51

our study area and are distributed throughout the outer 

shelf regions from the Bering Sea and Unimak Pass areas to Shelikof Strait, around Kodiak Island eastward 

to Prince William Sound, and along the western coast 

of North America off British Columbia, Washington, 

and Oregon. Larvae (6.1–30.2 mm) are found in late 

March–July, with the highest abundances observed in 

May and June.

Morphology (Tables 25, 26). Notochord fl exion in P. 

rothrocki begins at 9.6–10.8 mm and is complete by 

20.0 mm. Postfl exion larvae are 19.9–28.1 mm. Transformation can begin by 28 mm, and the juvenile stage 

begins by 38 mm. Larvae are slender-bodied with a 

mean body depth of 9.5% SL in the prefl exion stage, 

decreasing to 6.3% SL in the fl exion stage and increasing to 8.5% SL in the postfl exion stage. Relative head 

length remains constant through development, but relative snout length fi rst decreases then increases during 

development. Relative eye diameter and snout to anus 

length decrease with development.

Pigmentation (Fig. 27). Larvae of P. rothrocki are 

moderately pigmented. Among stichaeids, they have a 

unique caudal pigment pattern. The head is unpigmented through the early postflexion stage (<25 mm). In 

prefl exion larvae, the gut is distinctly pigmented laterally with a large spot just below the pectoral fi n. Continuing along the gut posteriorly from this spot, 2–3 

large melanophores are present on the dorsal surface, 

followed by a break and then 3–4 more large melanophores along the hindgut. Prefl exion larvae have a series of large PVM (about 38–39) beginning 3–4 myomeres posterior to the anus. Distinct from the PVM 

are about four larger melanophores in the area of the 

caudal peduncle just below the notochord. Larvae usually have about 4–12 dorsal midline melanophores that 

begin at about the posteriormost 15 myomeres. 

Late prefl exion larvae develop a spot on the isthmus, and the large melanophore below the pectoral fi n 

spreads onto the fi n base. The anteriormost dorsal midline melanophores become embedded.

By the fl exion stage, larvae develop the distinctive 

postanal melanophore pattern that easily distinguishes them from most other stichaeids. PVM increase in 

number to about 40–41 spots that remain through the 

postfl exion stage. Although still visibly larger, the posteriormost spots below the notochord in the peduncle 

area merge with the PVM. The dorsal midline melanophores migrate internally to just above the notochord 

and decrease in number from a maximum of 12 in prefl exion larvae to about 5–6 in fl exion larvae.

During the postfl exion stage, larvae ≤25 mm have 

several small melanophores in the isthmus area. A few 

internal melanophores are visible in the area of the 

hindbrain. The large melanophore at the base of the 

pectoral fi n remains but is more internal. The gut has 

about 4–6 large, variably spaced spots along its entire 

dorsal length, including the hindgut. The caudal peduncle area is distinctly pigmented with 3–5 large melanophores above and below the notochord. External pigment is present in the hypural area. 

Transformation-stage individuals have a covering of 

fi ne melanophores on the crown. Aggregations of larger melanophores on the snout and a dense patch posterior of the orbit give the appearance of a diagonally 

oriented streak passing through the eye. Additional melanophores are present in an arc-shaped patch on the 

medial area of the opercle. Five evenly spaced, irregularly shaped patches are present in an arc that extends 

from the lower jaw along the outer edge of the operculum to a point near the dorsal insertion of the pectoral 

fi n. Nine large connected patches of melanophores that 

resemble triangles or irregularly shaped diamonds are 

present on the dorsal half of the body, extending from 

just posterior of the head to the caudal peduncle. The 

apex of each patch is more heavily pigmented than the 

remainder, with some melanophores extending onto the 

dorsal fi n membrane. A crescent-shaped patch is present 

on the hypural region and is connected to the adjacent 

patch on the caudal peduncle. Pigmentation from this 

crescent-shaped patch extends dorsally and ventrally 

onto the caudal fi n. Irregularly spaced aggregations of 

melanophores are present along the distal margin of 

the dorsal fi n.The anal fi n is completely unpigmented. 

Several smaller, irregularly shaped and unevenly spaced 

ventrolateral patches are present from the pectoral-fi n 

base to the caudal peduncle. The patches posterior to 

the insertion of the fi rst anal-fi n spine are close to or 

touch the ventral midline of the body. The series of internal melanophores below the notochord described for 

earlier stages remains visible. These melanophores are 

small and closely spaced, extending farther anteriad 

than before because of their increased number (8–10). 

Meristic features (Table 26). Principal caudal-fi n rays 

are present between 10.0 and 12.0 mm, developed by 

18.8 mm, and ossifi ed by 24.0 mm. Procurrent caudalfi n rays are present by 22.5 mm. Ossifi cation begins at 

24.0 mm before all procurrent caudal-fi n rays are developed. All procurrent caudal-fi n rays are developed 

and ossifi ed by 26.0 mm. Anal-fi n elements are present by 17.9 mm, and dorsal-fi n spines are present by 

18.8 mm. Both are developed at 20.0 mm and ossifi ed 

by 22.5 mm. Pectoral-fi n rays are present by 12.0 mm, 

developed by 14.3 mm, and begin to ossify at 22.5 

mm; ossifi cation is complete by 24.0 mm. Pelvic-fi n elements are present at 18.8 mm. The pelvic-fi n spine is 

developed and ossifi ed by 24.0 mm, as are the rays by 

28.0 mm. Vertebral centra are present at 12.0 mm. All 
52 Professional Paper NMFS 15 

Figure 27

Larvae of Whitebarred Prickleback (Poroclinus rothrocki): (A) preflexion stage, 10.1 mm standard length (SL), University 

of Washington (UW) 145127; (B) flexion stage, 18.0 mm SL, UW 145128; (C) postflexion stage, 22.0 mm SL, UW 72252; 

and (D) transformation stage, 26.5 mm SL, UW 145891. Illustrations by B. Vinter.
53

Table 25

Body proportions of Whitebarred Prickleback (Poroclinus rothrocki) larvae. Except for standard length 

(SL), values given for each body proportion are expressed as percentage of SL or head length (HL): mean, 

± standard deviation, and range (in parentheses).

 Prefl exion Flexion Postfl exion

Sample size 9 50 31

Standard length (mm) 9.5±0.8 (8.6–10.8) 13.5±2.3 (9.6–18.4) 23.8±2.3 (19.9–28.1)

Head length/SL 16.5±1.6 (14.4–18.7)1 16.3±1.4 (12.2–21.5) 16.6±1.0 (14.4–19.0)2

Snout length/HL 23.9±3.7 (18.7–28.8)1 20.4±5.0 (9.8–29.8)3 29.0±3.4 (19.9–36.2)2

Eye diameter/HL 39.3±4.9 (29.9–46.2)1 31.0±4.9 (20.9–42.8)3 24.1±2.5 (17.5–28.6)2

Snout to anus length/SL 45.6±4.4 (42.0–55.7) 43.4±2.1 (38.8–50.3) 43.7±1.6 (39.1–47.2)4

Body depth/SL 9.5±1.0 (7.8–11.5) 6.3±0.9 (4.7–9.6) 8.5±0.6 (7.4–9.9)

1 n=8 3 n=49 2 n=30 4 n=29

Table 26

Meristic counts of cleared and stained Whitebarred Prickleback (Poroclinus rothrocki) larvae and juveniles. Counts are of 

ossifi ed elements only. Specimens between dashed lines (-----) were undergoing notochord fl exion.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 5.5 

 10.0 

 12.0 

 12.5 

 14.3 

 16.1 

 17.9 

 18.8 

 20.0 5 191 36 55 36 19 48 67 6+5

 22.5 LIX III,42 02 6 41 44 48 44 19 45 64 6+6

 24.0 LIX III,42 14 I,3 6 191 46 65 46 19 47 66 4,7+6,3

 26.0 LX III,43 14 I,3 6 191 44 63 44 19 45 64 5,7+6,4

 28.0 LIX III,42 14 I,3 6 191 45 64 45 19 46 65 5,7+6,4

 38.0 LIX III,* 14 I,3 6 181 46 64 46 18 47 65 *

 48.0 LIX III,43 14 I,3 6 191 46 65 47 19 48 67 5,7+6,4

1 All neural spines present but only four with stain noted as ossifi ed.

* Character damaged or missing.

Neural spines Centra

Spines, rays

vertebral centra and neural and haemal spines are 

developed by 16.1 mm and ossifi ed by 20.0 mm. The 

total count of 64 vertebral centra observed in the 22.5- 

and 26.0-mm specimens is lower than the total count 

previously reported for this species (65–68; Watson, 

1996). Ranges of adult meristic counts are given in 

Table 2.

Species comparisons. Larval P. rothrocki are similar 

to species of Lumpenus and Lumpenella longirostris

but can be distinguished by the irregularly spaced row 

of embedded melanophores above and below the notochord on the caudal peduncle. The break in the spacing 

of the melanophores dorsally along the gut in prefl exion and fl exion larvae is unique to P. rothrocki.
54 Professional Paper NMFS 15 

Subfamily Opisthocentrinae

Lumpenopsis hypochroma, Y-prickleback 

See “Other material” section of this guide.

Opisthocentrus ocellatus, Ocellated Blenny 

(Fig. 28, Table 27)

Literature. Tokuya and Amaoka (1980), Shiogaki 

(1981, 1982), Mecklenburg et al. (2002). 

Material examined. Seven specimens (13.6–15.8 mm) 

from the Sea of Japan (see Appendix for details).

Distribution and life history. Opisthocentrus ocellatus 

is found in the Sea of Okhotsk off southeastern Kamchatka, the Sea of Japan off North Korea, and the Kuril 

and Commander Islands; it has not been reported from 

North America. Ocellated Blenny has been collected at 

depths >300 m off Hokkaido, but this species is usually 

found among seaweed in shallow coastal areas <70 m 

deep (Mecklenburg et al., 2002). Adults can grow to 200 

mm TL (Mecklenburg et al., 2002) and reach maturity at 

1 year (Shiogaki, 1982). Adhesive masses of 700–3300 

eggs are spawned in narrow cavities under stones on 

muddy sand bottoms and guarded by the female. Eggs 

are 1.9–2.0 mm in diameter with one large and many 

small, yellow oil globules. Eggs hatch after 48 days at 

5–10°C; larvae hatch at 9–10 mm (Shiogaki, 1982).

Larval distribution. Larvae of Ocellated Blenny have 

not been collected from our study area. Five prefl exionstage and two fl exion-stage larvae (13.6–15.8 mm) collected at one location in April off Usujiri, southwest 

Hokkaido, Japan, were available for examination (see 

Appendix). 

Morphology (Table 27). Notochord fl exion in O. ocellatus begins at 14.4–15.8 mm, and it is unknown from 

available material when it is complete. As illustrated by 

Tokuya and Amaoka (1980), transformation appears 

to begin at about 19.5 mm before notochord fl exion is 

complete; the juvenile stage appears to begin before 22.0 

mm. Larvae are initially slender (mean body depth 9.0% 

SL), becoming moderately deep-bodied (11.8% SL) in 

the fl exion stage. Relative head and snout lengths and 

snout to anus lengths all increase through the fl exion 

stage, while relative eye diameter decreases.

Pigmentation (Fig. 28). Pigment description is based 

in part on Shiogaki (1982) and Tokuya and Amaoka 

(1980). Larvae of O. ocellatus are lightly pigmented. The 

head is unpigmented in prefl exion larvae. Initially, the 

nape region has several spots. A single spot occurs on the 

cleithrum. About six stellate melanophores are arrayed 

dorsolaterally along the gut, and several melanophores 

occur dorsally on the hindgut. A ventral series of evenly 

spaced melanophores extends along the midline of the 

gut from the cleithral pigment and continues to the anus. 

A PVM series (<40) of one melanophore per myoseptum 

extends from the second postanal myomere to the caudal 

peduncle. A melanophore occurs in the area where the 

hypural bones are forming.

The pigment pattern changes little in fl exion-stage larvae. Several spots are added in the opercular area, and a 

few PVM extend into the hypaxial area. 

In juveniles, melanophores completely cover the 

snout, crown, nape, and lateral body. Two darker streaks 

radiate posteriorly outward from the eye, with one projecting towards the crown and the other projecting ventrally onto the opercular region. In general, the opercular 

region is lightly pigmented, with the exception of this 

streak and some irregularly spaced melanophores scattered between the streak and heavier pigmentation that 

covers the crown and nape. Another dark streak of pigment that resembles a vertically projecting arc is present 

below the eye; an arc-shaped row of single melanophores 

connects it with the streak on the operculum. A triangular-shaped patch of melanophores with the tip pointing ventrally is present on the pectoral-fi n blade that is 

otherwise unpigmented. The lateral and ventral surfaces 

of the gut also are unpigmented. Numerous irregularly 

spaced areas on the lateral body, where the melanophore 

covering is slightly less dense, give the appearance of 

lighter spots. The base of the dorsal fi n is covered with a 

continuous line of dark pigment with four evenly spaced 

spots. The continuous line of dorsal pigmentation becomes increasingly thicker posteriorly over the caudal 

peduncle, covering from one-half to more than threefourths of the dorsal fi n. The anal fi n below the caudal 

peduncle is similarly pigmented but to a lesser extent. A 

large blotch extends posteriorly from the hypurals and 

covers about two-thirds of the caudal fi n surface. 

Meristic features. In the single cleared and stained prefl exion larva (12.5 mm), pectoral-fi n rays were present, 

but no development of other fi n elements was observed. 

Neural and haemal spines were developed but unossifi ed, 

and vertebral centra were present. Hypurals were present, but no caudal-fi n rays were present. All paired and 

median fi ns, except for the caudal fi n, are developed in 

the 22.1-mm juvenile illustrated by Tokuya and Amaoka 

(1980). Pelvic fi ns are absent. All fi ns are developed in 

the 33.0-mm juvenile illustrated by Tokuya and Amaoka 

(1980). Ranges of adult meristic counts are given in 

Table 2. 

Species comparisons. Larvae of O. ocellatus are similar 

to most species in the subfamily Xiphisterinae, particularly species of Anoplarchus, but can be distinguished by 

the evenly spaced row of large stellate melanophores that 
55

Figure 28

Larvae of Ocellated Blenny (Opisthocentrus ocellatus): (A) preflexion stage, 9.4 mm total length (TL); (B) late preflexion 

stage, 12.7 mm TL; (C) flexion stage, 15.0 mm standard length (SL), composite of 14.4- and 15.2-mm TL specimens, Hokkaido University Museum of Zoology (HUMZ-L) 02671; (D) flexion stage, 19.5 mm body length (BL); and (E) juvenile, 

33.0 mm TL. Illustrations (A–B, E) from Shiogaki (1982, A–B reared), (C) by A. Overdick, and (D) from Tokuya and 

Amaoka (1980).
56 Professional Paper NMFS 15 

occur ventrally along the entire length of the gut; in species of Anoplarchus, a line or row of small punctate melanophores extends only over the anterior two-thirds to 

three-fourths of its length. In addition, O. ocellatus has 

a longer mean snout to anus length (48.2–53.2% SL vs. 

41.1–43.2% SL) and more precaudal vertebrae (21–23) 

than species of Anoplarchus (17–19). These comparisons were made on the basis of examination of very few 

specimens.

Plectobranchus evides, Bluebarred Prickleback 

(Figs. 29, 30; Tables 28, 29)

Literature. Hart (1973), Eschmeyer et al. (1983), 

Matarese et al. (1989), Watson (1996), Mecklenburg and 

Sheiko (2004).

Material examined. 17 specimens (7.0–49.0 mm) from 

the coast of Oregon (see Appendix for details). 

Distribution and life history. Plectobranchus evides is 

found over mud, gravel, or sand bottoms 84–368 m deep 

(Eschmeyer et al., 1983; Mecklenberg and Sheiko, 2004) 

from central British Columbia to southern California. 

Adults can grow to 132 mm TL (Hart, 1973). Little is 

known of the early life history. Eggs are probably demersal and spawned in an adhesive mass; larvae are pelagic.

Larval distribution (Fig. 29). Larvae of Bluebarred 

Prickleback have not been collected dur ing AFSC ichthyoplankton surveys. Larvae (4.7–49.0 mm) are collected off the northern and central coast of Oregon during March–May and July. Larvae are found in winter 

and spring, primarily in February–April, during CalCOFI 

surveys (Watson, 1996).

Morphology (Table 28). Notochord flexion in P. 

evides begins after 11.0 mm (only one fl exion larva was 

available for examination) and is complete before 20.0 

mm. Postfl exion larvae are 20.0–50.0 mm. Transformation begins after 50.0 mm, and length at the beginning 

of the juvenile stage is unknown. Larvae are slender 

throughout development (mean body depth: 8.9% SL, 

7.9% SL, 9.5% SL, in the prefl exion–postfl exion stages). 

Relative head length increases throughout larval development. Relative snout to anus length decreases slightly 

from the prefl exion stage to the postfl exion stage. Relative snout length and eye diameter decrease during larval 

development. 

Pigmentation (Fig. 30). Larvae of P. evides are moderately pigmented. Among stichaeids, they have a unique 

pigment pattern composed of patches along the postanal 

body. The head is unpigmented through the early postfl exion stage (<27 mm). In prefl exion-stage larvae, three 

patches are present anterior to the anus: the fi rst patch is 

on the lateral surface of the gut near the base of the pectoral fi n, the second is above the dorsal margin of the gut 

midway between the pectoral fi n and anus, and the third 

patch is composed of melanophores on the hindgut and 

above the dorsal margin of the hindgut. Postanal pigment consists of three patches: the anteriormost is ventrolateral, the next is a lateral bar at about 85% SL, and 

the posterior patch encircles the tail with melanophores 

above and below the notochord tip as well as in the caudal peduncle and fi nfold area. 

During the fl exion stage, the melanophores above 

the gut increase in size and develop internally. Along 

the postanal body, the middle patch becomes primarily 

mediolateral and the melanophores in the posteriormost 

patch become larger but fewer in number. Mediolateral 

spots later appear in the caudal peduncle area, in addition to melanophores around the notochord tip, along 

the hypural edge, and extending onto the developing 

caudal-fi n rays. 

Table 27

Body proportions of Ocellated Blenny (Opisthocentrus ocellatus) larvae. Except for standard length (SL), 

values given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± 

standard deviation, and range (in parentheses). 

 Prefl exion Flexion Postfl exion

Sample size 5 2 

Standard length (mm) 14.5±0.9 (13.6–15.8) 14.8±0.6 (14.4–15.2) 

Head length/SL 14.6±0.9 (13.4–15.4) 19.8±0.3 (19.6–19.9) 

Snout length/HL 24.8±5.3 (18.3–32.4) 28.0±1.6 (26.9–29.1) 

Eye diameter/HL 35.7±4.1 (31.4–42.1) 28.1±0.3 (27.8–28.3) 

Snout to anus length/SL 48.2±1.1 (47.1–49.5) 53.2±1.6 (52.0–54.3) 

Body depth/SL 9.0±0.6 (8.6–10.0) 11.8±0.4 (11.5–12.1) 
57

Figure 29

Distribution of larvae of Bluebarred Prickleback (Plectobranchus evides) collected in the North Pacific Ocean off the Oregon coast. Each symbol may indicate more than one occurrence. Isobath=1000 m. 

By the early postfl exion stage, some pigment develops 

externally on the snout, at the jaw angle, and posteriorly 

on the mid- and hindbrain. The pigment near the base 

of the pectoral fi n is internal and may be diffi cult to see. 

Most of the melanophores in the remaining fi ve pigment 

patches become primarily internal, but external pigment 

remains along the hindgut and in the three postanal 

patches. Mediolateral pigment in the caudal peduncle 

area increases anteriorly, and melanophores are present 

on the caudal-fi n rays along the hypural edge. 

Meristic features (Table 29). No fl exion larvae were available for clearing and staining. All fi n elements except pelvics 

are developed and ossified in postflexion larvae by 23.0 

mm. The pectoral-fi n and anal-fi n rays, however, are only 

weakly ossifi ed. The count of two inferior procurrent caudal-fi n rays observed in the 23.0- and 39.5-mm specimens 

is lower than the count previously reported for this species 

(3–4; Watson, 1996). Vertebral centra and neural and haemal spines also are developed and ossifi ed by 23.0 mm. The 

counts of 21 precaudal centra and 61 total vertebral centra observed in the 34.0-mm specimen are greater than the 

counts previously reported for this species (20+39–40=59–

60; Watson, 1996). Pelvic-fi n elements are present by 23.0 

mm and developed and ossifi ed by 39.5 mm, perhaps earlier. 

Ranges of adult meristic counts are given in Table 2.
58 Professional Paper NMFS 15 

Figure 30

Larvae of Bluebarred Prickleback (Plectobranchus evides): (A) preflexion stage, 9.2 mm standard length (SL); (B) flexion 

stage, 16.9 mm SL; and (C) postflexion stage, 31.3 mm SL. Illustrations from Matarese et al. (1989).

Table 28

 Body proportions of Bluebarred Prickleback (Plectobranchus evides) larvae. Except for standard length 

(SL), values given for each body proportion are expressed as percentage of SL or head length (HL): mean, 

± standard deviation, and range (in parentheses).

 Prefl exion Flexion Postfl exion

Sample size 3 1 8

Standard length (mm) 9.5±1.7 (7.8–11.1) 15.11 34.0±10.8 (20.0–50.0)

Head length/SL 13.7±1.9 (11.6–15.3) 20.51 22.1±4.8 (17.8–28.1)

Snout length/HL 32.9±4.9 (29.4–36.4)2 25.81 24.9±5.7 (17.6–32.4)

Eye diameter/HL 29.1±6.6 (23.5–36.4) 19.41 19.0±3.5 (14.4–25.8)

Snout to anus length/SL 47.4±1.2 (46.2–48.4) 49.71 47.1±2.6 (43.2–50.9)

Body depth/SL 8.9±0.7 (8.1–9.5) 7.91 9.5±1.7 (7.5–12.2)

1 Original length 16.9 mm; 15.1 mm after preservative changed to ethanol.

2 n=2
59

Species comparisons. Larvae of P. evides are unusual 

among stichaeids because they have fi ve lateral melanophore patches that are almost evenly spaced and extend 

from midgut to the caudal region. The posteriormost 

patch extends anteriorly onto the caudle peduncle in postfl exion larvae. Plectobranchus evides is the only species in 

the study area that lacks PVM during larval stages. 

Subfamily Xiphisterinae

Alectridium aurantiacum, Lesser Prickleback

See “Other material” section of this guide.

Alectrias spp. (Fig. 31; Tables 30, 31)

Literature. Tokuya and Amaoka (1980), Shiogaki 

(1985, 1987), Coleman (1999), Hatooka (2002), Mecklenburg et al. (2002), Mecklenburg and Sheiko (2004).

Material examined. 19 specimens (10.7–16.6 mm) 

from the Sea of Japan (see Appendix for details). 

Distribution and life history. Stone Cockscomb (Alectrias alectrolophus) is found to depths of 100 m, usually at depths <50 m, commonly intertidally, and often in 

bays with bottoms of pebble-rock debris (Mecklenberg et 

al., 2002; Mecklenberg and Sheiko, 2004) in the northern Sea of Japan; Sea of Okhotsk; Norton Sound, Alaska; the Bering Sea; off southeastern Kamchatka; and the 

Commander Islands. A reported occurrence off Amchitka 

Island in the western Aleutian Island Archipelago is unconfi rmed (Mecklenberg et al., 2002). Adults can grow 

to 128 mm TL. Adhesive eggs are laid in a mass guarded 

by the female parent (Coleman, 1999); larvae are pelagic.

The only larvae available for examination for this 

study were from Japan, where several Alectrias species 

are found (A. alectrolophus, A. benjamini, A. mutsuensis). Six of the larvae, specifi cally those with 19–20 precaudal myomeres, were A. alectrolophus, according to 

information published by Hatooka (2002), but because 

there were no discernable differences in pigment among 

the larvae examined, this section is a description of the 

genus only.

Larval distribution. Larvae of the genus Alectrias have 

not been collected in our study area. Nineteen larvae 

(10.7–16.6 mm) collected in May and June from two 

locations off Muroran and Usujiri, southwest Hokkaido, 

Japan, were available for examination (see Appendix). 

Morphology (Table 30). Notochord fl exion in species 

of Alectrias begins before 10.7 mm and is complete as 

early as 12.1 mm. Postfl exion larvae are 12.1–16.6 mm. 

Lengths at which transformation and the juvenile stage 

begin are unknown. Body depth is moderate in the fl exion stage (11.5% SL) and decreases slightly (10.7% SL) 

in the postfl exion stage. Relative head length, snout to 

anus length, and eye diameter all decrease from the fl exion stage to the postfl exion stage, while relative snout 

length increases (an observation made on the basis of 

measurements from only one fl exion specimen). 

Pigmentation (Fig. 31). A total of 2 fl exion and 17 

postfl exion larvae were available for examination. Flexion and postfl exion larvae are lightly pigmented. There 

usually are one to several spots over the brain and a 

large internal spot at the nape. A single line of internal 

pigment occurs at the cleithral symphysis in the shape 

of a “V”. There are about four internal stellate melanophores dorsally along the gut and several external 

melanophores on the dorsal surface of the hindgut. A 

ventral midline series of melanophores on the gut begins anterior to the cleithral pigment and extends to just 

anterior to the anus. A PVM series (33–39), consisting 

Table 29

Meristic counts of cleared and stained Bluebarred Prickleback (Plectobranchus evides) larvae. Counts are of ossifi ed elements 

only. Specimen above dashed line (-----) was in the prefl exion stage, and the specimens below were in the postfl exion stage.

 Branchi-

 Standard Dorsal Anal Pectoral Pelvic ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 9.5 

 23.0 LV II,36 15 5 20 39 59 39 20 40 60 4,7+7,2

 34.01 21 39 60 39 21 40 61 

 39.5 LV II,35 15 I,3 5 20 39 59 39 20 40 60 4,7+7,2

 50.0 LIV II,35 15 I,3 5 20 39 59 39 20 40 60 4,7+7,3

1 Two specimens, neither had stained fi n elements; they likely were initally preserved improperly.

Neural spines Centra

Spines, rays
60 Professional Paper NMFS 15 

Figure 31

Larvae of Alectrias spp: (A) Stone Cockscomb (Alectrias alectrolophus): flexion stage, 10.7 mm standard length 

(SL), Hokkaido University Museum of Zoology (HUMZ-L) 02756; and (B) Alectrias sp.: postflexion stage, 16.0 mm 

SL, HUMZ-L 03915. Illustrations by A. Overdick.

of a melanophore each at most myosepta, extends from 

postanal myomere 4–5 to the caudal peduncle. Internal 

melanophores are present dorsally above the notochord 

in the peduncle area. Pigment occurs along the edge of 

the hypural bones.

Few changes occur in postfl exion larvae. The dorsal 

spots on the head decrease in size, becoming less prominent. Several spots are added on the lower jaw and opercular area. Pigment is added in the isthmus area. The 

number of melanophores on the dorsal surface of the gut 

increases to about eight, and all are smaller than they 

were during the fl exion stage. Although less prominent, 

pigment remains visible along the edges of the hypural 

bones. 

Meristic features (Table 31). Principal caudal-fi n rays 

are developed by 11.7 mm and ossifi ed by 14.0 mm. 

Procurrent caudal-fi n rays begin ossifying by 14.0 mm 

before all elements are developed, and they are developed and ossifi ed by 15.8 mm. Dorsal-fi n and anal-fi n 

elements are present by 11.7 mm. Anal-fi n rays are ossifi ed by 14.0 mm, and anal-fi n spines are ossifi ed by 15.8 

mm. Dorsal-fi n spines begin to ossify by 14.0 mm before 

all are developed; dorsal-fi n spines are developed and ossifi ed by 15.8 mm. Pectoral-fi n rays are present by 11.7 

mm and ossifi ed by 14.0 mm. This genus lacks pelvic 

fi ns. Vertebral centra and neural and haemal spines are 

developed by 11.7 mm; neural and haemal spines are 

weakly ossifi ed. By 14.0 mm, all vertebral centra are ossifi ed. Both cleared and stained postfl exion specimens 

had <19 precaudal vertebrae, and they were most likely 

A. benjamini or A. mutsuensis. Ranges of adult meristic 

counts are given in Table 2.

Species comparisons. Larval Xiphisterinae are most 

similar to larvae in the subfamilies Chirolophinae and 

Lumpeninae. Larval Xiphisterinae have longer guts than 

do most Chirolophinae larvae (snout to anus length: 

41–51% SL vs. 32–37% SL) and lack dorsal midline pigment and pelvic fi ns. Xiphisterines can be distinguished 

from lumpenine larvae by their deeper bodies (body 

depth: 9.0–11.0% SL vs. 6.6–8.5% SL), typically shorter 

guts (snout to anus length: 41–43% SL vs. 43–50% SL), 

and ventral pigment on the gut (vs. absent). 

Based on limited material, larvae of Alectrias species

are similar to most species of Xiphisterinae, particularly species of Anoplarchus. Larvae of Alectrias species cannot be readily distinguished from Anoplarchus 

insignis because of overlapping meristic counts and the 

presence of head pigment and internal pigment above 

the notochord in the caudal peduncle. However, species of Alectrias can be distinguished from Anoplarchus 
61

Table 30

Body proportions of Alectrias spp. larvae. Except for standard length (SL), values given for each body 

proportion are expressed as percentage of SL or head length (HL): mean, ± standard deviation, and range 

(in parentheses).

 Prefl exion Flexion Postfl exion

Sample size 1 17

Standard length (mm) 10.7 15.0±1.2 (12.1–16.6)

Head length/SL 16.6 15.3±1.0 (13.4–17.0)

Snout length/HL 26.4 29.9±4.0 (22.9–36.4)

Eye diameter/HL 29.6 28.3±3.0 (23.4–34.9)

Snout to anus length/SL 45.9 41.2±1.0 (38.8–42.5)

Body depth/SL 11.5 10.7±0.8 (9.3–11.8)

Table 31

Meristic counts of cleared and stained Alectrias spp. larvae1. Counts are of ossifi ed elements only. Specimen between dashed 

lines (-----) was undergoing notochord fl exion. This genus lacks pelvic fi ns.

 Branchi-

 Standard Dorsal Anal Pectoral ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 11.7 16 46 62 46 

 14.0 LIX I,41 10 5 17 45 62 45 17 46 63 3,6+6,2

 15.8 LVIII I,41 09 5 16 45 61 45 16 46 62 4,6+6,2

1 Counts of 14.0- and 15.8-mm larvae match A. benjamini and A. mutsuensis.

Neural spines Centra

Spines, rays

purpurescens by the presence of pigment on the head in 

the fl exion and postfl exion stages (absent in A. purpurescens), presence of internal pigment above the notochord 

on the caudal peduncle (also absent in A. purpurescens), 

usually more caudal myomeres (44–50 vs. 40–46), and 

usually more dorsal-fi n spines and anal-fi n rays in postfl exion larvae (LV–LXVI; I,39–45 vs. LIV–LX; I–II,35–

42). Larvae of Alectrias species are slightly deeper bodied 

than are larvae of both Anoplarchus species (mean body 

depth: 11.5–10.7% SL vs. 8.9–9.9% SL) during the fl exion and postfl exion stages. 

Anoplarchus insignis, Slender Cockscomb 

(Figs. 32, 33; Tables 32, 33)

Literature. Eschmeyer et al. (1983), Watson (1996), 

Mecklenburg and Sheiko (2004).

Material examined. 178 specimens (5.2–36.5 mm) from 

the Bering Sea, Gulf of Alaska, coast of Washington, and 

Puget Sound, Washington (see Appendix for details).

Distribution and life history. Anoplarchus insignis is 

found among rocks and algae from intertidal areas to a 

depth of 30 m, most frequently in subtidal areas (Mecklenberg and Sheiko, 2004), in the southeastern Bering 

Sea, and throughout the Aleutian Islands to northern 

California in the North Pacifi c Ocean. Adults grow to 

118 mm TL (Eschmeyer et al., 1983). Little is known of 

its life history. Eggs are probably demersal and spawned 

in an adhesive mass; larvae are pelagic.

Larval distribution (Fig. 32). The third most abundant 

stichaeid in our study area, larvae of Slender Cockscomb (5.2–31.1 mm) are found in April–June (most frequently in May) in the Bering Sea, west of and around 

Unimak Pass, into the Gulf of Alaska in nearshore and 

shelf areas, around Kodiak Island to the mouth of Cook 

Inlet, off the northern coast of Washington, and in Puget 

Sound, Washington. 

Morphology (Table 32). Notochord fl exion in A. insignis begins at 8.9 mm and is complete by 14.0 mm. 
62 Professional Paper NMFS 15 

Postfl exion larvae are 14.0–20.6 mm. Transformation can 

begin between 17.0 (note 17.0-mm specimen in Fig. 33) 

and 25.0 mm, and the juvenile stage begins before 35.0 

mm. Larvae are slender-bodied with a mean body depth of 

9.2% SL in the prefl exion stage that slightly decreases to 

9.0% SL in later stages. Relative head and snout lengths 

increase and relative eye diameter decreases from the prefl exion stage to the postfl exion stage. Snout to anus length 

remains constant throughout development.

Pigmentation (Fig. 33). Larvae of A. insignis are moderately pigmented. Prefl exion larvae as small as 6.5 mm 

Figure 32

Distribution and abundance of larvae of Slender Cockscomb (Anoplarchus insignis) collected by bongo net in the Bering 

Sea, Gulf of Alaska, and Puget Sound, Washington. A fourth-root transformation was computed for the data to normalize 

the abundance. Isobath=1000 m.

may have a melanophore on the hindbrain or nape area. 

A small melanophore at the base of the cleithrum persists 

throughout development. Five or more melanophores are 

present dorsally on the gut in addition to several that 

cover the hindgut. A ventral midline row of pigment extends three-fourths of the length of the gut. Prefl exion 

larvae have about 45–55 small, closely spaced PVM. 

Several melanophores are present in the ventral caudal 

fi nfold.

During the fl exion stage, several changes in pigmentation occur. By early fl exion, the prominent spot on the 

nape becomes embedded and pigment is present on the 
63

Figure 33

Larvae of Slender Cockscomb (Anoplarchus insignis): (A) preflexion stage, 7.8 mm standard length (SL), composite of 

7.3- and 7.8-mm SL specimens, University of Washington (UW) 145894 and UW 141551; (B) flexion stage, 8.9 mm SL, 

UW 141567; (C) late flexion stage, 12.1 mm SL, UW 143894; (D) postflexion stage, 18.4 mm SL, UW 140942; and (E) 

transformation stage, 17.0 mm SL, UW 40411. Illustrations by R. Cartwright.
64 Professional Paper NMFS 15 

Table 32

Body proportions of Slender Cockscomb (Anoplarchus insignis) larvae. Except for standard length (SL), 

values given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± 

standard deviation, and range (in parentheses). 

 Prefl exion Flexion Postfl exion

Sample size 70 71 27

Standard length (mm) 6.8±0.8 (5.2–8.9) 11.3±1.7 (8.9–13.9) 16.1±2.0 (14.0–20.6)

Head length/SL 13.8±1.3 (10.6–16.8) 13.2±1.6 (9.8–17.8)1 14.6±1.0 (12.4–16.7)

Snout length/HL 18.8±5.6 (7.7–28.6)2 23.2±4.7 (11.6–34.9)1 25.1±3.0 (18.6–31.8)

Eye diameter/HL 40.7±5.6 (30.7–55.5)3 32.5±4.6 (23.6–41.6)1 27.3±3.7 (20.2–34.3)

Snout to anus length/SL 41.3±2.0 (36.9–46.2) 41.1±1.5 (38.2–44.3) 41.6±1.5 (37.7–44.6)

Body depth/SL 9.2±1.7 (6.0–13.3)4 9.0±0.7 (7.2–10.3) 9.0±0.7 (8.0–11.6)

1 n=69 2 n=53 3 n=58 4 n=67

Table 33

Meristic counts of cleared and stained Slender Cockscomb (Anoplarchus insignis) larvae and juveniles. Counts are of ossifi ed elements only. Specimens between dashed lines (-----) were undergoing notochord fl exion. This species lacks pelvic fi ns.

 Branchi-

 Standard Dorsal Anal Pectoral ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 7.2 

 7.5 

 8.3 

 9.0 

 9.4 

 10.1 

 10.9 

 11.5 

 11.7 

 12.0 

 13.1 17 44 61 44 

 13.5 

 14.0 

 14.3 

 15.1 

 15.6 17 45 62 45 

 16.4 5 18 47 65 47 18 48 66 6+5

 16.4 5 19 46 65 46 19 47 66 2,6+6,1

 17.2 5 18 47 65 47 18 48 66 6+7,1

 18.2 5 18 47 65 47 18 48 66 6+6

 18.5 LX I,44 10 5 17 47 64 47 17 48 65 4,6+6,2

 19.2 LIX I,42 10 5 17 46 63 46 17 47 64 5,6+7,2

 20.0 LXI I,44 10 5 18 46 64 46 18 47 65 5,6+7,2

 36.5 LXI I,42 10 5 18 46 64 46 18 47 65 4,6+7,2

Neural spines Centra

Spines, rays
65

isthmus. Pigment increases along the dorsal surface of 

the gut and hindgut. The PVM coalesce to about 38–39, 

one per pterygiophore, and become more slash-like in 

appearance. Five or more internal melanophores form 

dorsally on the notochord; initially light and undefi ned, 

these melanophores become associated with the posterior epaxial myomeres. Additional head pigment usually 

develops during the postfl exion stage but is not present 

in every larva. The prominent nape spot becomes embedded but remains visible. The number of PVM decreases 

further to about 36, and the internal pigment on the notochord is not always easily visible. 

During transformation and early juvenile stages, melanophores cover the entire head and body. Two dark 

streaks radiate outward posteriorly from the eye, with 

the dorsalmost streak appearing as a nearly horizontal 

bar through the eye. Another short, diagonally oriented 

vertical bar is present near the ventral margin of the eye. 

This bar appears loosely joined to another streak along 

the outer edge of the preopercle, and together they form 

a “V” shape. A prominent spot is present at the dorsal fi n 

origin, and irregularly shaped clusters of melanophores 

are seen on the membrane along the entire length of 

the dorsal fi n. Up to 10 prominent, dash-shaped spots 

are present posteriorly along the base of the dorsal fi n. 

A similar pigmentation pattern forms along the entire 

length of the anal fi n, although the clusters of melanophores are concentrated more along the fi n rays rather 

than on the fi n membrane. Dark pigment spots are present on the dorsal and ventral margins of the hypurals, 

and a large patch of small melanophores covers most of 

the caudal fi n. 

Meristic features (Table 33). Principal caudal-fi n rays 

are present by 8.3 mm, developed by 12.0 mm, and ossifi ed by 16.4 mm. Procurrent caudal-fi n rays are present 

by 12.0 mm, and ossifi cation begins at 16.4 mm before 

all are developed. All procurrent caudal-fi n rays are ossifi ed by 18.5 mm. The ranges of 1–5 superior and 1–4 inferior procurrent caudal-fi n rays observed on radiographs 

are greater than the ranges previously reported for this 

species (3–4, 2–3; Watson, 1996). Anal-fi n elements are 

present by 13.1 mm, and dorsal-fi n spines are present 

between 14.0 and 15.1 mm. All elements in both fi ns 

are ossifi ed by 18.5 mm. The count of 38 anal-fi n rays 

observed on radiographs is lower than the count previously reported for this species (40–46; Watson, 1996). 

Pectoral-fi n rays are present by 10.9 mm, developed by 

14.3 mm, and ossifi ed by 18.5 mm. Pelvic fi ns are absent. 

Vertebral centra are present by 9.0 mm, developed by 

10.9 mm, and all of them are ossifi ed by 16.4 mm. The 

count of 42 caudal centra observed on radiographs is 

lower than the count previously reported for this species 

(44–49; Watson, 1996). Neural and haemal spines are 

ossifi ed by 15.6 mm (ossifi cation occurred at 13.1 mm in 

one specimen), before vertebral centra are ossifi ed. Ranges of adult meristic counts are given in Table 2.

Species comparisons. Larvae of A. insignis are most 

likely to be confused with other Xiphisterinae, particularly their congener A. purpurescens, but they also may 

be confused with species of Alectrias and the opisthocentrine Opisthocentrus ocellatus. Anoplarchus insignis

is generally more pigmented than is A. purpurescens

throughout development. A small melanophore present 

on the nape early in development, along with additional 

head pigment in later stages, is absent in A. purpurescens. Larvae of A. insignis typically have more PVM 

(>45) during the prefl exion stage than do larvae of A. 

purpurescens (≤40), and they also have a row of melanophores above the notochord in the caudal peduncle; this 

row is absent in A. purpurescens. Larvae of A. insignis

cannot be readily distinguished from species of Alectrias

because of overlapping counts of meristic features and 

the presence of head pigment and internal pigment above 

the notochord in the caudal peduncle in both taxa. Larvae of A. insignis can be distinguished from larvae of O. 

ocellatus by a line or row of small, punctate, ventral melanophores that extends along the anterior three-fourths 

of the length of the gut rather than an evenly spaced row 

of large stellate melanophores along its entire length. In 

addition, A. insignis has a shorter snout to anus length 

(mean: 41.1–41.6% SL vs. 48.2–53.2% SL) and fewer precaudal vertebrae (17–19) than does O. ocellatus

(21–23).

Anoplarchus purpurescens, High Cockscomb 

(Figs. 34, 35; Tables 34, 35)

Literature. Schultz and DeLacy (1932), Peppar (1965), 

Hart (1973), Marliave (1975a), Wang (1986), Matarese 

et al. (1989), Watson (1996).

Material examined. 204 specimens (4.7–37.0 mm) 

from the Bering Sea; Gulf of Alaska; Puget Sound, Washington; and the coasts of British Columbia, Canada, and 

Oregon (see Appendix for details).

Distribution and life history. Anoplarchus purpurescens is found from the Bering Sea near the Pribilof Islands, throughout the Aleutian Islands, and in nearshore 

areas in the Gulf of Alaska and North Pacifi c Ocean to 

southern California, mostly in intertidal areas to depths 

of 30 m among rocks and algae. Maximum length of 

adults is 200 mm TL (Hart, 1973); age at maturity is 

2–3 years (Peppar, 1965). In southern British Columbia, 

Washington, and California, spawning occurs in January–March (Marliave, 1975a; Schultz and DeLacy, 1932; 

Wang, 1986). Eggs 1.3–1.5 mm in diameter, with one 

large (0.33–0.41 mm in diameter) and one smaller oil 

globule, are laid in pedestal-shaped masses of 2000–3000 
66 Professional Paper NMFS 15 

eggs under or between rocks or shells and guarded by the 

female during a 3-week incubation period (Schultz and 

DeLacy, 1932; Peppar, 1965; Marliave, 1975a; Matarese 

et al., 1989). Larvae hatch at 7.5 mm TL (Peppar, 1965).

Larval distribution (Fig. 34). Larvae of High Cockscomb 

are found in the Bering Sea north of Unimak Island; near 

Amchitka Island in the western Aleutian Island Archipelago; west of and around Unimak Pass; into the Gulf of 

Alaska on the shelf and around Kodiak Island; Straits of 

Georgia, Canada; Puget Sound, Washington; and the coast 

of northern Oregon. In Alaskan waters, larvae (5.8–20.0 

mm) are collected in April–June with maximum occurrence observed in May; the larva off Oregon (20.0 mm) 

was collected in April. It is likely that spawning occurs 

earlier in the year at the lower latitudes.

Morphology (Tables 34, 35). Notochord flexion in 

A. purpurescens begins at 9.4 mm and is complete at 

12.0–13.0 mm. Postfl exion larvae are 12.1–20.0 mm. 

Transformation can begin by 12.0 mm, and it was 

complete soon after fl exion in reared specimens; our 

20.0-mm fi eld-collected specimen had no characteristics that indicated that transformation had begun. The 

Figure 34

Distribution and abundance of larvae of High Cockscomb (Anoplarchus purpurescens) collected in the Bering Sea; Gulf of 

Alaska; Straits of Georgia, British Columbia, Canada; Puget Sound, Washington; and North Pacific Ocean off the coast of 

Oregon. Squares indicate collections made by bongo net; black circles indicate additional nonquantitative collections (presence only) and may represent more than one occurrence. Isobath=1000 m.
67

Figure 35

Larvae of High Cockscomb (Anoplarchus purpurescens): (A) preflexion stage, 6.1 mm standard length (SL); (B) flexion 

stage, 9.0 mm SL; (C) late flexion stage, 12.0 mm SL; (D) postflexion stage, 14.3 mm SL, composite of 13.2- and 14.3-mm 

SL specimens, University of Washington (UW) 145895 and UW 145872; and (E) transformation stage, 12.0 mm SL. Illustrations (A–C, E) from Matarese et al. (1989, reared) and (D) by B. Vinter.
68 Professional Paper NMFS 15 

28.0- and 37.0-mm cleared and stained specimens were 

juveniles, indicating that this stage begins at <28.0 mm. 

Larvae are slender-bodied (mean body depth 9.0% SL, 

initially), increasing in size slightly (mean body depth 

9.9% SL) in the postfl exion stage. Relative snout length 

increases, and head length and snout to anus length increase slightly, from the prefl exion stage to the postfl exion stage. Relative eye diameter decreases from the prefl exion stage to the postfl exion stage. 

Pigmentation (Fig. 35). Larvae of A. purpurescens are 

lightly pigmented throughout larval development. There 

is no dorsal head pigment until after the postfl exion 

stage. A small melanophore persists at the base of the 

cleithrum through all stages. In prefl exion larvae, three 

or more dorsal pigment spots are present on the gut, and 

several spots cover the hindgut. A line of pigment extends three-fourths of the way along the ventral midline 

of the gut and persists through the transformation stage. 

Table 34

Body proportions of High Cockscomb (Anoplarchus purpurescens) larvae. Except for standard length (SL), 

values given for each body proportion are expressed as percentage of SL or head length (HL): mean, ± 

standard deviation, and range (in parentheses). 

 Prefl exion Flexion Postfl exion

Sample size 152 27 8

Standard length (mm) 7.3±1.1 (5.2–9.3) 10.3±0.8 (9.4–12.9) 15.2±2.2 (13.1–20.0)

Head length/SL 13.8±1.3 (10.5–18.1) 13.7±1.2 (11.5–16.4) 14.8±1.8 (12.4–17.3)

Snout length/HL 23.1±6.7 (10.5–39.3)1 24.9±4.3 (17.2–32.1)2 26.7±4.9 (20.8–33.9)

Eye diameter/HL 39.8±5.8 (26.5–57.1)3 32.6±3.3 (27.5–42.4) 27.2±3.2 (22.9–32.8)

Snout to anus length/SL 42.0±2.4 (37.0–49.3)4 41.8±1.8 (38.3–45.2) 43.2±1.6 (41.0–45.4)

Body depth/SL 9.0±1.2 (4.7–11.9) 8.9±0.7 (7.7–10.0) 9.9±0.3 (9.5–10.4)5

1 n=138 2 n=26 3 n=145 4 n=151 5 n=7

Table 35

Meristic counts of cleared and stained High Cockscomb (Anoplarchus purpurescens) larvae and juveniles. Counts are of 

ossifi ed elements only. Specimens between dashed lines (-----) were undergoing notochord fl exion. This species lacks pelvic 

fi ns.

 Branchi-

 Standard Dorsal Anal Pectoral ostegal Haemal Caudal-fi n

length (mm) fi n fi n fi n rays Abdominal Caudal Total spines Abdominal Caudal Total rays

 5.8 

 7.3 

 8.9 

 9.4 3 44 44 

 10.1 

 11.4 17 45 62 45 

 12.1 LVI I,40 5 17 43 60 43 17 44 61 4,6+6,2

 13.21 18 41 59 41 4,6+6,3

 14.5 XXXVII2 I,38 10 5 18 40 58 40 18 41 59 2,6+6,1

 17.1 LVII I,40 10 5 18 43 61 43 18 44 62 4,6+7,2

 28.0 LVII I,40 10 5 19 43 62 43 19 44 63 4,6+6,3

 37.0 LVII I,39 10 5 19 42 61 42 19 43 62 4,6+6,3

1 Dorsal-fi n and anal-fi n pterygiophores were stained, but fi n elements were not. 2 Anteriormost XVI dorsal-fi n spines weakly formed and unossifi ed.

Neural spines Centra

Spines, rays
69

Prefl exion larvae have a series of about 34–45 (usually 

<40) moderately spaced PVM. Melanophores (usually 

two) are present along the ventral margin of the notochord tip where the hypurals will form. 

Few pigmentation changes occur during the early fl exion stage. Up to four dorsal spots appear on the gut. The 

melanophores in the hypural area become prominent. 

Late fl exion larvae undergo several changes in pigment. 

Pigment appears laterally on the head. More pigment 

is visible in the area of the isthmus, with a short series 

of spots forming a ventral line. Up to eight dorsal spots 

are present on the gut, and the hindgut also has several 

spots. The PVM appear slash-like with melanophores 

that extend along the pterygiophores and hypaxial musculature. A few melanophores are on the dorsal midline 

at 75% and 90% SL and also on the caudal-fi n rays. 

Pigment on the dorsal surface of the gut is more diffi cult 

to see in postfl exion-stage larvae as the melanophores 

become embedded.

Beginning early in the transformation stage, the typical adult facial pigment pattern develops as settlement 

approaches. Although our 12.0-mm reared specimen 

(Fig. 35E) had juvenile pigmentation, we have not observed this pigment pattern in fi eld-caught specimens up 

to 20.0 mm. During transformation and early juvenile 

stages, melanophores cover nearly the entire head and 

body. Irregularly shaped patches and streaks of larger 

melanophores give the head a mottled appearance. The 

dorsal and ventral melanophores on the gut are retained 

from late larval stages during the early transformation 

stage. A row of regularly spaced unpigmented semicircular spots along the base of the dorsal fi n are more closely 

spaced anteriorly. Several large melanophores are present 

posteriorly near the distal margin of the dorsal fi n. Irregularly spaced PVM are present, some of which have 

an adjacent cluster of melanophores that extends onto 

the anal fi n. Slash-like PVM are present on the hypaxial 

myosepta just dorsally of the ventral midline. Pigment 

is also present on the dorsal and ventral margins of the 

hypural edge, and a large patch of small melanophores 

covers the central portion of the caudal fi n. Our 28- and 

37-mm specimens displayed juvenile pigmentation before 

they were cleared and stained.

Meristic features (Table 35). Principal caudal-fi n rays 

are present by 9.4 mm and ossifi ed by 12.1 mm. Procurrent caudal-fi n rays are present and begin to ossify 

by 12.1 mm before all are developed; all of these rays 

are ossifi ed by 13.2 mm. The counts of one superior and 

one inferior procurrent caudal-fi n rays observed on radiographs are lower than the counts previously reported 

for this species (3–4, 3–4; Watson, 1996). Anal-fi n elements and dorsal-fi n spines can be developed and ossifi ed 

by 12.1–17.1 mm. Pectoral-fi n rays are present by 11.5 

mm, developed by 13.2 mm, and ossifi ed by 14.5 mm. 

Pelvic fi ns are absent. Vertebral centra are present by 

9.5 mm and ossifi ed by 12.1–14.5 mm. The count of 16 

precaudal centra observed on radiographs is lower than 

the count previously reported for this species (17–19; 

Watson, 1996). Neural and haemal spines are ossifi ed 

before vertebral centra, by 11.4 mm, although, in one 

specimen, they were partially ossifi ed by 9.4 mm. Ranges 

of adult meristic counts are given in Table 2.

Species comparisons. Larvae of A. purpurescens are 

similar to not only other Xiphisterinae, particularly their 

congener A. insignis, but also species of Alectrias and the 

opisthocentrine Opisthocentrus ocellatus. Anoplarchus 

purpurescens is generally more lightly pigmented than 

is A. insignis throughout development. Unlike A. insignis, pigmentation on the head and nape is absent in A. 

purpurescens until near the end of the postfl exion stage, 

and melanophores posteriorly above the notochord are 

absent in A. purpurescens. Anoplarchus purpurescens

typically has fewer PVM (34–45, usually £40) during the 

prefl exion stage than does A. insignis (45–55). Anoplarchus purpurescens can be distinguished from species of 

Alectrias by the absence of pigment on the head in the 

fl exion and postfl exion stages (present in Alectrias spp.), 

absence of internal pigment above the notochord on the 

caudal peduncle (present in Alectrias spp.), usually fewer 

caudal myomeres (40–46 vs. 44–50), and usually fewer 

dorsal-fi n spines and anal-fi n rays (LIV–LX; I–II,35–42 

vs. LV–LXVI; I,39–45). Larvae of Anoplarchus purpurescens are more slender-bodied than are larvae of Alectrias

species (mean body depth: 8.9–9.9% SL vs. 11.5–10.7% 

SL) during the fl exion and postfl exion stages. Larvae of 

A. purpurescens can be distinguished from O. ocellatus by a line or row of small, punctate, ventral melanophores that extends along the anterior three-fourths of 

the length of the gut rather than an evenly spaced row 

of large, stellate melanophores ventrally along the entire length of the gut. In addition, A. purpurescens has a 

shorter snout to anus length (mean: 41.8–43.2% SL vs. 

48.2–53.2% SL) and fewer precaudal vertebrae (17–19) 

than has O. ocellatus (21–23).

Cebidichthys violaceus, Monkeyface Prickleback 

(Figs. 36, 37; Table 36)

Literature. Fitch and Lavenburg (1971), Burge and 

Schultz2, Eschmeyer et al. (1983), Wang (1986), Marshall 

and Wyllie-Echeverria (1992), Lea and Reilly (2001), 

PG&E3.

2 Burge, R. T., and S. A. Schultz. 1973. The marine environment 

in the vicinity of Diablo Cove with special reference to abalones 

and bony fi shes. Mar. Res. Tech. Rep. No. 19, 433 p. Calif. Dep. 

Fish Game, Long Beach, CA.

3 PG&E (Pacifi c Gas & Electric Co.). Diablo Canyon Power Plant 

316(b) Demonstration Project. Document No. E9-055.0. Prepared for Pacifi c Gas & Electric Co. by Tenera Environmental 

Services. March 2000.
70 Professional Paper NMFS 15 

Material examined. 10 specimens (6.2–14.4 mm) from 

the coast of California (see Appendix for details).

Distribution and life history. Cebidichthys violaceus

is common in tidepools and rocky intertidal and subtidal habitats to depths of 24 m (Burge and Schultz2; 

Eschmeyer et al., 1983) from southern Oregon to northern Baja California, Mexico. This species is the only 

stichaeid for which there is a small, specialized recreational fi shery (Lea and Reilly, 2001). Adults grow to 

760 mm TL and may live to 18 years (Eschmeyer et al., 

1983; Marshall and Wyllie-Echeverria, 1992). Males 

and females reach 50% maturity at fi ve years and 100% 

maturity at eight years. Fecundity ranges from 17,500 

to 46,000 eggs, which are deposited on subtidal rocky 

surfaces from January to May, with a peak in February–April (Marshall and Wyllie-Echeverria, 1992); eggs 

(about 2 mm diameter) are laid in the form of a ball 

with 6000–8000 eggs in a cluster (Fitch and Lavenburg, 

1971). Nests are guarded by one or both parents (Lea 

and Reilly, 2001). Hatch size is about 4.8 mm; larvae are 

pelagic (PG&E3).

Larval distribution (Fig. 36). Larvae of Monkeyface 

Prickleback have not been collected during AFSC ichthyoplankton surveys. Larvae at lengths of 4.8–24.6 mm 

Figure 36

Distribution of larvae of Monkeyface Prickleback (Cebidichthys violaceus) collected in the North Pacific Ocean off the California coast. Each symbol may indicate more than one occurrence. Isobath=100 m. Inset shows location of area expanded 

in the main map.
71

(most specimens were 6.6–7.6 mm) have been collected 

off central California between Point Buchon and Point 

San Luis in January–July; greatest abundances occur in 

April–May (PG&E3). Larvae also have been collected in 

San Francisco Bay, California (Wang, 1986).

Morphology (Table 36). Notochord fl exion in C. violaceus begins by 8.6 mm and is complete by 13.6 mm. The 

two postfl exion larvae examined were 13.6–14.4 mm; 

longer postfl exion larvae would probably be found in the 

fi eld. Lengths at which transformation and the juvenile 

stage begin are unknown. Larvae are slender-bodied with 

a mean body depth of 8.2% SL in the prefl exion stage 

(n=2), increasing to 8.6% SL and 9.6% SL in the fl exion 

and postfl exion stages. Relative head length and snout to 

anus length increase slightly through development, and 

relative snout length and eye diameter decrease.

Pigmentation (Fig. 37). Larvae are moderately pigmented. Among stichaeids, only larvae of C. violaceus have 

lateral pigment in a single bar on the postanal body. The 

head is unpigmented through at least the early postfl exion stage. In prefl exion larvae, ventral pigment is present 

below the pectoral-fi n base anteriorly on the gut. A series 

of about 8–10 melanophores occurs dorsally along the 

gut with 3–4 additional spots on the hindgut. Anteriorly, 

Figure 37

Larvae of Monkeyface Prickleback (Cebidichthys violaceus): (A) preflexion stage, 6.7 mm standard length (SL), University of Washington (UW) 144863; (B) flexion stage, 9.2 mm SL, UW 144865; and (C) postflexion stage, 14.4 mm SL, 

UW 144869. Illustrations by R. Cartwright.
72 Professional Paper NMFS 15 

a few melanophores are present laterally on the gut near 

the base of the pectoral fi n. Postanal pigment consists of 

one large bar and a series of PVM. The bar is composed 

of small melanophores between postanal myomeres 24 

and 33. 

During the fl exion stage, the dorsal melanophores on 

the gut become internal, but anterior and hindgut melanophores remain external. The postanal pigment bar 

melanophores and PVM are small and numerous. A single melanophore is present on the ventral margin of the 

caudal peduncle. Spots appear along the hypural edge 

and extend onto the developing caudal-fi n rays. By the 

early postfl exion stage, a few spots are present externally around the base of the pectoral fi n, but pigmentation 

otherwise changes little. Light external pigment along 

the lateral surface of the gut and hindgut is still visible. 

The postanal body pigment bar remains approximately 

between myomeres 24 and 33. Pigment is relatively unchanged on the caudal peduncle and on the caudal-fi n 

rays along the hypural edge. 

Meristic features. No specimens of C. violaceus were 

available to clear and stain. Development of principal 

caudal-fi n and pectoral-fi n rays was evident by 9.2 mm 

in unstained specimens of mid-fl exion larvae. All median 

and paired fi n elements appear developed in postfl exion 

larvae by 14.4 mm. The posterior two-thirds of the dorsal fi n is composed of soft rays. Pelvic fi ns are absent. 

Ranges of adult meristic counts are given in Table 2.

Species comparisons. Adults and larvae of C. violaceus are notably different in appearance from all other 

Stichaeidae in our study area. Most notably, the posterior two-thirds of the dorsal fi n is composed of soft rays. 

Although somewhat similar in general morphology to 

species of Anoplarchus, larval C. violaceus have a longer 

mean snout to anus length (48.6–50.6% SL vs. 41.1–

43.2% SL) and no ventral pigment on the gut; in contrast, species of Anoplarchus have a row or line of pigment on the gut. The most noticeable difference between 

C. violaceus and other xiphisterine larvae is the presence 

of a bar of small melanophores that cover an area of the 

postanal body at about 75–85% SL, just anterior of the 

caudal peduncle. Larvae of C. violaceus also have more 

preanal myomeres (vertebrae) than do larvae of both 

A. insignis and A. purpurescens (23–25 vs. 16–19) and 

more total vertebrae than has A. purpurescens (65–71 

vs. 58–64). 

Phytichthys chirus, Ribbon Prickleback 

See “Other material” below.

Xiphister atropurpureus, Black Prickleback 

See “Other material” below.

Xiphister mucosus, Rock Prickleback 

See “Other material” below.

Other material

This section provides brief accounts and descriptions of 

larval stages of eight stichaeid taxa represented by very 

limited material from our collections. On the basis of 

general morphology, pigmentation, and counts of meristic features, these specimens, except Acantholumpenus 

mackayi (Tokuya and Amaoka, 1980), were tentatively 

identifi ed. All individuals shared the following characters: bodies generally slender, guts short (snout to anus 

length of about 50% SL or less), PVM typically present, 

myomeres 50–83, and other meristic features that overlap with at least one known stichaeid species. 

A single prefl exion larva, 10.6 mm (Fig. 38A), collected in the Beaufort Sea, bore a strong resemblance to 

Stichaeus punctatus (subfamily Stichaeinae, Fig. 3) and 

Table 36

Body proportions of Monkeyface Prickleback (Cebidichthys violaceus) larvae. Except for standard length 

(SL), values given for each body proportion are expressed as percentage of SL or head length (HL): mean, 

± standard deviation, and range (in parentheses). 

 Prefl exion Flexion Postfl exion

Sample size 2 4 2

Standard length (mm) 6.4±0.3 (6.2–6.6) 9.7±1.1 (8.6–10.7) 14.0±0.6 (13.6–14.4)

Head length/SL 15.1±1.0 (14.4–15.8) 16.5±2.1 (13.8–18.8) 16.7±2.5 (14.9–18.5)

Snout length/HL 29.9±3.3 (27.6–32.2) 24.7±4.4 (21.3–31.1) 20.8±6.2 (16.4–25.2)

Eye diameter/HL 32.2±1.1 (31.5–33.0) 27.5±3.4 (23.5–31.8) 22.2±3.3 (20.0–24.7)

Snout to anus length/SL 48.6±1.4 (47.6–49.5) 49.6±1.7 (47.5–51.2) 50.6±0.3 (50.3–50.8)

Body depth/SL 8.2±0.6 (7.8–8.6) 8.6±0.6 (8.1–9.4) 9.6±1.2 (8.8–10.4)
73

was very likely Eumesogrammus praecisus. This larva 

had a short gut (37.5% SL) with pigment along its ventral surface, melanophores on the dorsal midline, internal pigment above the notochord, PVM, and, most notably, slash-like hypaxial pigment along the myosepta. 

In addition, this larva had 50 myomeres, which is within 

the vertebral range (50–53) for E. praecisus; the count 

of 53 total vertebral centra observed on radiographs is 

greater than the count previously published for this species (50–52; Mecklenburg et al., 2002). This larva differed from prefl exion S. punctatus by having a more 

slender body (body depth: about 5.0% SL vs. 7.9–11.2% 

SL), a continuous line of ventral pigment along the entire 

length of the gut (vs. restricted to the anterior one-half 

of the gut), and larger size during the prefl exion stage 

(10.6 mm SL vs. 6.4–9.0 mm SL). Adult E. praecisus occur in the northern Bering, Chukchi, and Beaufort seas, 

Canadian Arctic Ocean, off the coast of Greenland, and 

in the North Atlantic Ocean. Larvae are likely unknown 

because few ichthyoplankton collections have been made 

within this species’ range. With the increasing interest in 

Arctic ecosystems, it is likely that more of these larvae 

will be collected in the future.

Grigor’ev (1992a) described and illustrated two postfl exion larvae that he identifi ed as Chirolophis snyderi 

(28.0 and 33.0 mm TL) collected near Kamchatka, 

Russia. However, because of the poor quality of these 

illustrations, it is unclear how larval C. snyderi are 

distinguished from C. decoratus and C. nugator other 

than that the collection locality reported for C. snyderi

by Grigor’ev (1992a) excludes C. decoratus and C. nugator.

We examined 12 larvae of Chirolophis cf. snyderi that 

were clearly not C. decoratus or C. nugator. These larvae 

were collected with bongo and neuston nets and Tucker 

trawls; 11 larvae were caught in the Gulf of Alaska and 1 

was collected in the Bering Sea. Of these specimens, two 

were illustrated (Fig. 38B–C) and four were cleared and 

stained. Flexion larvae of C. cf. snyderi are most likely to 

be confused with C. decoratus and C. nugator, but they 

can be distinguished by a row of evenly spaced, internal melanophores above the urostyle that extends anteriorly only to the caudal peduncle. In C. decoratus and 

C. nugator, urostyle pigment is absent, but the internal 

notochord pigment extends anteriorly from the caudal 

peduncle to midbody or farther. Larvae of C. cf. snyderi 

lack dorsal melanophores on the gut, unlike C. decoratus and C. nugator. However, the illustrated 18.5-mm C. 

cf. snyderi specimen had two deeply embedded melanophores that were just above the dorsal surface of the gut 

at about 75% gut length and not visible from a lateral 

view. Mean body depth is greater in postfl exion C. cf.

snyderi than in C. decoratus or C. nugator (14% SL vs. 

8.9% SL and 9.9% SL). Although there is minimal overlap, meristic counts of C. snyderi, particularly vertebrae, 

typically are intermediate between C. decoratus and C. 

nugator (61–65 vs. 65–69 and 55–61). Another species, 

Matcheek Warbonnet (Chirolophis tarsodes; Jordan and 

Snyder, 1902), has meristic counts similar to C. snyderi

(Table 2), but it is unlikely that these examined larvae 

were C. tarsodes because this species is known from very 

few adult specimens (Mecklenburg et al., 2002). Until we 

obtain and identify additional specimens, we cannot be 

certain of this larval identifi cation. 

A single late-postfl exion larva of Acantholumpenus 

mackayi (36.5 mm) was collected in Norton Sound, Alaska (subfamily Lumpeninae, Fig. 38D; see Tokuya and 

Amaoka, 1980: fi g. 11). This larva was similar to other 

larval Lumpeninae, particularly Lumpenella longirostris, 

in having a slender body (body depth 8.5% SL), a long 

snout, a long gut (snout to anus length 47% SL) with 

melanophores on its lateral surface, and what appeared 

to be a very faint row of embedded PVM that were visible only in ventral view. As with the other members of 

the subfamily Lumpeninae, the presence of mediolateral 

pigment indicates that an A. mackayi larva will soon 

enter the transformation stage. Postfl exion larvae of 

A. mackayi are distinguished from L. longirostris and 

Lumpenus sagitta by the presence of mediolateral pigment by about 35 mm (vs. absence before 40 mm). In addition, A. mackayi has more precaudal (27–30 vs. 22–25) 

and total (76–80 vs. 70–75) vertebrae than does L. longirostris. Meristic counts of A. mackayi and L. sagitta 

generally overlap, but A. mackayi may have fewer caudal vertebrae (47–50 vs. 46–55). Fin-ray counts for the 

examined larva (D:LXVIII; A:II,41; P1:14; P2:I,3) were 

consistent with A. mackayi (Table 2). This specimen appeared to have >75 myomeres, a number that is within 

the known vertebral range for this species (76–80).

Two prefl exion larvae with lenths of 10.3 and 10.8 

mm were collected near Vancouver Island, British Columbia, Canada (Fig. 39A) and another larva (length unknown4) was sampled from southeast Alaska. These three 

larvae were very likely Lumpenopsis hypochroma (Hastings and Walker, 2003; Mecklenburg and Sheiko, 2004). 

These specimens were somewhat similar to Opisthocentrus ocellatus (subfamily Opisthocentrinae) in having a 

relatively long gut (about 47% SL) with a row of large 

melanophores along its ventral surface. These larvae had 

an unusual feature: 11–15 internal anterior PVM separated by an unpigmented space of 3–4 myomeres from 

4 external PVM. Two mediolateral melanophores were 

directly above the posterior PVM. From their general appearance, and considering species for which there is no 

larval description, the most likely identity of these larvae was L. hypochroma. Hastings and Walker (2003) reported 54–55 vertebrae for this species (counts based on 

three specimens), and our larvae had 54–55 myomeres. 

Because they had similar vertebral counts, our larvae 

were very possibly L. hypochroma; however, until more 

adults are available from which to obtain additional 

4 Specimen was examined by M. S. Busby, but it is now lost.
74 Professional Paper NMFS 15 

Figure 38

Larvae from other material: (A) Fourline Snakeblenny (Eumesogrammus praecisus), preflexion stage, 10.6 mm standard 

length (SL), University of Washington (UW) 145179; (B) Chirolophis cf. snyderi, flexion stage, 18.5 mm SL, UW 145178; 

(C) Chirolophis cf. snyderi, postflexion stage, 21.0 mm SL, UW 144864; and (D) Blackline Prickleback (Acantholumpenus 

mackayi), late-postflexion stage, 36.5 mm SL, UW 145177. Illustrations by A. Overdick.
75

Figure 39

Larvae from other material: (A) Y-prickleback (Lumpenopsis hypochroma), preflexion stage, 10.3 mm standard length 

(SL), University of Washington (UW) 145180; and (B) Lesser Prickleback (Alectridium aurantiacum), postflexion stage, 

21.0 mm SL, UW 145871 (original specimen lost). Illustrations by A. Overdick. 

counts and we locate additional larval specimens, we 

cannot be absolutely certain of the identifi cation. 

Two postflexion larvae with lengths of 20.0 and 

21.0 mm (Fig. 39B) collected during a 1997 cruise in 

the southeastern Bering Sea resembled Stichaeus punctatus (subfamily Stichaeinae, Fig. 3) and were tentatively 

identifi ed as Eumesogrammus praecisus (Fig. 38A). These 

larvae had a short gut (about 40% SL) with a line of 

ventral pigment, melanophores on the dorsal midline, internal pigment above the notochord, PVM, and slash-like 

hypaxial pigment along the myosepta. However, the best 

match for the specimens’ meristic counts (D: LIII, LXIII; 

A: I,40 or 41, I,43 or 44; P1: 12; P2: absent; vertebrae: 63 

[20+43], 67 [21+46]) is Alectridium aurantiacum (subfamily Xiphisterinae), among stichaeid species reported 

from the study area. All Xiphisterinae and their larvae 

lack pelvic fi ns, but pelvic fi ns are present in Stichaeinae. 

Matarese et al. (1989) provided species accounts for 

Phytichthys chirus, Xiphister atropurpureus, and X. mucosus larvae on the basis of material collected in Barkley 

Sound, British Columbia, Canada, by Marliave (1975a, 

1975b; as presented in Matarese et al., 1989). Wang 

(1986) reported information on early life history of Xiphister species from the Sacramento-San Joaquin estuary. After reexamination of the material from Marliave 

(1975a, 1975b, as presented in Matarese et al., 1989) 

and examination of new material, it became apparent 

that the use of pigmentation to distinguish prefl exionand fl exion-stage larvae of these three taxa is problematic. Yolk-sac larvae (Fig. 40A) of X. atropurpureus from 

known parents hatch at about 6.3 mm with an ambercolored yolk sac and a short gut (about 38% SL). Larvae 

have six evenly spaced dorsal melanophores on the gut 

with a single spot on the hindgut, about 38 PVM with 

the posteriormost 5–6 appearing dash-like, and about 

5–6 small, widely spaced melanophores on the dorsal 

midline that begin at about midbody and extend to the 

caudal peduncle. Notochord fl exion in X. atropurpureus

occurs between 12.0 and 16.0 mm. In our limited material, individuals at lengths of about 12.0 mm showed 

early hypural development but no fl exion of the notochord, and individuals at lengths of 15.8 and 16.0 mm 

were clearly in the postfl exion stage (Fig. 40B).

We have three wild-caught postfl exion specimens of 

Phytichthys chirus from Puget Sound, Washington (Fig. 

40C). Phytichthys chirus and species of Xiphister can 

usually be identifi ed with counts of abdominal vertebrae 

(precaudal myomeres) (Table 2). We measured, cleared, 

and stained larvae of each prospective taxon and, after 

verifying our identifi cations, determined that larvae of P. 

chirus are generally similar to species of Xiphister. However, P. chirus is deeper bodied during the postfl exion 
76 Professional Paper NMFS 15 

Figure 40

Larvae from other material: (A) Black Prickleback (Xiphister atropurpureus), yolk-sac stage, 6.3 mm standard length 

(SL), University of Washington (UW) 144848; (B) X. atropurpureus, postflexion stage, 16.0 mm SL, UW 145176; 

and (C) Ribbon Prickleback (Phytichthys chirus), postflexion stage, 19.0 mm SL, UW 145181. Illustrations by A. 

Overdick.

stage (13.5% SL) compared to both Xiphister species (10.5–12.0% SL), and this species has a longer gut 

(snout to anus length 47% SL) than does X. atropurpureus (43% SL). Phytichthys chirus typically has more 

precaudal vertebrae (24–25) than does X. atropurpureus

(22–24) and fewer than does X. mucosus (29–31). Lastly, 

postfl exion P. chirus has a double row of irregularly 

spaced melanophores along the dorsal midline. Similar 

pigmentation is present in X. atropurpureus, but it is 

less pronounced and does not extend as far anteriorly. 

From the limited material examined, dorsal midline pigment was apparently absent in X. mucosus (specimens 

not suitable for illustration). However, it is presently unclear how useful pigment characters are in distinguishing 

among these three species because very few specimens 

were available for study. Additional larvae of all three 

taxa could likely be collected by intertidal and nearshore 

ichthyoplankton surveys. 
77

Discussion

With this study, at least one early life history stage is 

now known for most of the 27 stichaeid species reported 

from the NEP and Bering Sea; exceptions are Eumesogrammus praecisus, Lumpenopsis hypochroma, and Alectridium aurantiacum, all tentatively identifi ed and illustrated herein. Illustrations of complete developmental series (prefl exion, fl exion, and postfl exion; some previously 

published) are collectively presented in this guide for 14 

species, and there are at least 2 developmental stages 

presented for an additional 6 species. New, original illustrations of 16 prefl exion, 17 fl exion, 22 postfl exion, 

and 6 transformation stages are presented for 25 species. 

This guide represents a comprehensive source of original morphological, meristic, and pigmentation characters 

that will ensure accurate identifi cation of the early life 

history stages of this family of cryptic nearshore fi shes. 

Additional work on stichaeids, such as molecular studies on phylogenetic relationships and effects of ecosystem changes on species assemblages, will benefi t from the 

completion of this guide. However, there are still gaps 

in our knowledge, especially of egg and juvenile stages 

(Table 37). Additional collections, particularly from intertidal and nearshore habitats, are needed to fi ll some of 

these gaps. Eggs could be obtained from adults spawning 

in captivity, from extensive intertidal surveys, or by divers in subtidal areas. Larvae and juveniles might also be 

collected by similar methods in these habitats. It is likely 

that juveniles of some species are already represented in 

regional fi sh collections, but location and examination 

of them was beyond the scope of this study. Species of 

Phytichthys and Xiphister are likely collected as larvae 

in intertidal and nearshore surveys. Ichthyoplankton surveys in the more northern reaches of the Bering Sea and 

Arctic Basin likely will provide additional specimens and 

undescribed early life history stages of Acantholumpenus 

mackayi and E. praecisus. Additional material might also 

help clarify some of the diffi culties in identifi cation of 

larvae of the Arctic species of Lumpenus, particularly L. 

fabricii and L. medius. 

Meristic, morphological, and pigmentation characters are all useful for species-level identifi cation of larval and early juvenile pricklebacks (Table 38). Although 

many of the character states and meristic values reported 

here are new, some are updated from previous studies. 

Examples include myomere counts for Stichaeus punctatus that Faber (1976) reported as <50, but we (and 

others) found 51–55, and PVM for Lumpenus maculatus

that had been usually 30–40 (also from Faber, 1976), but 

we report 34–41. We also found length at transformation 

of S. punctatus begins at about 28 mm and is complete 

by 35 mm, lengths that are greater than Faber’s (1976) 

value of about 25 mm. Grigor’ev (1992b) reported 14 

precaudal myomeres for L. medius and 40–42 PVM for 

L. fabricii, but we found 21–25 precaudal myomeres for 

L. medius and 33–45 PVM (prefl exion–postfl exion) for 

L. fabricii. 

Counts of precaudal, caudal, and total vertebrae 

(myomeres) and anal-fi n elements are the most helpful 

meristic characters for generic and species-level identifi -

cations. Body depth and gut length (snout to anus length) 

are the most important morphological characters. Number of PVM can sometimes be a distinguishing pigmentation characteristic or at least can help to narrow the 

possible choices (e.g., Chirolophinae and Xiphisterinae). 

Although some stichaeid taxa have very distinct pigmentation patterns (e.g., S. punctatus, Plectobranchus evides, 

and Cebidichthys violaceus), others have more subtle differences within their respective subfamilies (e.g., Chirolophinae: Bryozoichthys and Chirolophis; Lumpeninae: 

Lumpenella, Lumpenus, and Poroclinus) or genera (e.g., 

Anoplarchus and Xiphister). In the genus Lumpenus, 

however, pigmentation alone is not useful to distinguish 

species. 

Another useful character to aid identifi cation is the 

length at stage of development, particularly the size at 

onset of transformation. It is worth noting, however, that 

we observed variation within taxa in length at transformation, particularly between reared and wild-caught 

individuals. Laboratory-reared larvae may transform 

early as a result of the proximity of suitable substrate 

for settlement in the aquaria. Larval Xiphisterinae transform at ≤35 mm, but Lumpeninae are generally >35 mm 

at transformation (Table 38). We observed a wide range 

of lengths at transformation for wild-caught individuals 

in these subfamilies, including species of Lumpenus and 

Anoplarchus. It has been suggested that larvae of some 

intertidal fi shes are locally retained by physical forces 

and are, therefore, genetically isolated populations distinctly adapted to their environments (Marliave, 1986). 

Adaptation to physical forces could infl uence length at 

transformation because it has been suggested that larvae 

of some species may be able to delay transformation until they encounter suitable juvenile habitat (Moser, 1981; 

McCormick, 1999). Larval behavior might also be involved, especially for these species that sometimes demonstrate prolonged larval stages (e.g., Lumpenus). Most 

of the specimens we examined were collected in the fi eld, 

but some were reared in the laboratory. Transformation 

of reared specimens at smaller sizes, compared to wildcaught specimens, is evident in our developmental series 

of Anoplarchus purpurescens (Fig. 35D wild-caught; 

A–C, E reared). 

Our observations that ossifi cation does not occur until 

postfl exion in some species, even when the larvae appear 

well developed and in good condition, indicate that there 

may be problems with some of the specimens that were 

cleared and stained. In some instances, smaller specimens 

showed indications of ossifi cation, but larger specimens 

did not (e.g., Chirolophis nugator, Lumpenus maculatus, L. sagitta, Anoplarchus insignis, A. purpurescens). 
78 Professional Paper NMFS 15 

Table 37

Summary of knowledge of early life history stages of the family Stichaeidae in the northeastern Pacifi c 

Ocean and Bering Sea based on this study.

Life history stage

Taxon Egg Prefl exion Flexion Postfl exion Juvenile

STICHAEINAE 

Eumesogrammus praecisus X1 

Stichaeus punctatus X X X X X

 

CHIROLOPHINAE 

Bryozoichthys lysimus X X X X

Bryozoichthys marjorius X X X X

Chirolophis decoratus X X X X

Chirolophis nugator X X X X 

Chirolophis snyderi X1

Gymnoclinus cristulatus X 

 

LUMPENINAE 

Acantholumpenus mackayi X 

Lumpenella longirostris X X X X

Lumpenus fabricii X X X X

Lumpenus maculatus X X X X

Lumpenus medius X X X X

Lumpenus sagitta X X X 

Poroclinus rothrocki X X X X

 

OPISTHOCENTRINAE 

Lumpenopsis hypochroma X1 

Opisthocentrus ocellatus X X X X

Plectobranchus evides X X 

 

XIPHISTERINAE 

Alectrias spp. X X X 

Alectridium aurantiacum X1

Anoplarchus insignis X X X X

Anoplarchus purpurescens X X X X X

Cebidichthys violaceus X X X X 

Phytichthys chirus X X1 X1 X 

Xiphister atropurpureus X X1 X1 X 

Xiphister mucosus X X1 X1 X1

1 Identifi cation is tentative.

There were also instances when results differed between 

two cleared and stained specimens of the same length 

(e.g., L. maculatus, L. sagitta). In these cases, it is possible that preservation in acidic formaldehyde could have 

decalcifi ed bones in early ossifi cation. It should also be 

noted that the typical alcian blue solution used to stain 

cartilage contains acetic acid, which may also decalcify 

skeletal elements just beginning to ossify. Acquisition 

of additional specimens of larvae and early juveniles of 

these species would permit more clearing and staining 

and would provide additional osteological examinations. 

A general observation made during this study was that 

specimens initially preserved in marble chip buffered 

formalin cleared more slowly and stained with less consistent results than did specimens preserved in sodium 

borate buffered formalin. The difference in preservation 

technique is likely the cause of the variability of results 

observed in cleared and stained specimens.

Larval characters: relationships within family 

Stichaeidae

The phylogeny of stichaeid fi shes is poorly understood, 

and relationships of the subfamilies have not been investigated. Stoddard (1985), Yatsu (1986), Kartavtsev 

et al. (2009), and Radchenko et al. (2009) have 
79

suggested that the current classifi cations (subfamilies 

of Makushok (1958) are not monophyletic groups. We 

have followed the subfamily classifi cation of Mecklenburg and Sheiko (2004), which differs from Makushok 

(1958) only in that species from Makushok’s Alectriinae are included in Xiphisteriinae. A phylogeny of 

the family might be resolved through the use of modern cladistic analysis with multiple character sets and 

inclusion of all known taxa. In addition to adult morphological characters, some early life history features 

noted here will be a valuable addition to the systematic 

data set (Table 38) and might help resolve relationships 

within the family, particularly among the subfamilies. 

To include early life history characters into a proposed 

phylogeny of the family Stichaeidae and establish relationships within and among the subfamilies, it will be 

necessary to establish primitive (ancestral) larval character states through selection of multiple outgroup taxa, 

both within and outside Zoarcoidei.

Traditionally, the ronquils (family Bathymasteridae), 

have been considered the basal zoarcoid family, closely 

related to stichaeids, and would be an appropriate outgroup family (Anderson, 1994). Larvae of the bathymasterid Ronquilus jordani have been described ( Stevenson 

and Matarese, 2008: fi g. 1) and share several general 

morphological and pigment characters with Stichaeus 

punctatus (Fig. 3). Both of these species have deep bodies in postfl exion (≥12.5% SL) (more so for R. jordani) 

and short guts (<40% SL) with ventral pigment, rows of 

melanophores along the dorsal midline, internal pigment 

both above and below the notochord in the area of the 

caudal peduncle, and slash-like hypaxial melanophores 

along the myosepta. These attributes would be considered primitive larval characters, because they are shared 

with R. jordani, supporting the Stichaeinae as the basal 

stichaeid subfamily. The larva we identifi ed tentatively 

as Eumesogrammus praecisus also shares these characters, although postfl exion body depth was unknown (Fig. 

38A). In addition, Stichaeinae have the lowest counts of 

vertebrae in the family, and Anderson (1994) considered 

low vertebral counts a primitive character state among 

zoarcoid fi shes. With the exception of a deep body (9% 

SL vs. >12% SL), the slash-like hypaxial melanophores, 

and low vertebral counts, the genus Chirolophis, in the 

subfamily Chirolophinae, shares short gut and general 

pigmentation characters with the genus Stichaeus. 

Shared larval characters within other subfamilies support their close relationships. The Lumpeninae generally 

share the characters of sparse pigmentation with a row 

of PVM, melanophores present dorsally or laterally on 

gut, and slender bodies with relatively long guts (snout 

to anus lengths). Xiphisterine larvae are also very similar to one another with PVM, relatively short guts, medium body depth, and a lack of pelvic fi ns (Table 38). 

It should be noted, however, that counts of PVM are 

variable among taxa within the subfamilies and probably 

not phylogenetically informative. Although only notable 

in postfl exion larvae and adults, absence (loss) of pelvic 

fi ns might be a derived character.

Within other subfamilies, however, some species have 

marked differences in some of their larval characters. 

For example, within the Chirolophinae, postfl exion-stage 

Gymnoclinus cristulatus is deeper bodied (16% SL vs. 

6.3–10% SL) and more heavily pigmented than are other 

members of the subfamily (the genera Bryozoichthys and 

Chirolophis), which are all similar with short guts, PVM, 

internal pigment above the notochord, and pigment on 

the dorsal midline in varying degrees. In Opisthocentrinae, Opisthocentrus ocellatus larvae possess features 

more similar to some members of Xiphisterinae, including a long gut (snout to anus length 40–50% SL) with a 

row of melanophores on its ventral surface, PVM, and 

the absence of pelvic fi ns. Other Opisthocentrinae, such 

as the genera Kasatkia (Mecklenburg and Sheiko, 2004) 

and Pholidapus (Hatooka, 2002), lack pelvic fi ns, but 

their larvae are unknown. 

One unusual observation is that our larvae tentatively 

identifi ed as Alectridium aurantiacum (Fig. 39B) shared 

pigment characters of the Stichaeinae, but it was placed 

in Xiphisterinae primarily because it lacked pelvic fi ns. 

The absence of pelvic fi ns in Xiphisterinae (and some 

Opisthocentrinae) is an important character that distinguishes this subfamily from Stichaeinae. On the basis 

of meristic features, it is possible this larva was an undescribed species of Stichaeinae that has not yet developed pelvic fi ns. The striking similarities in pigmentation characters between postfl exion A. aurantiacum and 

S. punctatus, however, indicate a possible relationship 

that warrants further investigation. These similarities are 

pigment along the ventral midline of the gut and dorsal 

midline, PVM, and slash-like hypaxial pigment along the 

myosepta.

Larval characters: relationships within suborder 

Zoarcoidei

The phylogenetic relationships of the family Stichaeidae within the suborder Zoarcoidei (Bathymasteridae, 

Zoarcidae, Ptilichthyidae, Zaproridae, Anarhichadidae, 

Cryptacanthodidae, Pholidae, and Scytalinidae) are not 

well understood and need further study. Larvae of scytalinids are unknown, and zoarcids lack a larval stage. 

In addition to adult morphological characters, some 

early life history features are noted on the basis of unpublished sources (Ichthyoplankton Information System, 

http://access.afsc.noaa.gov/ichthyo/index.cfm), our observations during the course of this study, and ongoing 

research on bathymasterid and cryptacanthodid larvae. Bathymasterids, stichaeids, and pholids have similar pigment patterns with PVMs; zaprorids and cryptacanthodids share similar heavy pigment patterns and 
80 Professional Paper NMFS 15 Table 38 Summary of some important characters useful for identifi cation of larval and early juvenile stichaeids. Morphological measurements are mean percentages of standard length for larvae in the postfl exion stage, except for Ocellated Blenny (Opisthocentrus ocellatus), which was in the fl exion stage; postanal ventral me- lanophore (PVM) counts are for postfl exion stage. Lengths at developmental stage are standard length in millimeters unless otherwise noted. BD=body depth; SNA=snout to anus length; Post=postfl exion; Trans=transformation; abs=absent. Vertebrae Fins Pigment Morphology1 Length at developmental stage Taxon Precaudal Caudal Total Pelvic Anal PVM BD SNA Hatch Flexion Post Trans STICHAEINAE Eumesogrammus praecisus 15–17 35–37 50–53 I,3 I–II,29–32,II–III 40 Stichaeus punctatus 14–16 36–40 51–55 I,4 I–II,32–37 32–35 12.5 40.0 <6.4 9.0–16.4 16.4–28.2 >28.0,<35.0 CHIROLOPHINAE Bryozoichthys lysimus 16–17 53–58 69–75 I,3 I,47–53 46–48 8.5 34.0 <6.9 12.0–22.0 21.6–35.6 >36.0 Bryozoichthys marjorius 15–17 55–59 71–76 I,3 I,51–59 53–54 6.5 32.0 <7.7 11.0–24.0 21.4–40.0 >40.0,<55.0 Chirolophis decoratus 16–17 50–53 65–69 I,3–4 I–II,44–49 >40 9.0 35.0 <8.7 13.0–19.5 19.5–31.6 >32.0 Chirolophis nugator 14–16 42–46 55–61 I,3–4 I–II,37–42 35–37 10.0 37.0 <8.7 12.3–17.0 17.0–25.0 >25.0 Chirolophis snyderi 16–17 44–48 63–65 I,4 I–II,43–47 Chirolophis tarsodes 15 47–50 61–65 I,3–4 I,43–46 Gymnoclinus crisulatus 18–19 46–47 65 I,2 40–43 Variable 16.0 47.0 18.8–26.8 >26.8 LUMPENINAE Acantholumpenus mackayi 27–30 47–50 76–80 I,3 II,41–48 40 8.5 47.0 ?–35.0 >35.0 Lumpenella longirostris 22–25 47–49 70–75 I,2–3 II–V,36–42 42–44 6.5 43.0 <8.0 13.0–25.0 24.9–40.4 >40.0–50.0 Lumpenus fabricii 23–28 43–49 70–75 I,3 I,38–44 33–37 7.0 43.0 <11.9 13.0–21.0 20.0–46.0 45.0–50.0 Lumpenus maculatus 25–30 39–43 66–72 I,3–4 I–II,34–40 34–41 7.5 50.0 <8.0 13.0–25.0 20.0–66.2 >50.0 Lumpenus medius 21–25 42–50 65–70 I,3 I,37–42 29–38 8.0 44.0 <9.6 11.0–14.5 ?–46.5 >35.0 Lumpenus sagitta 26–29 46–55 75–82 I,3 I–II,44–50 44–53 6.5 44.0 <5.02 15.5–25.0 25.0–40.0 >40.0 Poroclinus rothrocki 18–20 45–48 64–68 I,3 III,40–44 40–41 8.5 44.0 <5.5 9.5–20.0 20.0–30.0 <28.0 OPISTHOCENTRINAE Lumpenopsis hypochroma 17–18 36–38 54–55 I,3 I,30–32 11–15+43 Opisthocentrus ocellatus 21–23 39–44 61–67 abs I–II,32–39 40 12.0 53.0 9.0–10.04 14.0–? <22.0 Plectobranchus evides 20–21 39–40 59–61 I,3 II–III,34–36 abs 9.5 47.0 <4.7 11.0–20.0 20.0–49.0 >49.0 XIPHISTERINAE Alectrias alectrolophus 17–20 47–50 64–70 abs I,41–45 Alectrias spp. 15–20 44–50 60–70 abs I,39–45 <40 11.0 41.0 <10.7 11.0 12.0–16.5 Alectridium aurantiacum 20–21 47 60–68 abs I,41–44 45 20.0–? Anoplarchus insignis 17–19 42–49 60–69 abs I,38–46 36 9.0 42.0 <5.2 9.0–14.0 14.0–20.6 <35.0 7.5 mm TL5; Anoplarchus purpurescens 16–19 40–46 58–64 abs I–II,35–42 <40 10.0 43.0 <4.76 9.0–13.0 12.1–21.0 <28.0 Cebidichtys violaceus 23–25 40–47 65–71 abs I–II,39–42 ≥40 9.5 51.0 ≤4.87 8.6–13.6 13.6–14.4 Phytichthys chirus 24–25 50–52 75–76 abs II–III,40–45 Xiphister atropurpureus 22–24 51–56 73–80 abs 0–I,49–55 Xiphister mucosus 29–31 44–53 73–83 abs 0–I,46–50 1 Morphometric values rounded to the nearest 0.5 mm. 2 Hart (1973). 3 Internal anterior PVM separated by an unpigmented space of 3–4 myomeres from four external PVM. 4 Shiogaki (1982). 5 Peppar (1965). 6 This study. 7 PG&E (text footnote 3).
81

development milestones; and larvae of both anarhichadids and ptilichthyids are unique in terms of morphology and pigment. However, there are some notable morphological and pigmentation similarities between larval 

stichaeids and other zoarcoids, particularly Bathymasteridae and Pholidae. 

For example, as previously noted, postfl exion Stichaeus punctatus (Fig. 3C) shares several general morphological and pigment characters with Ronquilus jordani

(Stevenson and Matarese, 2008: fi g. 1). Both of these species have deep bodies (≥12.5% SL) (R. jordani more so) 

and short guts (<40% SL) with ventral pigment, rows of 

melanophores along the dorsal midline, internal pigment 

both above and below the notochord in the area of the 

caudal peduncle, and slash-like hypaxial melanophores 

along the myosepta. All of these characters, except the 

slash-like hypaxial melanophores, also are shared with 

species of Bathymaster (Stevenson and Matarese, 2008: 

fi g. 9). Chirolophis (present study) also is similar in appearance to S. punctatus and R. jordani but lacks the 

slash-like hypaxial myomeres and is more slenderbodied. Many stichaeid larvae also resemble pholids with 

a long, slender body with a row of PVM; in most cases, 

however, pholids have a longer gut (>50% SL) and more 

vertebrae (>80). 

Systematics of the Stichaeidae requires further study 

as monophyly of the group has not been demonstrated. 

Since Makushok (1958), there have been few additional 

studies and attempts at classifi cation (Makushok, 1961; 

Gosline, 1968; Anderson, 1984; Stoddard, 1985; Yatsu, 

1986; Anderson, 1994). In each of these studies, the 

taxonomic scope was limited, specimens of some species 

were not examined, or early life history characters were 

not considered. 

Yabe and Uyeno (1996) reported similarities between 

zoarcoids and some cottoids. More recent molecular and 

morphological studies have provided interesting insight 

into the relationships of other zoarcoids (Zoarcidae, 

Pholidae) to gasterosteiforms (e.g., Aulorhynchidae, Gasterosteidae) and the cottoid lineage of the scorpaeniform 

(e.g., Cyclopteridae, Cottidae, Anoplopomatidae, Hexagrammidae) fi shes (Imamura and Yabe, 2002; Chen et 

al., 2003; Miya et al., 2003; Smith and Wheeler, 2004; 

Kartavtsev et al., 2009; Radchenko et al., 2009). Wiley 

and Johnson (2010) have formally recognized zoarcoids 

within the order Cottiformes, but a detailed study of the 

morphology and systematics of the Zoarcoidei is still 

needed. A more robust reconstruction of stichaeid phylogenetic relationships will require additional specimens 

with a more complete character set. New morphological 

data from adults and larvae will contribute to the overall 

understanding of the phylogenetic relationships of the 

Stichaeidae. 

With the completion of this guide, many early life history characters are now available to be used in studies of 

systematics in conjunction with molecular data (see Roje, 

2010). Larval fi shes possess unique characters and adaptations for planktonic life that are often very different 

from those of adults and are useful in studies of taxonomy and systematics (Moser and Ahlstrom, 1974). Early 

life history characters of stichaeid fi shes show promise 

for use in phylogenetic analyses and, along with molecular data, should be considered in future systematic studies of this very interesting family of fi shes. 

Acknowledgments 

Larval fi sh taxonomists at the Alaska Fisheries Science 

Center have been examining stichaeid larvae for more 

than 30 years. J. Dunn, B. Vinter, and K. Waldron initiated work in the 1970s by separating larvae into groups. 

B. Vinter compiled, measured, and illustrated many of 

the larvae that formed the basis of this study.

A joint project in the late 1970s and early 1980s 

with the University of Washington (UW) comparing inshore and offshore larval fi shes enabled further taxonomic studies that led to the identifi cation of unknown 

stichaeids. More species were identifi ed as part of zoarcoid research in the mid-1980s for the Ahlstrom Symposium. Studies were resumed in the late 1990s when 

UW student E. MacDonald began her thesis research on 

larvae of the genus Lumpenus.

We thank the following for providing hard-to-obtain 

specimens: K. Garrison and E. MacDonald (formerly UW); J. Marliave (Vancouver Public Aquarium); W. 

Watson (NOAA Southwest Fisheries Science Center); G. 

McGowen (Los Angeles County Museum); H. Imamura 

(Hokkaido University Museum of Zoology); D. Markle 

(Oregon State University); N. Laman (AFSC); C. Ehrler 

(Tenera Environmental Services); T. Pietsch, K. Maslenikov, and D. Roje (UW); C. Mecklenberg (California 

Academy of Sciences); and J. Paulic (Canada Department 

of Fisheries and Oceans).

Many scientists and interns at AFSC assisted throughout this study: B. Vinter (compiling and measuring 

specimens; illustrations while at fi rst employed by and 

later under contract to AFSC); K. Mier (statistics); S. 

Picquelle, formerly of AFSC, and J. Benson (GIS mapping); A. Fleischer and R. Takamine (interns; data collection and radiographs); A. Overdick (data collection, radiographs, and illustrations); L. De Forest (clearing and 

staining, preliminary review of manuscript); S. Remple, 

formerly of AFSC (clearing and staining and compiling 

data for materials examined); R. Cartwright, formerly 

of AFSC (data compilation and illustrations); and N. 

Laman, J. Orr, and D. Stevenson (of the “Tuesday Fish 

Lunch Group” for helpful discussions). Some original illustrations drawn by B. Vinter and R. Cartwright were 

later revised by A. Overdick.

We thank L. De Forest and D. Stevenson also for 

their review of the draft manuscript. W. Watson greatly 
82 Professional Paper NMFS 15 

improved the accuracy and quality of the draft manuscript by providing a detailed review and sharing his vast 

reserve of knowledge of larval fi sh taxonomy. The fi nal 

draft benefi ted from thoughtful reviews by D. Markle, E. 

Hilton, and an anonymous reviewer.

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86 Professional Paper NMFS 15 

Appendix

This appendix provides details, such as number and 

length of specimens and location, depth, sampling gear, 

and date of collection, on the materials examined for 

identifi cation of larval and early juvenile pricklebacks 

and on other material for other illustrated species. 

UW=University of Washington, HUMZ-L=Hokkaido 

University Museum of Zoology (larvae), OS=Oregon 

State University, MOCNESS=Multiple Opening/Closing 

Net and Environmental Sensing System.

Material examined

Stichaeus punctatus. 80 specimens (6.4–35.0 mm) 

examined. Chukchi Sea: UW 140709, 1 (19.4 mm), 

68°31.5′ N, 171°27.2′ W, 0–45 m depth, bongo net (net 

1), 15 August 2004; UW 140854, 1 (19.0 mm), 68°31.5′

N, 171°27.2′ W, 0–45 m depth, bongo net (net 2), 15 

August 2004. Bering Sea: UW 144841, 1 (8.9 mm), 

54°6.0′ N, 166°14.1′ W, 0–76 m depth, bongo net (haul 

5), 28 April 1994; UW 144842, 1 (8.9 mm), 54°6.0′ N, 

166°14.1′ W, 0–76 m depth, bongo net (haul 6), 28 April 

1994; UW 144859, 1 (23.5 mm), 62°21.6′ N, 179°36.0′

W, 0 m depth, neuston net, 19 August 1994; UW 144860, 

1 (9.9 mm), 55°15.1′ N, 165°13.9′ W, 0–105 m depth, 

bongo net, 16 May 1995; UW 140705, 1 (25.2 mm), 

57°59.7′ N, 167°1.1′ W, 0–55 m depth, Methot net, 22 

July 1995; UW 140706, 5 (23.1–24.4 mm), 58°30.5′ N, 

166°0.1′ W, 0–35 m depth, Methot net, 23 July 1995; 

UW 144845, 1 (12.1 mm), 57°20.9′ N, 170°8.3′ W, 0–53 

m depth, bongo net, 5 July 1997; UW 144846, 1 (22.3 

mm), 57°18.5′ N, 170°9.5′ W, 0–42 m depth, bongo net, 

5 July 1997; UW 144847, 2 (15.0–22.0 mm), 57°3.0′ N, 

170°16.9′ W, 0–45 m depth, bongo net, 6 July 1997; UW 

144844, 1 (20.2 mm), 56°37.8′ N, 169°34.4′ W, 0–40 m 

depth, Methot net, 10 July 1997; UW 144843, 1 (21.0 

mm), 56°42.6′ N, 169°35.9′ W, 25–64 m depth, Tucker trawl, 10 July 1997. Gulf of Alaska: UW 144817, 1 

(24.2 mm), 56°47.5′ N, 153°40.1′ W, 0 m depth, neuston 

net, 24 June 1978; UW 144818, 1 (23.0 mm), 56°33.3′

N, 154°51.2′ W, 0–18 m depth, bongo net, 25 June 1978; 

UW 144819, 1 (28.8 mm), 57°16.6′ N, 151°17.8′ W, 0 m 

depth, neuston net, 26 June 1978; UW 140857, 2 (24.4–

25.4 mm), 56°42.1′ N, 153°33.6′ W, 0 m depth, neuston 

net, 29 June 1978; UW 140827, 1 (25.0 mm), 56°42.2′

N, 153°33.1′ W, 0 m depth, neuston net, 29 June 1978; 

UW 144816, 1 (26.0 mm), 56°25.4′ N, 152°37.4′ W, 0 

m depth, neuston net, 1 July 1978; UW 144820, 1 (9.0 

mm), 57°43.3′ N, 154°47.0′ W, 0–103 m depth, bongo 

net, 10 May 1985; UW 144836, 1 (18.5 mm), 56°18.1′

N, 156°16.7′ W, 0–102 m depth, Tucker trawl, 4 June 

1988; UW 67888, 1 (9.3 mm), 56°46.7′ N, 154°28.1′

W, 0–63 m depth, bongo net, 11 May 1989; UW 49815, 

1 (35.0 mm), 60°39.4′ N, 147°23.2′ W, 0–30 m depth, 

beam trawl, 21 September 1989; UW 62383, 1 (10.4 

mm), 57°28.6′ N, 154°42.4′ W, 0–59 m depth, bongo 

net, 9 May 1990; UW 62705, 1 (10.1 mm), 59°2.2′

N, 152°18.1′ W, 0–100 m depth, bongo net, 15 May 

1990; UW 72426, 1 (18.0 mm), 57°31.9′ N, 155°28.5′

W, 0–105 m depth, 4 June 1990; UW 144830, 1 (9.0 

mm), 57°16.8′ N, 155°55.5′ W, 0–100 m depth, bongo 

net, 8 May 1991; UW 144837, 1 (10.0 mm), 56°16.4′

N, 158°4.9′ W, 0–64 m depth, bongo net, 22 May 1991; 

UW 144838, 1 (12.9 mm), 55°46.7′ N, 156°17.0′ W, 

0–101 m depth, bongo net, 23 May 1991; UW 144831, 

1 (7.9 mm), 57°9.2′ N, 156°9.1′ W, 0–77 m depth, bongo 

net, 8 May 1992; UW 144832, 1 (8.4 mm), 56°49.3′ N, 

154°56.1′ W, 0–55 m depth, bongo net, 8 May 1992; 

UW 144833, 1 (8.1 mm), 56°56.0′ N, 157°3.9′ W, 40–

58 m depth, MOCNESS, 12 May 1992; UW 144834, 

1 (11.4 mm), 57°1.2′ N, 156°14.6′ W, 0–100 m depth, 

bongo net, 14 May 1992; UW 144839, 1 (13.6 mm), 

56°55.3′ N, 155°25.2′ W, 0–99 m depth, bongo net, 26 

May 1992; UW 144840, 2 (9.7–9.8 mm), 57°26.8′ N, 

154°47.6′ W, 0–101 m depth, bongo net, 11 May 1993; 

UW 144850, 1 (7.7 mm), 57°8.4′ N, 156°5.7′ W, 0–30 m 

depth, Tucker trawl, 9 May 1994; UW 144849, 1 (10.3 

mm), 57°8.7′ N, 156°0.6′ W, 0–57 m depth, Tucker trawl, 

10 May 1994; UW 144861, 1 (13.5 mm), 57°16.7′ N, 

155°55.2′ W, 0–101 m depth, bongo net, 28 May 1995; 

UW 144857, 1 (9.6 mm), 58°2.9′ N, 153°43.2′ W, 0–190 

m depth, bongo net, 3 May 1996; UW 144856, 1 (8.6 

mm), 56°34.7′ N, 156°54.8′ W, 0–120 m depth, 9 May 

1996; UW 144858, 1 (12.4 mm), 55°31.7′ N, 160°4.7′

W, 0–101 m depth, bongo net, 31 May 1996; UW 

144862, 1 (17.0 mm), 57°21.7′ N, 155°40.1′ W, 0–101 

m depth, bongo net, 29 May 1997; UW 144823, 4 (6.4–

7.6 mm), 55°33.7′ N, 161°31.4′ W, 0–29 m depth, bongo 

net, 24 May 1999; UW 144822, 1 (6.5 mm), 55°21.8′ N, 

161°26.4′ W, 0–49 m depth, bongo net, 24 May 1999; 

UW 144827, 2 (7.2–8.2 mm), 55°21.8′ N, 161°26.4′ W, 

0–40 m depth, bongo net, 24 May 1999; UW 144851, 

1 (10.5 mm), 56°10.1′ N, 156°28.2′ W, 0–101 m depth, 

bongo net, 28 May 1999; UW 144824, 1 (9.7 mm), 

56°38.8′ N, 156°4.2′ W, 0–100 m depth, bongo net, 

30 May 1999; UW 144825, 1 (10.6 mm), 57°3.1′ N, 

156°24.5′ W, 0–101 m depth, bongo net, 31 May 1999; 

UW 144852, 1 (11.5 mm), 57°24.0′ N, 156°17.2′ W, 

0–21 m depth, bongo net, 31 May 1999; UW 144853, 

4 (10.2–12.0 mm), 57°23.3′ N, 156°20.4′ W, 0–35 m 

depth, bongo net, 31 May 1999; UW 144826, 1 (15.4 

mm), 57°32.2′ N, 152°2.2′ W, 0–65 m depth, bongo net, 

2 June 1999; UW 144828, 7 (14.8–18.0 mm), 57°43.0′

N, 152°20.2′ W, 0–102 m depth, bongo net, 5 June 1999; 

UW 144854, 1 (19.5 mm), 57°24.1′ N, 156°14.1′ W, 

0–30 m depth, bongo net, 1 June 2000; UW 144855, 1 

(21.5 mm), 57°43.5′ N, 152°19.0′ W, 0–100 m depth, 

bongo net, 4 June 2000; UW 140831, 1 (12.0 mm), 

59°31.0′ N, 149°48.8′ W, 0–194 m depth, bongo net, 

15 May 2001; UW 144835, 1 (13.1 mm), 55°17.9′ N, 

159°36.8′ W, 0–100 m depth, bongo net, 26 May 2002; 
87

UW 140707, 1 (7.9 mm), 60°32.4′ N, 147°48.0′ W, 0–11 

m depth, MOCNESS, 3 May 2003; UW 140708, 1 (10.7 

mm), 60°32.1′ N, 147°48.2′ W, 10–20 m depth, MOCNESS, 2 June 2003. Location, depth, gear, and date of 

collection unknown for the following specimens: UW 

145892, 1 (16.4 mm); UW 145893, 1 (18.9 mm). 

Bryozoichthys lysimus. 82 specimens (6.9–49.0 mm) 

examined. Bering Sea: UW 144927, 3 (13.8–16.5 mm), 

56°17.4′ N, 171°20.4′ W, depth unknown, bongo net, 

6 May 1976; UW 144929, 1 (17.3 mm), 54°12.6′ N, 

166°26.4′ W, 0–233 m depth, bongo net, 16 April 1977; 

UW 144932, 1 (20.5 mm), 55°4.2′ N, 165°9.0′ W, 0 m 

depth, neuston net, 18 April 1977; UW 145158, 1 (21.1 

mm), 55°9.0′ N, 168°18.0′ W, 0–224 m depth, bongo 

net, 22 April 1977; UW 144933, 1 (21.4 mm), 56°31.8′

N, 166°18.0′ W, 0 m depth, neuston net, 23 April 1977; 

UW 144928, 1 (21.9 mm), 55°13.8′ N, 165°27.6′ W, 

0–123 m depth, bongo net, 24 April 1977; UW 144931, 

4 (20.6–24.7 mm), 55°22.2′ N, 166°9.0′ W, 0 m depth, 

neuston net, 24 April 1977; UW 144934, 6 (19.6–23.8 

mm), 54°22.2′ N, 166°7.8′ W, 0 m depth, neuston net, 

25 April 1977; UW 144930, 1 (22.5 mm), 54°7.8′ N, 

166°8.4′ W, 0 m depth, neuston net, 26 April 1977; UW 

144938, 5 (21.2–24.1 mm), 53°7.8′ N, 169°14.4′ W, 0 m 

depth, neuston net, 3 May 1977; UW 144937, 3 (20.5–

24.5 mm), 54°31.2′ N, 166°25.8′ W, 0 m depth, neuston 

net, 5 May 1977; UW 144935, 1 (18.2 mm), 55°31.2′ N, 

163°27.0′ W, 0–85 m depth, bongo net, 8 May 1977; UW 

144936, 1 (29.0 mm original measurement when illustrated; remeasured at 26.8 mm for this study), 56°27.0′

N, 171°17.4′ W, 0 m depth, neuston net, 12 May 1977; 

UW 144939, 1 (13.2 mm), 54° N, 167°8.8′ W, 0 m depth, 

neuston net, 3 March 1978; UW 65424, 1 (21.6 mm), 

54°49.8′ N, 168°5.6′ W, 0 m depth, neuston net, 1 June 

1979; UW 144917, 1 (32.4 mm), 56°3.5′ N, 166°33.9′

W, 0 m depth, neuston net, 2 June 1979; UW 144911, 

1 (20.3 mm), 54°4.9′ N, 171°26.2′ W, 0 m depth, neuston net, 7 June 1979; UW 144912, 3 (27.9–29.6 mm), 

52°45.6′ N, 174°48.0′ W, 0 m depth, neuston net, 16 

June 1979; UW 144913, 3 (32.6–35.6 mm), 54°45.8′

N, 168°51.6′ W, 0 m depth, neuston net, 17 June 1979; 

UW 144914, 1 (34.2 mm), 57°4.4′ N, 167°58.8′ W, 0 m 

depth, neuston net, 24 June 1979; UW 144915, 1 (31.9 

mm), 56°23.0′ N, 170°22.0′ W, 0 m depth, neuston net, 

26 June 1979; UW 144916, 2 (30.8–31.5 mm), 52°30.8′

N, 178°36.0′ W, 0 m depth, neuston net, 2 July 1979; 

UW 144918, 1 (12.0 mm), 54°11.0′ N, 166°56.8′ W, 

0–399 m depth, bongo net, 15 April 1993; UW 144919, 

1 (10.4 mm), 54°5.9′ N, 167°54.0′ W, 0–399 m depth, 

bongo net, 18 April 1993; UW 144922, 2 (19.0–21.5 

mm), 54°47.5′ N, 167°30.0′ W, 0–400 m depth, bongo net, 22 April 1993; UW 144920, 3 (6.9–8.0 mm), 

54°44.8′ N, 167°52.7′ W, 0–400 m depth, bongo net, 

22 April 1993; UW 144923, 1 (17.0 mm), 54°46.0′ N, 

168°1.6′ W, 0–400 m depth, bongo net, 27 April 1993; 

UW 144926, 1 (14.5 mm), 54°28.2′ N, 166°2.8′ W, 

0–401 m depth, bongo net, 23 April 1995; UW 144940, 

1 (49.0 mm), 51°56.2′ N, 176°50.3′ W, 0–87 m depth, 

gear unknown, 17 June 2001; UW 144909, 1 (25.0 mm), 

54°27.1′ N, 166°59.9′ W, 0 m depth, neuston net, 16 

May 2002. Gulf of Alaska: UW 69706, 1 (17.4 mm), 

56°39.5′ N, 155°25.1′ W, 0 m depth, 12 April 1978; 

UW 69324, 4 (12.0–13.7 mm), 58°4.0′ N, 151°41.5′ W, 

0–128 m depth, bongo net, 29 May 1978; UW 33190, 1 

(10.2 mm), 58°10.0′ N, 152°14.0′ W, depth unknown, 

bongo net, 6–16 March 1979; UW 144902, 1 (12.9 

mm), 57°3.0′ N, 155°53.0′ W, 0 m depth, neuston net, 

19 March 1981; UW 60160, 1 (17.1 mm), 57°10.9′ N, 

155°8.0′ W, 0–208 m depth, bongo net, 1 April 1981; 

UW 64990, 1 (22.3 mm), 56°41.0′ N, 156°26.0′ W, 0 

m depth, neuston net, 21 April 1981; UW 145156, 1 

(29.4 mm), 55°14.1′ N, 159°2.0′ W, 0 m depth, neuston 

net, 29 May 1982; UW 144904, 1 (27.1 mm), 54°54.1′

N, 158°39.0′ W, 0 m depth, neuston net, 29 May 1982; 

UW 144897, 1 (28.9 mm), 55°59.5′ N, 157°21.0′ W, 0 m 

depth, neuston net, 23 May 1983; UW 144903, 1 (23.4 

mm), 57°11.5′ N, 154°44.0′ W, 0 m depth, neuston net, 

20 April 1984; UW 55919, 3 (14.8–18.0 mm), 58°44.2′

N, 152°51.0′ W, 0 m depth, neuston net, 29 March 1985; 

UW 64569, 1 (29.3 mm), 54°8.0′ N, 164°14.0′ W, 0 m 

depth, neuston net, 6 June 1985; UW 64631, 5 (28.4–

30.4 mm), 53°34.8′ N, 165°18.7′ W, 0 m depth, neuston 

net, 7 June 1985; UW 64723, 1 (25.3 mm), 53°0.5′ N, 

167°59.0′ W, 0 m depth, 8 June 1985; UW 144898, 2 

(15.4–16.2 mm), 53°59.3′ N, 162°10.0′ W, 0 m depth, 

neuston net, 18 April 1986; UW 39750, 1 (14.1 mm), 

53°40.0′ N, 165°3.0′ W, 0–171 m depth, bongo net, 

19 April 1986; UW 144900, 1 (30.4 mm), 56°24.0′ N, 

153°23.4′ W, 0–60 m depth, Tucker trawl, 27 May 1986; 

UW 144925, 1 (32.0 mm), 55°46.6′ N, 156°16.7′ W, 

0–100 m depth, bongo net, 28 May 1994. 

Bryozoichthys marjorius. 90 specimens (7.7–53.0 mm) 

examined. Bering Sea: UW 144953, 2 (18.8–19.7 mm), 

54°22.2′ N, 166°8.3′ W, 0 m depth, neuston net, 25 

April 1977; UW 144983, 2 (31.0–31.6 mm), 54°7.8′ N, 

166°9.3′ W, 0 m depth, neuston net, 26 April 1977; UW 

144954, 3 (18.7–20.5 mm), 53°8.0′ N, 169°14.4′ W, 0 m 

depth, neuston net, 3 May 1977; UW 144969, 2 (23.3–

25.8 mm), 54°22.2′ N, 166°8.4′ W, 0 m depth, neuston 

net, 5 May 1977; UW 144977, 1 (25.8 mm), 55°26.9′ N, 

168°17.7′ W, 0 m depth, neuston net, 10 May 1977; UW 

144959, 1 (20.6 mm), 55°26.6′ N, 171°18.8′ W, 0–219 m 

depth, bongo net, 13 May 1977; UW 144974, 3 (25.1–

28.3 mm), 52°45.6′ N, 174°48.0′ W, 0 m depth, neuston 

net, 16 June 1979; UW 66115, 1 (31.9 mm), 58°16.6′

N, 172°48.6′ W, 0 m depth, neuston net, 17 June 1979; 

UW 144985, 2 (31.5–35.9 mm), 57°45.8′ N, 168°51.6′

W, 0 m depth, neuston net, 17 June 1979; UW 65595, 1 

(32.9 mm), 55°38.9′ N, 167°3.6′ W, 0 m depth, neuston 

net, 18 June 1979; UW 65607, 1 (26.6 mm), 55°47.6′ N, 
88 Professional Paper NMFS 15 

166°54.2′ W, 0 m depth, neuston net, 19 June 1979; UW 

144981, 2 (30.3–35.8 mm), 55°28.8′ N, 167°46.2′ W, 

0 m depth, neuston net, 20 June 1979; UW 144995, 1 

(36.2 mm), 56°23.0′ N, 170°22.0′ W, 0 m depth, neuston 

net, 26 June 1979; UW 65874, 1 (38.0 mm), 56°29.8′ N, 

171°55.7′ W, 0 m depth, neuston net, 28 June 1979; UW 

65956, 2 (36.8–38.0 mm), 56°27.5′ N, 172°9.1′ W, 0 m 

depth, neuston net, 9 July 1979; UW 65974, 1 (36.0 mm), 

56°59.6′ N, 173°33.9′ W, 0 m depth, neuston net, 9 July 

1979; UW 66011, 1 (34.6 mm), 57°43.2′ N, 174°6.1′

W, 0 m depth, neuston net, 11 July 1979; UW 66153, 1 

(34.9 mm), 59°19.6′ N, 174°11.0′ W, 0 m depth, neuston 

net, 19 July 1979; UW 66179, 1 (32.1 mm), 59°50.1′ N, 

175°26.3′ W, 0 m depth, neuston net, 20 July 1979; UW 

144964, 1 (22.5 mm), 54°59.4′ N, 168°12.6′ W, 0–202 

m depth, bongo net, 21 April 1994; UW 144998, 1 (7.7 

mm), 54°20.33′ N, 165°25.8′ W, 0–144 m depth, bongo 

net, 4 May 1995; UW 144963, 1 (22.5 mm), 54°28.1′

N, 166°2.2′ W, 0 m depth, neuston net, 13 May 2002; 

UW 144973, 1 (25.0 mm), 54°46.1′ N, 167°47.3′ W, 0 m 

depth, neuston net, 20 May 2002; UW 111242, 1 (53.0 

mm), 52°12.8′ N, 174°56.4′ W, 206 m depth, benthic 

bag, 13 June 2002; UW 144943, 1 (13.7 mm), 56°32.3′

N, 151°46.1′ W, 0–602 m depth, bongo net, 15 February 2003. Gulf of Alaska: UW 144942, 1 (11.4 mm), 

56°28.0′ N, 156°12.0′ W, 0–105 m depth, bongo net, 

29 April 1972; UW 69029, 1 (22.8 mm), 56°15.5′ N, 

153°21.2′ W, 0 m depth, neuston net, 14 April 1978; UW 

60255, 1 (14.5 mm), 56°29.5′ N, 156°14.1′ W, 0–212 

m depth, bongo net, 4 April 1981; UW 57125, 1 (15.8 

mm), 55°27.2′ N, 159°18.0′ W, 0 m depth, neuston net, 

23 May 1981; UW 36366, 2 (30.5–32.3 mm), 55°45.0′

N, 157°21.0′ W, 0 m depth, neuston net, 23 May 1983; 

UW 144991, 1 (34.4 mm), 55°15.3′ N, 157°49.7′ W, 0 

m depth, neuston net, 26 May 1983; UW 57459, 1 (22.6 

mm), 57°53.0′ N, 148°39.0′ W, 0 m depth, neuston net, 

14 April 1984; UW 144967, 3 (22.7–23.6 mm), 57°11.5′

N, 154°44.0′ W, 0 m depth, neuston net, 20 April 1984; 

UW 83134, 1 (21.4 mm), 55°34.0′ N, 156°35.0′ W, 0 m 

depth, neuston net, 1 May 1984; UW 83138, 2 (21.9–

22.8 mm), 55°28.0′ N, 158°58.0′ W, 0 m depth, neuston net, 5 May 1984; UW 55946, 2 (13.8–17.1 mm), 

58°34.5′ N, 153°13.0′ W, 0 m depth, neuston net, 29 

March 1985; UW 144947, 8 (15.9–18.2 mm), 58°44.2′

N, 152°51.0′ W, 0 m depth, neuston net, 29 March 1985; 

UW 144952, 1 (18.0 mm), 58°18.8′ N, 153°26.0′ W, 0 m 

depth, neuston net, 30 March 1985; UW 56216, 2 (18.0–

21.6 mm), 57°12.2′ N, 155°26.0′ W, 0 m depth, neuston 

net, 4 April 1985; UW 56255, 1 (17.9 mm), 56°50.2′ N, 

154°48.0′ W, 0–40 m depth, bongo net, 5 April 1985; 

UW 56581, 1 (19.8 mm), 57°38.5′ N, 150°32.0′ W, 0 m 

depth, neuston net, 17 April 1985; UW 63509, 5 (24.3–

33.0 mm), 57°34.0′ N, 155°20.0′ W, 0 m depth, neuston 

net, 23 May 1985; UW 63908, 1 (34.0 mm), 57°18.4′

N, 155°50.9′ W, 0 m depth, neuston net, 27 May 1985; 

UW 63918, 1 (25.4 mm), 56°29.5′ N, 155°30.0′ W, 0 m 

depth, neuston net, 27 May 1985; UW 64168, 1 (29.6 

mm), 55°35.5′ N, 157°15.2′ W, 0 m depth, neuston net, 

30 May 1985; UW 64178, 1 (24.9 mm), 55°48.4′ N, 

157°32.9′ W, 0 m depth, neuston net, 30 May 1985; UW 

64277, 2 (19.7–27.3 mm), 55°39.4′ N, 158°35.7′ W, 0 

m depth, neuston net, 1 June 1985; UW 144975, 1 (25.2 

mm), 53°34.8′ N, 165°18.7′ W, 0 m depth, neuston net, 

7 June 1985; UW 82725, 2 (19.3–20.5 mm), 57°20.0′ N, 

150°56.2′ W, 0 m depth, neuston net, 10 April 1986; UW 

60536, 1 (17.7 mm), 57°16.1′ N, 155°37.1′ W, 0–270 

m depth, bongo net, 4 May 1986; UW 144945, 1 (14.9 

mm), 57°18.7′ N, 155°16.8′ W, 0–238 m depth, bongo 

net, 7 April 1987; UW 35711, 1 (23.3 mm), 55°40.0′ N, 

157° W, 0–66 m depth, bongo net, 23 April 1987; UW 

52038, 1 (10.9 mm), 56°57.1′ N, 155°57.2′ W, 0–267 m 

depth, bongo net, 10 April 1988; UW 144961, 1 (22.0 

mm), 56°44.7′ N, 156°46.0′ W, 0–104 m depth, Tucker 

trawl, 1 June 1989; UW 144958, 1 (20.4 mm), 56°1.8′

N, 156°49.4′ W, 0–54 m depth, Tucker trawl, 4 June 

1989; UW 52670, 1 (16.7 mm), 57°32.8′ N, 154°49.9′

W, 0–226 m depth, bongo net, 10 April 1990; UW 

144970, 1 (23.9 mm), 56°32.2′ N, 156°19.6′ W, 0–107 

m depth, bongo net, 8 May 1991; UW 144984, 1 (31.5 

mm), 57°40.7′ N, 154°46.5′ W, 0–100 m depth, bongo 

net, 11 May 1993; UW 144989, 1 (33.0 mm), 56°32.9′

N, 155°51.2′ W, 0–101 m depth, bongo net, 31 May 

1993; UW 144979, 1 (28.0 mm), 57°43.1′ N, 155°15.9′

W, 0–145 m depth, bongo net, 28 May 1995. Location, 

depth, gear, and date of collection unknown for the following specimen: UW 145874, 1 (40.0 mm). 

Chirolophis decoratus. 70 specimens (8.7–42.5 mm) 

examined. Bering Sea: UW 145014, 1 (22.5 mm), 

55°38.7′ N, 163°9.5′ W, 0 m depth, neuston net, 15 May 

2002; UW 145160, 1 (32.0 mm), 54°46.0′ N, 167°47.1′

W, 0 m depth, neuston net, 11 May 2006; UW 145161, 1 

(22.0 mm), 54°35.1′ N, 165°39.1′ W, 0 m depth, neuston 

net, 13 May 2006. Gulf of Alaska: UW 110397, 1 (42.5 

mm), 60°20.2′ N, 146°37.3′ W, 45 m depth, beam trawl, 

date unknown; UW 107634, 3 (11.1–14.9 mm), 56°44.8′

N, 154°17.9′ W, 0–14 m depth, Tucker trawl, 11 April 

1978; UW 69705, 1 (21.8 mm), 56°39.5′ N, 155°25.1′

W, 0 m depth, neuston net, 12 April 1978; UW 145023, 1 

(28.2 mm), 57°12.1′ N, 152°44.5′ W, 0 m depth, neuston 

net, 24 June 1978; UW 64891, 1 (27.0 mm), 55°57.8′

N, 153°31.0′ W, 0 m depth, neuston net, 19 April 1981; 

UW 37072, 1 (14.7 mm), 57°22.2′ N, 151°56.0′ W, 0–61 

m depth, bongo net, 5 April 1982; UW 37157, 1 (17.4 

mm), 57°30.5′ N, 155°31.0′ W, 0 m depth, neuston 

net, 8 April 1982; UW 37175, 1 (17.6 mm), 56°59.0′

N, 155°50.0′ W, 0 m depth, neuston net, 8 April 1982; 

UW 59889, 1 (23.0 mm), 54°52.2′ N, 157°32.0′ W, 0 m 

depth, neuston net, 28 May 1982; UW 145016, 1 (24.9 

mm), 55°14.1′ N, 159°2.0′ W, 0 m depth, neuston net, 

29 May 1982; UW 145018, 1 (26.6 mm), 54°54.1′ N, 

158°39.0′ W, 0 m depth, neuston net, 29 May 1982; 
89

UW 59978, 1 (31.6 mm), 55°4.2′ N, 158°51.0′ W, 0 m 

depth, neuston net, 29 May 1982; UW 60031, 1 (30.6 

mm), 54°32.2′ N, 159°16.0′ W, 0 m depth, neuston net, 

30 May 1982; UW 60023, 1 (30.8 mm), 54°32.2′ N, 

159°16.0′ W, 0–82 m depth, bongo net, 30 May 1982; 

UW 36026, 1 (27.2 mm), 57°0.5′ N, 155°32.8′ W, 0 m 

depth, neuston net, 22 May 1983; UW 36198, 1 (31.1 

mm), 56°14.8′ N, 156°26.6′ W, 0 m depth, neuston 

net, 24 May 1983; UW 145013, 1 (21.9 mm), 57°7.0′

N, 155°56.0′ W, 0 m depth, neuston net, 4 April 1985; 

UW 62916, 3 (29.5–31.2 mm), 59°2.9′ N, 148°30.0′ W, 

0 m depth, neuston net, 18 May 1985; UW 145026, 1 

(29.2 mm), 57°43.9′ N, 155°1.4′ W, 0 m depth, neuston 

net, 23 May 1985; UW 63937, 1 (15.9 mm), 56°52.0′

N, 155°28.0′ W, 0–202 m depth, bongo net, 27 May 

1985; UW 63987, 1 (27.8 mm), 56°20.8′ N, 156°29.0′

W, 0 m depth, neuston net, 28 May 1985; UW 64156, 1 

(28.6 mm), 55°24.5′ N, 156°55.1′ W, 0 m depth, neuston 

net, 30 May 1985; UW 64269, 1 (26.3 mm), 55°27.5′

N, 158°16.6′ W, 0 m depth, neuston net, 1 June 1985; 

UW 64343, 1 (30.3 mm), 54°12.0′ N, 158°29.0′ W, 0 

m depth, neuston net, 2 June 1985; UW 60443, 1 (21.0 

mm), 57°29.0′ N, 154°47.8′ W, 0–62 m depth, bongo 

net, 3 May 1986; UW 60667, 1 (11.9 mm), 56°47.1′ N, 

155°54.6′ W, 0–294 m depth, bongo net, 7 May 1986; 

UW 145030, 1 (30.7 mm), 57°21.6′ N, 151°25.2′ W, 

0–45 m depth, Tucker trawl, 26 May 1986; UW 145019, 

1 (26.6 mm), 56°24.0′ N, 153°23.4′ W, 0–44 m depth, 

Tucker trawl, 27 May 1986; UW 37745, 1 (10.2 mm), 

56°19.7′ N, 154°19.0′ W, 0–80 m depth, bongo net, 

2 April 1988; UW 52204, 1 (21.5 mm), 57°35.0′ N, 

155°21.0′ W, 0–307 m depth, bongo net, 7 April 1989; 

UW 52786, 1 (13.1 mm), 57°15.6′ N, 155°35.1′ W, 

0–278 m depth, bongo net, 11 April 1990; UW 145009, 

1 (19.5 mm), 57°21.4′ N, 155°27.1′ W, 0–259 m depth, 

bongo net, 9 April 1993; UW 145025, 1 (28.5 mm), 

55°27.9′ N, 158°39.8′ W, 0–100 m depth, bongo net, 26 

May 1994. British Columbia, Canada: UW 145169, 9 

(8.7–9.7 mm), 49°21.6′ N, 123°29.3′ W, reared, 7 February 1979; UW 145170, 5 (10.9–11.5 mm), 49°21.6′

N, 123°29.3′ W, reared, 14 February 1979. Washington 

coast: UW 145168, 1 (21.5 mm), 48°16.4′ N, 123°33.6′

W, 0–50 m depth, bongo net, 24 February 1976; UW 

145000, 1 (10.1 mm), 48°25.4′ N, 124°41.7′ W, depth 

unknown, bongo net, 12 January 1977; UW 51382, 1 

(17.2 mm), 43°19.9′ N, 124°30.4′ W, 0–75 m depth, 

bongo net, 21 January 1987; UW 144999, 2 (10.0–10.5 

mm), 41°59.1′ N, 124°22.3′ W, 0–56 m depth, bongo 

net, 23 January 1987. Location, depth, gear, and date 

of collection unknown for the following specimens: UW 

145164, 1 (10.3 mm); UW 145166, 2 (10.2–11.6 mm); 

UW 145165, 1 (11.1 mm); UW 145162, 2 (21.0–21.9 

mm); UW 145163, 1 (17.9 mm); UW 145875, 1 (13.6 

mm); UW 145876, 1 (14.3 mm); UW 145877, 1 (14.8 

mm); UW 145878, 1 (14.9 mm). 

Chirolophis nugator. 76 specimens (8.7–25.0 mm) examined. Bering Sea: UW 145050, 1 (15.5 mm), 54°26.6′

N, 164°58.8′ W, 0–41 m depth, bongo net, 11 May 1997; 

UW 145048, 1 (18.5 mm), 56°27.4′ N, 169°28.4′ W, 

0–96 m depth, Methot net, 12 July 1997. Gulf of Alaska: 

UW 32896, 2 (10.1–10.4 mm), 58°10.0′ N, 152°14.0′

W, 0–10 m depth, Tucker trawl, 21 April 1978; UW 

145045, 1 (20.1 mm), 57°12.1′ N, 152°44.5′ W, 0 m 

depth, neuston net, 24 June 1978; UW 50335, 1 (10.4 

mm), 57°43.0′ N, 155°6.6′ W, 0–176 m depth, bongo 

net, 15 March 1981; UW 56975, 1 (19.3 mm), 56°30.0′

N, 155°14.0′ W, 0–23 m depth, bongo net, 18 March 

1981; UW 145044, 2 (14.0–17.1 mm), 56°20.4′ N, 

152°2.4′ W, 0–58 m depth, Tucker trawl, 27 May 1986; 

UW 35681, 1 (12.3 mm), 55°40.0′ N, 155°45.0′ W, 

0–78 m depth, bongo net, 22 April 1987; UW 145038, 

1 (10.4 mm), 56°24.3′ N, 154°7.9′ W, 0–40 m depth, 

bongo net, 20 April 1991; UW 145049, 1 (10.9 mm), 

56°27.5′ N, 157°30.3′ W, 0–91 m depth, bongo net, 

22 April 1991; UW 145036, 1 (9.6 mm), 57°16.8′ N, 

155°55.5′ W, 0–100 m depth, bongo net, 8 May 1991; 

UW 145052, 1 (16.1 mm), 54°55.0′ N, 161°31.4′ W, 

0–121 m depth, bongo net, 27 May 2000; UW 145051, 

1 (19.0 mm), 55°49.6′ N, 155°49.0′ W, 0–49 m depth, 

bongo net, 30 May 2000; UW 145047, 1 (10.4 mm), 

57°18.0′ N, 151°17.9′ W, 0 m depth, neuston net, 17 

February 2003; UW 145046, 1 (25.0 mm), 56°2.6′ N, 

157°6.3′ W, 0–100 m depth, bongo net, 29 May 2003. 

British Columbia, Canada: UW 145033, 2 (9.2–10.5 

mm), 48°32.3′ N, 124°42.2′ W, 0–50 m depth, bongo 

net, 24 February 1976; UW 145062, 3 (8.7–9.7 mm), 

49°21.6′ N, 123°29.3′ W, reared, 6 March 1984. Puget 

Sound, Washington: UW 145059, 1 (24.8 mm), 47°33.7′

N, 122°32.1′ W, 0 m depth, dip net, 12 April 1988; UW 

145060, 3 (20.1–20.8 mm), 47°33.7′ N, 122°32.1′ W, 0 

m depth, dip net, 19 April 1988; UW 145040, 2 (19.5–

22.1 mm), 47°33.7′ N, 122°32.1′ W, 0 m depth, dip net, 

6 April 1989; UW 145061, 2 (20.8–22.3 mm), 47°33.7′

N, 122°32.1′ W, 0 m depth, dip net, 11 April 1989; UW 

145056, 2 (16.8–20.1 mm), 47°33.7′ N, 122°32.1′ W, 0 

m depth, dip net, 6 April 1990; UW 145057, 2 (18.0–

23.1 mm), 47°33.7′ N, 122°32.1′ W, 0 m depth, dip net, 

4 May 1990; UW 145058, 8 (19.2–23.7 mm), 47°33.7′

N, 122°32.1′ W, 0 m depth, dip net, 20 April 1991; UW 

145053, 6 (10.6–22.5 mm), 47°33.7′ N, 122°32.1′ W, 

0 m depth, dip net, 26 February 1992; UW 145054, 13 

(13.1–18.7 mm), 47°33.7′ N, 122°32.1′ W, 0 m depth, 

dip net, 12 March 1992; UW 145055, 4 (14.0–20.3 

mm), 47°33.7′ N, 122°32.1′ W, 0 m depth, dip net, 28 

March 1992; UW 145039, 1 (11.2 mm), 48°19.4′ N, 

123° W, 0 m depth, dip net, 8 April 1993. Oregon coast: 

UW 55217, 1 (11.2 mm), 45°18.0′ N, 124°6.0′ W, 0–88 

m depth, bongo net, 22 April 1987. California coast: UW 

51287, 1 (9.2 mm), 44°41.2′ N, 124°10.4′ W, 0–60 m 

depth, bongo net, 19 January 1987; UW 145034, 2 (9.3–

9.9 mm), 41°59.1′ N, 124°22.3′ W, 0–56 m depth, bongo 
90 Professional Paper NMFS 15 

net, 23 January 1987. Location, depth, gear, and date 

of collection unknown for the following specimens: UW 

145170, 4 (9.8–12.9 mm); UW 145879, 1 (23.6 mm); 

UW 145880, 1 (24.1 mm).

Gymnoclinus cristulatus. 8 specimens (18.8–26.8 

mm) examined (depths and gear unknown). Sea of Japan: HUMZ-L 02569, 3 (23.2–26.8 mm), 41°56.0′ N, 

140°57.0′ E, 30 April 1983; HUMZ-L 03113, 5 (18.8–

22.3 mm), 41°56.0′ N, 140°57.0′ E, 26 April 1984.

Lumpenella longirostris. 126 specimens (8.0–64.0 mm) 

examined. Bering Sea: UW 65743, 2 (26.2–32.5 mm), 

56°45.3′ N, 167°29.6′ W, 0–82 m depth, neuston net, 

23 June 1979. Gulf of Alaska: UW 27447, 1 (49.0 mm), 

57°8.2′ N, 153°21.1′ W, 0–118 m depth, 1.5-m net, 4 

March 1978; UW 32916, 3 (8.0–8.5 mm), 58°10.0′

N, 152°14.0′ W, 0–70 m depth, gear unknown, 3 May 

1978; UW 145086, 1 (32.5 mm), 57°12.1′ N, 152°44.5′

W, 0–121 m depth, bongo net, 24 June 1978; UW 

145093, 1 (40.0 mm), 56°42.7′ N, 153°33.4′ W, 28–71 

m depth, Tucker trawl, 29 June 1978; UW 145081, 1 

(32.0 mm), 56°37.4′ N, 153°26.8′ W, 0–56 m depth, 

bongo net, 30 June 1978; UW 145082, 1 (32.7 mm), 

56°37.5′ N, 153°26.8′ W, 29–100 m depth, Tucker trawl, 

30 June 1978; UW 145087, 1 (38.0 mm), 56°37.5′ N, 

153°26.8′ W, 0–100 m depth, 30 June 1978; UW 66781, 

1 (9.8 mm), 57°34.8′ N, 155°17.5′ W, 0–200 m depth, 

bongo net, 28 April 1981; UW 66786, 2 (9.8–12.2 mm), 

57°31.3′ N, 155°27.5′ W, 0–216 m depth, bongo net, 28 

April 1981; UW 66838, 4 (12.7–13.3 mm), 57°0.8′ N, 

156°19.7′ W, 0–72 m depth, bongo net, 29 April 1981; 

UW 66879, 1 (14.8 mm), 56°49.9′ N, 155°57.1′ W, 

0–218 m depth, bongo net, 29 April 1981; UW 66907, 3 

(8.1–10.5 mm), 57°18.5′ N, 155°21.8′ W, 0–217 m depth, 

bongo net, 30 April 1981; UW 66916, 5 (10.8–12.2 mm), 

57°24.2′ N, 154°56.0′ W, 0–202 m depth, bongo net, 30 

April 1981; UW 66922, 3 (9.2–10.4 mm), 57°19.3′ N, 

155°5.2′ W, 0–208 m depth, bongo net, 30 April 1981; 

UW 66929, 2 (11.7–14.2 mm), 57°14.4′ N, 155°15.1′

W, 0–208 m depth, 30 April 1981; UW 66971, 1 (15.8 

mm), 56°56.3′ N, 155°32.4′ W, 0–209 m depth, bongo 

net, 1 May 1981; UW 66978, 1 (16.1 mm), 56°49.9′ N, 

155°19.4′ W, 0–207 m depth, bongo net, 1 May 1981; 

UW 70563, 3 (11.7–17.3 mm), 58°9.0′ N, 153°24.3′ W, 

0–208 m depth, bongo net, 20 May 1981; UW 70575, 

1 (13.5 mm), 58°11.5′ N, 153°32.5′ W, 0–166 m depth, 

bongo net, 20 May 1981; UW 70586, 4 (13.1–16.0 mm), 

58°14.5′ N, 153°39.4′ W, 0–166 m depth, bongo net, 20 

May 1981; UW 70645, 5 (14.1–16.5 mm), 57°56.1′ N, 

153°50.5′ W, 0–220 m depth, bongo net, 20 May 1981; 

UW 70652, 1 (14.8 mm), 57°59.5′ N, 153°58.0′ W, 

0–163 m depth, bongo net, 20 May 1981; UW 70665, 

1 (17.1 mm), 58°2.9′ N, 154°6.0′ W, 0–208 m depth, 

bongo net, 20 May 1981; UW 70704, 2 (15.4–15.8 mm), 

57°48.4′ N, 154°31.5′ W, 0–208 m depth, bongo net, 

21 May 1981; UW 70744, 1 (14.9 mm), 57°40.0′ N, 

154°51.3′ W, 0–208 m depth, 21 May 1981; UW 70754, 

4 (20.9–24.6 mm), 57°28.5′ N, 154°45.3′ W, 0–126 m 

depth, bongo net, 21 May 1981; UW 70875, 1 (18.8 

mm), 57°18.7′ N, 155°22.7′ W, 0–208 m depth, bongo 

net, 21 May 1981; UW 70989, 1 (19.9 mm), 56°52.8′ N, 

155°25.2′ W, 0–208 m depth, bongo net, 22 May 1981; 

UW 70995, 2 (19.8–24.9 mm), 56°56.3′ N, 155°25.2′ W, 

0–208 m depth, bongo net, 22 May 1981; UW 68784, 

2 (22.0–27.4 mm), 57°3.0′ N, 154°56.0′ W, 0–93 m 

depth, bongo net, 23 May 1981; UW 71018, 1 (24.5 

mm), 57°4.5′ N, 155°47.6′ W, 0–208 m depth, bongo 

net, 23 May 1981; UW 71027, 1 (25.3 mm), 57°8.0′ N, 

155°54.4′ W, 0–208 m depth, bongo net, 23 May 1981; 

UW 71072, 4 (21.3–23.5 mm), 57°6.9′ N, 156°18.4′ W, 

0–75 m depth, bongo net, 23 May 1981; UW 71095, 

3 (15.7–25.3 mm), 56°54.8′ N, 155°47.6′ W, 0–208 m 

depth, bongo net, 23 May 1981; UW 71145, 1 (23.2 

mm), 56°34.0′ N, 155°45.5′ W, 0–208 m depth, bongo 

net, 23 May 1981; UW 68834, 1 (21.4 mm), 56°54.8′

N, 155°42.0′ W, 0–197 m depth, bongo net, 24 May 

1981; UW 71164, 1 (23.8 mm), 56°45.0′ N, 156°6.2′

W, 0–208 m depth, bongo net, 24 May 1981; UW 

71174, 1 (21.6 mm), 56°52.1′ N, 156°19.5′ W, 0–167 

m depth, bongo net, 24 May 1981; UW 71224, 1 (22.2 

mm), 56°28.2′ N, 155°53.2′ W, 0–208 m depth, bongo 

net, 24 May 1981; UW 68859, 1 (24.9 mm), 56°32.0′ N, 

156°15.0′ W, 0–210 m depth, bongo net, 25 May 1981; 

UW 68867, 2 (25.1–27.2 mm), 56°23.2′ N, 156°6.0′ W, 

0–22 m depth, bongo net, 25 May 1981; UW 36209, 1 

(23.4 mm), 56°15.2′ N, 156°26.1′ W, 0–213 m depth, 

bongo net, 24 May 1983; UW 36257, 1 (17.3 mm), 

56°43.5′ N, 156°53.2′ W, 0–62 m depth, bongo net, 24 

May 1983; UW 36387, 1 (24.6 mm), 56° N, 157°21.0′

W, 0–94 m depth, bongo net, 25 May 1983; UW 36395, 

1 (17.5 mm), 56°15.8′ N, 157°20.6′ W, 0–105 m depth, 

bongo net, 25 May 1983; UW 57781, 1 (9.9 mm), 

56°51.5′ N, 156°8.0′ W, 0–216 m depth, bongo net, 

24 April 1984; UW 63379, 1 (10.9 mm), 58°13.9′ N, 

153°55.0′ W, 0–200 m depth, bongo net, 22 May 1985; 

UW 63420, 1 (10.8 mm), 58°4.1′ N, 153°53.0′ W, 0–185 

m depth, bongo net, 22 May 1985; UW 60600, 1 (9.8 

mm), 56°51.8′ N, 155°46.2′ W, 0–272 m depth, bongo 

net, 5 May 1986; UW 60755, 1 (11.0 mm), 56°22.2′ N, 

156°26.5′ W, 0–218 m depth, bongo net, 13 May 1986; 

UW 67064, 2 (10.3–11.1 mm), 56°54.7′ N, 155°32.6′ W, 

0–275 m depth, bongo net, 20 May 1987; UW 67070, 

1 (12.8 mm), 56°43.0′ N, 155°24.3′ W, 0–198 m depth, 

bongo net, 21 May 1987; UW 67120, 4 (9.6–13.2 mm), 

56°46.4′ N, 155°55.6′ W, 0–303 m depth, bongo net, 

21 May 1987; UW 67138, 1 (10.5 mm), 56°37.0′ N, 

155°33.2′ W, 0–190 m depth, bongo net, 21 May 1987; 

UW 82889, 1 (64.0 mm), 57°37.5′ N, 155°18.7′ W, 

332 m depth, sled trawl, 15 April 1989; UW 94969, 1 

(11.9 mm), 56°50.1′ N, 156°26.0′ W, 0–100 m depth, 

bongo net, 13 May 1992; UW 145094, 1 (19.0 mm), 
91

56°7.8′ N, 157°27.8′ W, 0–101 m depth, bongo net, 

22 May 1992; UW 145085, 1 (20.0 mm), 56°10.5′ N, 

156°2.5′ W, 0–100 m depth, bongo net, 29 May 1996; 

UW 145084, 2 (25.5–26.5 mm), 57°7.2′ N, 156°5.4′ W, 

0–79 m depth, bongo net, 29 May 1997; UW 145089, 

1 (15.9 mm), 56°46.3′ N, 155°51.5′ W, 0–100 m depth, 

bongo net, 30 May 2001; UW 145088, 1 (22.5 mm), 

57°22.1′ N, 155°14.2′ W, 0–100 m depth, bongo net, 

31 May 2001; UW 145092, 1 (19.8 mm), 57°43.2′ N, 

155°15.5′ W, 0–180 m depth, bongo net, 20 May 2002; 

UW 145091, 1 (17.6 mm), 56°24.3′ N, 156°29.4′ W, 

0–56 m depth, bongo net, 29 May 2002; UW 145070, 

1 (8.4 mm), 60°32.1′ N, 147°48.3′ W, 0–12 m depth, 

MOCNESS, 1 May 2003; UW 145071, 1 (8.9 mm), 

60°32.1′ N, 147°48.3′ W, 21–41 m depth, MOCNESS, 

1 May 2003; UW 145072, 4 (8.2–9.7 mm), 60°32.1′ N, 

147°48.2′ W, 20–41 m depth, MOCNESS, 2 May 2003; 

UW 145076, 3 (9.0–10.1 mm), 60°32.1′ N, 147°48.2′ W, 

0–14 m depth, MOCNESS, 2 May 2003; UW 145073, 

1 (10.2 mm), 60°10.8′ N, 147°38.5′ W, 0–11 m depth, 

MOCNESS, 3 May 2003; UW 145077, 1 (12.0 mm), 

60°32.4′ N, 147°48.0′ W, 20–40 m depth, MOCNESS, 

3 May 2003; UW 145074, 1 (10.3 mm), 60°32.3′ N, 

147°48.3′ W, 0–14 m depth, MOCNESS, 4 May 2003; 

UW 145075, 1 (10.2 mm), 60°32.3′ N, 147°48.3′ W, 

21–41 m depth, MOCNESS, 4 May 2003; UW 145078, 

1 (9.4 mm), 60°10.8′ N, 147°38.5′ W, 11–21 m depth, 

MOCNESS, 4 May 2003; UW 145079, 1 (9.0 mm), 

60°10.8′ N, 147°38.5′ W, 0–11 m depth, MOCNESS, 4 

May 2003; UW 145090, 2 (12.9–13.3 mm), 55°24.9′ N, 

159°50.7′ W, 0–101 m depth, bongo net, 27 May 2003. 

Location, depth, gear, and date of collection unknown 

for the following specimens: UW 145882, 1 (35.6 mm); 

UW 145883, 1 (39.1 mm); UW 145884, 1 (40.4 mm).

Lumpenus fabricii. 22 specimens (12.1–57.0 mm) examined. Chukchi Sea: UW 145069, 2 (55.0–57.0 mm), 

69°58.4′ N, 165°53.3′ W, 0 m depth, Cantrawl, 6 September 2007. Beaufort Sea: UW 145881, 1 (22.5 mm), 

geographic coordinates and depth unknown, bongo net, 

10 August 1977; UW 145067, 1 (19.5 mm), 69°57.7′ N, 

133°8.4′ W, 0–9 m depth, bongo net, 25 June 2003; UW 

145068, 1 (35.0 mm), 69°45.0′ N, 133°21.7′ W, 0–12 m 

depth, bongo net, 25 June 2003. Bering Sea: UW 94975, 

1 (46.0 mm), 57°56.0′ N, 163°30.0′ W, 0–90 m depth, 

Methot net, 6 September 1994; UW 94974, 1 (12.5 mm), 

55°47.3′ N, 167°17.1′ W, 0–125 m depth, bongo net, 8 

May 1995; UW 145063, 2 (20.5–30.0 mm), 67°25.0′ N, 

173°37.5′ W, 0–22 m depth, bongo net, 16 August 2004; 

UW 145064, 1 (37.0 mm), 71°23.6′ N, 174°54.6′ W, 

0–48 m depth, bongo net, 21 August 2004; UW 145065, 

1 (22.5 mm), 71°54.5′ N, 175°29.3′ W, 0–44 m depth, 

bongo net, 21 August 2004; UW 145066, 1 (41.0 mm), 

62° N, 171°0.2′ W, 0–40 m depth, bongo net, 20 September 2004. Gulf of Alaska: UW 94971, 2 (34.0–39.0 mm), 

57°16.0′ N, 152°55.0′ W, 0–90 m depth, Tucker trawl, 

20 June 1978; UW 94976, 1 (34.0 mm), 57°16.0′ N, 

152°55.0′ W, depth unknown, bongo net, 20 June 1978; 

UW 94972, 1 (45.0 mm), 57°16.0′ N, 152°55.0′ W, 

depth unknown, bongo net, 26 July 1978; UW 94973, 1 

(12.1 mm), 57°19.0′ N, 153°2.0′ W, depth, gear, and date 

unknown; UW 123507, 1 (length unknown), 57°37.6′ N, 

152°25.3′ W, 0–50 m depth, bongo net, 18 May 1981; 

UW 123508, 1 (length unknown), 57°40.9′ N, 152°21.7′

W, 0–42 m depth, bongo net, 18 May 1981; UW 123509, 

1 (length unknown), 57°38.7′ N, 152°23.6′ W, 0–79 m 

depth, bongo net, 27 May 1981; UW 94970, 1 (13.3 

mm), 56°24.0′ N, 158°5.0′ W, 0–99 m depth, bongo net, 

25 May 1994. Location, depth, and gear unknown for 

the following specimen: UW 145896, 1 (41.0 mm), 1978. 

Lumpenus maculatus. 80 specimens (8.1–72.1 mm) 

examined. Bering Sea: UW 95047, 3 (27.5–33.0 mm), 

60°6.0′ N, 174°39.6′ W, 0 m depth, neuston net, 3 August 

1994; UW 95049, 1 (29.0 mm), 62°22.3′ N, 179°36.0′

W, 0–50 m depth, Methot net, 19 August 1994; UW 

95032, 1 (10.4 mm), 55°58.0′ N, 163°58.5′ W, 0–77 m 

depth, bongo net, 24 April 1995; UW 95036, 1 (11.0 

mm), 56°14.7′ N, 166°44.4′ W, 0–107 m depth, bongo 

net, 27 April 1995; UW 94977, 1 (28.0 mm), 56°29.5′

N, 166°59.4′ W, 0–98 m depth, Methot net, 21 July 

1995; UW 94978, 1 (36.7 mm), 57°30.2′ N, 167°58.8′

W, 0–66 m depth, Methot net, 22 July 1995; UW 95044, 

1 (61.0 mm), 57°2.6′ N, 170°37.7′ W, 0–73 m depth, 

Methot net, 21 September 1995; UW 94979, 1 (46.0 

mm), 55°30.7′ N, 166°0.8′ W, 0–100 m depth, Methot 

net, 22 July 1996; UW 14506, 1 (42.0 mm), 56°29.6′ N, 

168°57.6′ W, 0–90 m depth, Methot net, 22 July 1997; 

UW 94980, 3 (40.0–45.5 mm), 56°0.4′ N, 165°59.3′ W, 

0–100 m depth, Methot net, 22 July 1997; UW 95045, 

1 (57.5 mm), 56°27.4′ N, 169°28.4′ W, 0–92 m depth, 

Methot net, 11 September 1997; UW 14507, 1 (44.0 

mm), 56°30.6′ N, 168°60.0′ W, 0–116 m depth, Methot 

net, 23 July 2001; UW 145102, 1 (52.8 mm), 56°29.9′

N, 168°58.7′ W, 0–95 m depth, Methot net, 8 August 

2002. Gulf of Alaska: UW 14509, 1 (72.1 mm), 59°54.0′

N, 144°44.4′ W, 0–182 m depth, bongo net, 8 November 

1978; UW 64838, 2 (8.1–8.2 mm), 56°33.8′ N, 153°6.0′

W, 0–90 m depth, bongo net, 18 April 1981; UW 57475, 

1 (10.5 mm), 58°20.8′ N, 149°39.0′ W, 0–131 m depth, 

bongo net, 14 April 1984; UW 63531, 1 (17.0 mm), 

57°32.8′ N, 154°44.5′ W, 0–185 m depth, bongo net, 

23 May 1985; UW 61502, 1 (13.5 mm), 57°19.6′ N, 

155°5.9′ W, 0–223 m depth, bongo net, 28 April 1988; 

UW 145103, 1 (22.4 mm), 56°40.1′ N, 155°12.5′ W, 

0–62 m depth, Tucker trawl, 22 May 1988; UW 145105, 

1 (31.1 mm), 55°58.8′ N, 157°2.1′ W, 0–76 m depth, 

Tucker trawl, 6 June 1988; UW 145100, 1 (19.4 mm), 

56°9.0′ N, 157° W, 0–90 m depth, Tucker trawl, 4 June 

1989; UW 62423, 1 (15.5 mm), 57°17.9′ N, 155°27.8′

W, 0–100 m depth, bongo net, 9 May 1990; UW 145095, 

1 (11.8 mm), 57°18.0′ N, 155°42.9′ W, 0–258 m depth, 
92 Professional Paper NMFS 15 

bongo net, 27 April 1991; UW 94984, 5 (36.0–42.0 

mm), 56°12.3′ N, 156°40.7′ W, 0–77 m depth, Methot net, 26 July 1991; UW 94985, 3 (50.0–53.0 mm), 

56°11.6′ N, 157°21.7′ W, 0–129 m depth, Methot net, 

27 July 1991; UW 94986, 3 (38.9–45.0 mm), 54°38.6′

N, 160°6.6′ W, 0–73 m depth, Methot net, 29 July 1991; 

UW 94987, 3 (47.0–56.5 mm), 54°47.3′ N, 162°33.8′

W, 0–37 m depth, Methot net, 30 July 1991; UW 94981, 

1 (66.2 mm), 58°6.7′ N, 154°10.5′ W, 0–148 m depth, 

Tucker trawl, 30 March 1994; UW 94989, 6 (9.5–12.0 

mm), 57°42.7′ N, 155°13.4′ W, 0–271 m depth, bongo 

net, 4 May 1994; UW 94990, 1 (10.9 mm), 56°25.5′ N, 

155°37.1′ W, 0–69 m depth, bongo net, 6 May 1994; 

UW 94995, 3 (12.1–13.6 mm), 57°34.6′ N, 154°34.0′

W, 0–93 m depth, bongo net, 8 May 1994; UW 95002, 

1 (16.0 mm), 57°6.5′ N, 155°59.5′ W, 0–30 m depth, 

Tucker trawl, 9 May 1994; UW 95010, 1 (16.2 mm), 

57°8.0′ N, 155°56.1′ W, 29–64 m depth, Tucker trawl, 

10 May 1994; UW 95012, 2 (13.1–14.2 mm), 57°3.7′

N, 155°53.1′ W, 56–101 m depth, Tucker trawl, 11 May 

1994; UW 95013, 1 (12.0 mm), 57°3.7′ N, 155°53.1′ W, 

0–56 m depth, Tucker trawl, 11 May 1994; UW 95015, 

1 (15.0 mm), 56°58.6′ N, 155°52.7′ W, 0–52 m depth, 

Tucker trawl, 11 May 1994; UW 95016, 1 (14.0 mm), 

56°58.6′ N, 155°52.7′ W, 52–108 m depth, Tucker trawl, 

11 May 1994; UW 95018, 2 (13.0–14.1 mm), 56°55.6′

N, 155°53.4′ W, 0–294 m depth, bongo net, 11 May 

1994; UW 95020, 1 (17.1 mm), 57°36.3′ N, 155°0.4′ W, 

0–227 m depth, bongo net, 24 May 1994; UW 95022, 

2 (18.8 mm), 57°1.8′ N, 156°21.1′ W, 0–89 m depth, 

bongo net, 25 May 1994; UW 95024, 2 (20.0–21.5 

mm), 55°19.8′ N, 159°15.6′ W, 0–101 m depth, bongo 

net, 26 May 1994; UW 95025, 1 (24.0 mm), 55°22.6′

N, 156°56.4′ W, 0–86 m depth, bongo net, 27 May 

1994; UW 95027, 1 (24.9 mm), 56°17.1′ N, 156°46.8′

W, 0–75 m depth, bongo net, 29 May 1994; UW 95028, 

1 (23.9 mm), 56°31.4′ N, 156°47.1′ W, 0–100 m depth, 

bongo net, 30 May 1994; UW 95029, 2 (18.2–23.0 mm), 

56°52.0′ N, 156°44.7′ W, 0–99 m depth, bongo net, 

30 May 1994; UW 145099, 1 (17.1 mm), 55°31.7′ N, 

160°4.9′ W, 0–100 m depth, bongo net, 27 May 2000; 

UW 145104, 1 (25.2 mm), 57°20.6′ N, 156°2.8′ W, 0–59 

m depth, bongo net, 1 June 2000; UW 95046, 2 (57.0–

62.0 mm), 55°27.8′ N, 158°32.6′ W, 0–134 m depth, 

shrimp trawl, 6 September 2000; UW 145096, 1 (12.0 

mm), 58°50.2′ N, 151°20.8′ W, 0–116 m depth, bongo 

net, 29 April 2001; UW 145098, 1 (16.0 mm), 57°35.3′

N, 155°11.9′ W, 50–60 m depth, MOCNESS, 5 May 

2001; UW 145101, 1 (20.8 mm), 54°55.4′ N, 161°4.9′

W, 0–88 m depth, bongo net, 26 May 2001; UW 145097, 

1 (13.1 mm), 58°12.1′ N, 153°18.5′ W, 0–101 m depth, 

bongo net, 30 May 2006. 

Lumpenus medius. 78 specimens (9.6–46.5 mm) examined. Arctic Ocean: UW 145110, 1 (40.0 mm), 

73°20.0′ N, 165° W, 0–57 m depth, bongo net, 12 September 2003. Beaufort Sea: UW 145112, 1 (32.0 mm), 

69°46.8′ N, 133°8.1′ W, 0–4 m depth, bongo net, 15 

August 2002; UW 145111, 1 (35.0 mm), 69°57.7′ N, 

133°8.4′ W, 0–9 m depth, bongo net, 16 August 2002. 

Gulf of Alaska: UW 33044, 10 (9.8–16.3 mm), 57°16.0′

N, 152°55.0′ W, 70 m depth, Tucker trawl, 5 April 1978; 

UW 33045, 2 (10.2–13.0 mm), 56°52.0′ N, 153°35.0′

W, 0–38 m depth, bongo net, 16 April 1978; UW 33049, 

12 (31.0–41.5 mm), 57°16.0′ N, 152°55.0′ W, 0–90 m 

depth, Tucker trawl, 20 June 1978; UW 33050, 8 (33.0–

38.0 mm), 57°16.0′ N, 152°55.0′ W, 70 m depth, Tucker 

trawl, 20 June 1978; UW 33051, 1 (36.0 mm), 57°16.0′

N, 152°55.0′ W, 90 m depth, Tucker trawl, 20 June 

1978; UW 33052, 1 (39.0 mm), 57°50.0′ N, 152°19.0′

W, 103 m depth, sled trawl, 29 June 1978; UW 33053, 2 

(41.0–44.5 mm), 57°16.0′ N, 152°55.0′ W, 87 m depth, 

sled trawl, 13 July 1978; UW 33064, 2 (40.0–42.5 mm), 

57°50.0′ N, 152°19.0′ W, 130 m depth, sled trawl, 3 

August 1978; UW 33065, 1 (46.0 mm), 57°1.0′ N, 

153°29.0′ W, 116 m depth, sled trawl, 8 August 1978; 

UW 33074, 2 (11.5–12.3 mm), 57°19.0′ N, 153°2.0′ W, 

0–81 m depth, bongo net, 15 March 1979; UW 33075, 

1 (10.0 mm), 57°16.0′ N, 152°55.0′ W, 0–92 m depth, 

bongo net, 15 March 1979; UW 33076, 1 (14.4 mm), 

57°16.0′ N, 152°37.0′ W, 0–47 m depth, bongo net, 15 

March 1979; UW 33079, 6 (10.4–11.9 mm), 57°18.0′ N, 

153°6.0′ W, 0–27 m depth, bongo net, 15 March 1979; 

UW 33080, 15 (9.6–12.2 mm), 57°20.0′ N, 152°55.0′ W, 

0–33 m depth, bongo net, 15 March 1979; UW 33073, 2 

(10.6–11.0 mm), 57°4.0′ N, 153°36.0′ W, 0–81 m depth, 

bongo net, 16 March 1979; UW 145885, 1 (31.0 mm), 

54°17.2′ N, 164°16.1′ W, 0–65 m depth, bongo net, 25 

May 2001. Location, depth, gear, and date of collection unknown for the following specimens: UW 33056, 

1 (11.4 mm); UW 33057, 1 (45.0 mm); UW 33058, 2 

(43.5–45.5 mm); UW 33059, 2 (39.5–43.0 mm); UW 

33063, 1 (46.5 mm); UW 33069, 1 (11.4 mm).

Lumpenus sagitta. 92 specimens (5.8–53.0 mm) examined. Bering Sea: UW 142296, 1 (19.5 mm), 56°5.1′ N, 

160°57.7′ W, 0–15 m depth, bongo net, 20 May 2003. 

Gulf of Alaska: UW 141556, 1 (13.8 mm), 56°32.0′ N, 

155°44.0′ W, 0–152 m depth, bongo net, 29 April 1972; 

UW 141552, 1 (15.3 mm), 57°1.2′ N, 153°27.3′ W, 

0–115 m depth, bongo net, 1 May 1972; UW 33120, 

1 (31.2 mm), 57°16.0′ N, 152°55.0′ W, 0–70 m depth, 

Tucker trawl, 26 May 1978; UW 33125, 1 (51.0 mm), 

57°16.0′ N, 152°55.0′ W, 0–10 m depth, Tucker trawl, 

4 June 1978; UW 33126, 1 (49.5 mm), 57°16.0′ N, 

152°55.0′ W, 0–50 m depth, Tucker trawl, 4 June 1978; 

UW 141253, 1 (15.7 mm), 57°7.3′ N, 155°34.5′ W, depth 

unknown, bongo net, 10 April 1980; UW 60222, 1 (15.4 

mm), 56°22.8′ N, 155°26.1′ W, 0–30 m depth, bongo 

net, 3 April 1981; UW 66714, 1 (30.0 mm), 57°51.7′ N, 

154°40.6′ W, 0–192 m depth, bongo net, 28 April 1981; 

UW 66839, 1 (21.0 mm), 57°49.4′ N, 154°56.0′ W, 0–72 
93

m depth, bongo net, 29 April 1981; UW 70716, 1 (30.0 

mm), 57°52.0′ N, 154°40.8′ W, 0–208 m depth, bongo 

net, 21 May 1981; UW 70755, 1 (30.0 mm), 57°28.5′ N, 

154°45.3′ W, 0–126 m depth, bongo net, 21 May 1981; 

UW 70841, 1 (35.1 mm), 57°29.5′ N, 155°43.0′ W, 0–65 

m depth, bongo net, 21 May 1981; UW 70885, 1 (31.0 

mm), 57°23.9′ N, 154°57.3′ W, 0–208 m depth, bongo 

net, 22 May 1981; UW 37176, 1 (17.3 mm), 56°40.0′

N, 155°27.0′ W, 0–194 m depth, bongo net, 8 April 

1982; UW 95054, 1 (25.5 mm), 55°14.1′ N, 159°2.0′ W, 

0–189 m depth, bongo net, 29 May 1982; UW 37525, 

2 (15.6–16.0 mm), 56°13.4′ N, 156°3.4′ W, 0–219 m 

depth, bongo net, 26 March 1985; UW 60334, 1 (21.5 

mm), 57°0.9′ N, 155°37.0′ W, 0–244 m depth, bongo 

net, 6 May 1985; UW 60353, 1 (22.0 mm), 56°54.4′ N, 

156°14.0′ W, 0–164 m depth, bongo net, 7 May 1985; 

UW 62957, 1 (44.5 mm), 59°7.6′ N, 150°21.0′ W, 0–162 

m depth, bongo net, 19 May 1985; UW 63889, 1 (36.0 

mm), 55°57.0′ N, 155°54.0′ W, 0–64 m depth, bongo 

net, 27 May 1985; UW 50984, 1 (12.6 mm), 57°44.0′

N, 155°5.8′ W, 0–294 m depth, bongo net, 7 April 1986; 

UW 60450, 1 (25.0 mm), 58°0.4′ N, 154°14.1′ W, 0–196 

m depth, bongo net, 6 May 1986; UW 144705, 1 (36.0 

mm), 56°24.0′ N, 153°23.4′ W, 0–60 m depth, Tucker 

trawl, 27 May 1986; UW 95115, 3 (28.0–36.0 mm), 

53°25.8′ N, 166°31.8′ W, 0–58 m depth, Tucker trawl, 

4 June 1986; UW 60932, 1 (15.5 mm), 57°40.0′ N, 

155°18.1′ W, 0–285 m depth, bongo net, 5 April 1987; 

UW 60962, 2 (16.5–18.0 mm), 57°51.3′ N, 154°44.6′

W, 0–262 m depth, bongo net, 5 April 1987; UW 

95069, 1 (16.1 mm), 57°51.4′ N, 154°22.2′ W, 0–205 

m depth, bongo net, 5 April 1987; UW 61006, 1 (15.9 

mm), 57°33.0′ N, 154°52.7′ W, 0–222 m depth, bongo 

net, 6 April 1987; UW 51920, 1 (15.8 mm), 58°14.2′ N, 

154°2.0′ W, 0–256 m depth, bongo net, 5 April 1988; 

UW 51747, 1 (28.0 mm), 57°43.4′ N, 154°56.4′ W, 

0–262 m depth, 7 April 1988; UW 51661, 1 (19.0 mm), 

57°55.9′ N, 154°29.5′ W, 0–285 m depth, bongo net, 

11 April 1988; UW 61461, 1 (23.5 mm), 57°34.2′ N, 

155°17.7′ W, 0–282 m depth, bongo net, 29 April 1988; 

UW 61539, 1 (9.2 mm), 57°23.6′ N, 155°38.8′ W, 0–238 

m depth, bongo net, 29 April 1988; UW 61551, 1 (27.5 

mm), 57°27.0′ N, 155°44.6′ W, 0–147 m depth, bongo 

net, 29 April 1988; UW 142201, 1 (32.5 mm), 56°38.7′

N, 156°31.8′ W, 0–145 m depth, Tucker trawl, 21 May 

1988; UW 95095, 1 (41.0 mm), 55°52.1′ N, 156°47.3′ W, 

0–102 m depth, Tucker trawl, 6 June 1988; UW 67470, 

3 (26.0–30.0 mm), 56°43.4′ N, 155°26.2′ W, 0–93 m 

depth, Tucker trawl, 24 May 1989; UW 52644, 1 (17.0 

mm), 57°47.5′ N, 155°6.4′ W, 0–258 m depth, bongo 

net, 10 April 1990; UW 71418, 1 (40.0 mm), 58°0.1′ N, 

152°0.3′ W, 0–100 m depth, bongo net, 28 May 1990; 

UW 141255, 1 (15.5 mm), 57°56.0′ N, 154°46.6′ W, 

0–208 m depth, bongo net, 17 April 1991; UW 95076, 1 

(17.5 mm), 57°46.8′ N, 155°7.7′ W, 0–259 m depth, bongo net, 26 April 1991; UW 95086, 1 (14.8 mm), 57°38.5′

N, 155°4.3′ W, 0–246 m depth, bongo net, 2 May 1991; 

UW 95087, 1 (19.5 mm), 57°15.8′ N, 156°3.3′ W, 0–105 

m depth, bongo net, 2 May 1991; UW 95097, 1 (11.2 

mm), 56°16.4′ N, 158°4.9′ W, 0–64 m depth, bongo net, 

22 May 1991; UW 141557, 4 (12.9–17.3 mm), 57°46.7′

N, 155°0.2′ W, 0–309 m depth, bongo net (net 2), 7 April 

1992; UW 143892, 1 (15.0 mm), 57°46.7′ N, 155°0.2′

W, 0–309 m depth, bongo net (net 1), 7 April 1992; UW 

143372, 1 (12.2 mm), 56°54.3′ N, 156°16.0′ W, 0–100 

m depth, bongo net, 9 May 1992; UW 95089, 1 (13.2 

mm), 56°45.4′ N, 156°31.0′ W, 0–117 m depth, bongo 

net, 13 May 1992; UW 144493, 1 (5.8 mm), 56°56.1′ N, 

155°10.9′ W, 0–101 m depth, bongo net, 10 May 1993; 

UW 95099, 1 (25.5 mm), 57°54.9′ N, 154°47.7′ W, 

0–101 m depth, bongo net, 11 May 1993; UW 95103, 1 

(27.0 mm), 56°18.8′ N, 155°22.4′ W, 0–41 m depth, bongo net, 6 May 1994; UW 95101, 1 (24.0 mm), 57°8.7′

N, 156°0.6′ W, 0–57 m depth, Tucker trawl, 10 May 

1994; UW 95102, 1 (29.0 mm), 57°3.7′ N, 155°53.1′

W, 0–101 m depth, Tucker trawl, 11 May 1994; UW 

95110, 1 (29.0 mm), 57°16.7′ N, 155°55.2′ W, 0–101 

m depth, bongo net, 28 May 1995; UW 95107, 1 (24.0 

mm), 57°36.3′ N, 155°0.3′ W, 0–201 m depth, bongo 

net, 4 May 1996; UW 94983, 1 (44.0 mm), 54°10.2′ N, 

164°10.9′ W, 0–58 m depth, Methot net, 19 June 1998; 

UW 94982, 5 (45.5–53.0 mm), 55°3.0′ N, 161°4.0′ W, 

0–101 m depth, Methot net, 20 June 1998; UW 95113, 

1 (48.0 mm), 55°31.8′ N, 159°31.8′ W, 0–59 m depth, 

Methot net, 21 June 1998; UW 141719, 1 (19.0 mm), 

54°39.8′ N, 162°47.6′ W, 0–68 m depth, bongo net, 

26 May 2000; UW 141566, 1 (25.0 mm), 57°27.1′ N, 

155°45.6′ W, 0–101 m depth, bongo net, 1 May 2001; 

UW 14454, 1 (29.5 mm), 56°31.3′ N, 156°16.4′ W, 

0–199 m depth, bongo net, 4 May 2001; UW 141549, 1 

(22.0 mm), 58°8.3′ N, 151°48.9′ W, 0–58 m depth, bongo net, 4 May 2003; UW 144287, 1 (46.0 mm), 56°47.0′

N, 155°0.7′ W, 0–63 m depth, bongo net, 30 May 2004; 

UW 141554, 1 (33.0 mm), 57°27.0′ N, 155°45.8′ W, 

0–101 m depth, bongo net, 29 May 2005. Puget Sound, 

Washington: UW 95119, 1 (37.0 mm), 47°20.5′ N, 

122°19.4′ W, 0 m depth, dip net, 6 September 1988; UW 

95111, 2 (34.0–42.0 mm), 48°19.8′ N, 123°8.9′ W, 0 m 

depth, dip net, 6 June 1989; UW 95112, 8 (11.7–14.1 

mm), 48°19.4′ N, 123°0.3′ W, 0 m depth, dip net, 7 April 

1992; UW 142295, 1 (34.3 mm), 47°20.5′ N, 122°19.4′

W, 0 m depth, dip net, 14 May 1995. Location and depth 

unknown for the following specimen: UW 141561, 1 

(42.5 mm), bongo net, May 1979. Location, depth, and 

date of collection unknown for the following specimen: 

UW 95118, 1 (36.0 mm), bongo net. 

Poroclinus rothrocki. 70 specimens (5.5–48.0 mm) examined. Bering Sea: UW 66398, 1 (12.6 mm), 56°3.0′ N, 

166°35.5′ W, 25–26 m depth, Tucker trawl, 3 June 1979; 

UW 65629, 1 (15.5 mm), 55°57.3′ N, 166°44.6′ W, 

0–123 m depth, bongo net, 19 June 1979; UW 129895, 
94 Professional Paper NMFS 15 

2 (26.0–28.0 mm), 56°27.4′ N, 169°28.4′ W, 0–96 m 

depth, Methot net, 12 June 1997. Gulf of Alaska: UW 

65106, 1 (13.0 mm), 55°5.4′ N, 157°47.0′ W, 0–62 m 

depth, bongo net, 24 April 1981; UW 68662, 2 (10.6–

12.1 mm), 56°53.2′ N, 153°23.0′ W, 0–127 m depth, 

bongo net, 21 May 1981; UW 68682, 1 (12.6 mm), 

56°33.8′ N, 153°5.0′ W, 0–68 m depth, bongo net, 21 

May 1981; UW 68770, 2 (11.2–11.4 mm), 56°38.0′ N, 

155°25.0′ W, 0–96 m depth, bongo net, 23 May 1981; 

UW 68790, 7 (11.4–13.1 mm), 56°3.0′ N, 154°56.0′ W, 

0–93 m depth, bongo net, 23 May 1981; UW 71106, 

4 (11.0–12.3 mm), 56°40.4′ N, 155°22.0′ W, 0–122 m 

depth, bongo net, 23 May 1981; UW 36019, 1 (14.3 

mm), 56°44.9′ N, 155°32.9′ W, 0–203 m depth, bongo 

net, 22 May 1983; UW 36380, 1 (16.0 mm), 55°45.3′ N, 

157°21.5′ W, 0–78 m depth, bongo net, 25 May 1983; 

UW 36440, 3 (15.0–15.7 mm), 55°30.9′ N, 157°47.2′

W, 0–60 m depth, bongo net, 26 May 1983; UW 36484, 

2 (14.7–15.6 mm), 55°15.3′ N, 158°41.4′ W, 0–196 m 

depth, bongo net, 27 May 1983; UW 60324, 1 (13.3 

mm), 56°48.2′ N, 155°8.0′ W, 0–83 m depth, bongo 

net, 6 May 1985; UW 60378, 1 (13.1 mm), 55°50.9′ N, 

156°49.0′ W, 0–90 m depth, bongo net, 8 May 1985; 

UW 62968, 1 (13.8 mm), 58°52.2′ N, 149°55.0′ W, 

0–208 m depth, bongo net, 19 May 1985; UW 63121, 1 

(15.3 mm), 59° N, 151°18.5′ W, 0–104 m depth, bongo 

net, 20 May 1985; UW 63205, 1 (18.3 mm), 58°55.7′ N, 

152°51.5′ W, 0–168 m depth, 21 May 1985; UW 63751, 

2 (14.8–15.8 mm), 56°25.8′ N, 153°29.0′ W, 0–62 m 

depth, 25 May 1985; UW 64286, 1 (13.9 mm), 55°16.8′

N, 158°40.8′ W, 0–190 m depth, bongo net, 1 June 

1985; UW 145119, 1 (11.0 mm), 55°56.2′ N, 156°23.8′

W, 45–60 m depth, MOCNESS, 15 May 1986; UW 

60840, 1 (10.4 mm), 56°22.4′ N, 156°45.2′ W, 0–114 

m depth, bongo net, 16 May 1986; UW 67222, 1 (14.1 

mm), 56°41.1′ N, 156°44.7′ W, 0–139 m depth, bongo 

net, 22 May 1987; UW 67299, 1 (11.8 mm), 56°32.7′ N, 

156°32.8′ W, 0–191 m depth, bongo net, 23 May 1987; 

UW 145118, 1 (9.6 mm), 56°33.6′ N, 156°32.6′ W, 30–

45 m depth, MOCNESS, 24 May 1987; UW 145121, 

1 (11.5 mm), 56°33.9′ N, 156°32.7′ W, 1–15 m depth, 

MOCNESS, 25 May 1987; UW 145117, 1 (10.6 mm), 

56°35.1′ N, 156°31.9′ W, 30–45 m depth, MOCNESS, 

25 May 1987; UW 145138, 1 (10.0 mm), 56°16.7′ N, 

157°10.0′ W, 0–104 m depth, Tucker trawl, 23 May 

1988; UW 145127, 1 (10.1 mm), 56°38.9′ N, 156°31.5′

W, 0–101 m depth, Tucker trawl, 2 June 1988; UW 

145128, 1 (18.0 mm), 56°21.4′ N, 157°3.8′ W, 23–46 

m depth, Tucker trawl, 7 June 1988; UW 72252, 1 (22.0 

mm), 57°1.0′ N, 156°18.9′ W, 0–95 m depth, bongo 

net, 3 June 1990; UW 129519, 1 (5.5 mm), 56°37.0′ N, 

156°20.0′ W, 0–101 m depth, bongo net, 13 May 1993; 

UW 129629, 2 (12.0 mm), 55°29.0′ N, 158°8.2′ W, 0–101 

m depth, bongo net, 23 May 1995; UW 129645, 1 (14.3 

mm), 58°40.5′ N, 152°45.5′ W, 0–100 m depth, bongo 

net, 1 June 1996; UW 129681, 1 (22.5 mm), 55°31.8′ N, 

159°31.8′ W, 0–60 m depth, bongo net, 21 June 1998; 

UW 129893, 1 (20.0 mm), 55°31.8′ N, 159°31.8′ W, 

0–59 m depth, Methot net, 21 June 1998; UW 145133, 

1 (24.0 mm), 59°15.3′ N, 149°34.4′ W, 0–113 m depth, 

bongo net, 31 May 2002; UW 145132, 1 (16.1 mm), 

58°18.4′ N, 148°58.5′ W, 40–60 m depth, MOCNESS, 

5 June 2002; UW 145134, 2 (17.9–18.8 mm), 59°35.2′

N, 147°0.4′ W, 0–200 m depth, bongo net, 12 May 

2003; UW 113858, 1 (48.0 mm), 58° N, 152° W, 0–156 

m depth, gear unknown, 25 July 2003; UW 145135, 1 

(12.5 mm), 57°40.7′ N, 155°11.0′ W, 0–281 m depth, 

31 May 2004; UW 145891, 1 (26.5 mm), 56°33.0′ N, 

157°44.4′ W, 39–74 m depth, Tucker trawl, 14 September 2009. British Columbia, Canada: UW 42915, 1 (38.0 

mm), 48°52.5′ N, 126°53.0′ W, depth and gear unknown, 

8 September 1964. Washington coast: UW 145125, 1 

(17.8 mm), 47°6.3′ N, 124°12.5′ W, 0–55 m depth, 1-m 

net, 6 May 1971; UW 145126, 1 (18.4 mm), 47°6.3′ N, 

124°12.5′ W, 0–37 m depth, 1-m net, 6 May 1971; UW 

58120, 1 (10.7 mm), 47°20.0′ N, 124°25.0′ W, 0–140 

m depth, bongo net, 24 April 1980. Oregon coast: UW 

145122, 3 (12.8–16.5 mm), 46° N, 124°8.0′ W, 0–91 m 

depth, 1-m net, 4 May 1971; UW 58255, 1 (17.3 mm), 

46° N, 124°25.0′ W, 0–131 m depth, bongo net, 26 April 

1980; UW 54125, 1 (16.5 mm), 43°40.0′ N, 124°21.5′

W, 0–109 m depth, bongo net, 22 May 1981. Location 

and depth unknown for the following specimen: UW 

145114, 1 (16.2 mm), 1-m net, 3–6 May 1971.

Opisthocentrus ocellatus. 7 specimens (13.6–15.8 

mm) examined (depths and gear unknown). Sea of Japan: HUMZ-L 02671, 2 (14.4–15.2 mm), 41°56.0′ N, 

140°57.0′ E, 20 April 1983; HUMZ-L 03502, 5 (13.6–

15.8 mm), 41°56.0′ N, 140°57.0′ E, 12 April 1985.

Plectobranchus evides. 17 specimens (7.0–49.0 mm) 

examined. Oregon coast: UW 145887, 1 (37.0 mm), 

44°39.0′ N, 124°39.9′ W, 130 m depth, midwater 

trawl, 28 June 1970; OS 13628, 1 (7.0 mm), 45°23.7′

N, 124°10.2′ W, 0–138 m depth, 0.7-m bongo net, 28 

March 1972; OS 13626, 1 (11.1 mm), 44°10.4′ N, 

124°13.8′ W, 0–75 m depth, 0.7-m bongo net, 29 March 

1972; OS 13627, 1 (7.8 mm), 44°2.4′ N, 124°4.8′ W, 

0–80 m depth, 0.7-m bongo net, 29 March 1972; UW 

145886 1 (20.0 mm), 44°17.5′ N, 124°20.6′ W, 0–86 

m depth, 0.7-m bongo net, 20 April 1973; UW 145889, 

1 (16.9 mm original length, 15.1 mm after preservative changed to ethanol), 44°6.0′ N, 124°22.1′ W, 75 m 

depth, 0.7-m bongo net, 20 April 1973; UW 145888, 1 

(9.2 mm), 45°4.0′ N, 124°15.0′ W, 105 m depth, 0.7-m 

bongo net, 24 April 1973; UW 145113, 1 (34.0 mm), 

44° N, 124°36.2′ W, 0–124 m depth, bongo net, 22 

May 1981; UW 145115, 2 (23.0–28.0 mm), 43°57.6′ N, 

124°16.3′ W, 0–75 m depth, bongo net, 14 May 1982; 

UW 55422, 1 (9.5 mm), 43°40.0′ N, 124°17.1′ W, 0–82 

m depth, bongo net, 24 March 1984; UW 145116, 5 
95

(33.5–49.0 mm), 44°23.6′ N, 124°22.8′ W, 0–266 m 

depth, mid water trawl, 20 July 2007. Location, depth, 

gear, and date of collection unknown for the following 

specimen: UW 145890, 1 (39.5 mm).

Alectrias spp. 13 specimens (12.8–16.5 mm) examined 

(depths and gear unknown). Sea of Japan: HUMZ-L 

02756, 2 (12.8–16.5 mm), 41°56.0′ N, 140°57.0′ E, 26 

May 1983; HUMZ-L 02920, 1 (14.4 mm), 41°56.0′ N, 

140°57.0′ E, 15 June 1985; HUMZ-L 03915, 10 (14.0–

16.2 mm), 42°18.0′ N, 140°58.0′ E, 15 June 1984.

Alectrias alectrolophus. 6 specimens (10.7–16.6 

mm) examined (depths and gear unknown). Sea of Japan: HUMZ-L 02576, 3 (10.7–16.6 mm), 41°56.0′ N, 

140°57.0′ E, 26 May 1983; HUMZ-L 02920, 2 (15.0–

16.4 mm), 41°56.0′ N, 140°57.0′ E, 15 June 1985; 

HUMZ-L 03439, 1 (14.1 mm), 41°56.0′ N, 140°57.0′ E, 

12 June 1985.

Anoplarchus insignis. 178 specimens (5.2–36.5 mm) 

examined. Bering Sea: UW 141553, 1 (6.5 mm), 54°0.6′

N, 166°34.3′ W, 0–110 m depth, bongo net, 13 May 

2002; UW 141737, 1 (6.6 mm), 54°14.6′ N, 165°49.6′

W, 0–79 m depth, bongo net (net 1), 18 May 2003; UW 

141551, 4 (6.8–9.5 mm), 54°14.6′ N, 165°49.6′ W, 0–79 

m depth, bongo net (net 2), 18 May 2003; UW 141735, 

1 (6.3 mm), 54°7.3′ N, 166°12.1′ W, 0–80 m depth, 

bongo net (net 1), 18 May 2003; UW 141861, 2 (7.2 

mm), 54°7.3′ N, 166°12.1′ W, 0–80 m depth, bongo net 

(net 2), 18 May 2003; UW 143915, 2 (11.6–15.2 mm), 

54°24.8′ N, 165°8.9′ W, 0–146 m depth, bongo net, 18 

May 2003; UW 140950, 4 (6.7–13.6 mm), 54°29.4′ N, 

165°4.1′ W, 0–53 m depth, bongo net (net 1), 19 May 

2003; UW 141567, 4 (6.4–8.9 mm), 54°29.4′ N, 165°4.1′

W, 0–53 m depth, bongo net (net 2), 19 May 2003; 

UW 141550, 3 (6.4–7.9 mm), 54°42.7′ N, 165°16.2′

W, 0–132 m depth, bongo net (net 1), 19 May 2003; 

UW 144184, 2 (6.3–7.9 mm), 54°42.7′ N, 165°16.2′

W, 0–132 m depth, bongo net (net 2), 19 May 2003; 

UW 141565, 5 (7.6–12.4 mm), 54°27.8′ N, 165°0.3′ W, 

0–70 m depth, bongo net (net 1), 19 May 2003; UW 

141560, 7 (6.6–12.2 mm), 54°27.8′ N, 165°0.3′ W, 

0–70 m depth, bongo net (net 2), 19 May 2003; UW 

144452, 1 (10.0 mm), 54°56.6′ N, 164°30.6′ W, 0–164 

m depth, bongo net, 19 May 2003; UW 145894, 1 (7.3 

mm), 55°9.2′ N, 165°41.7′ W, 0–114 m depth, bongo 

net, 23 May 2003. Gulf of Alaska: UW 141558, 1 (17.6 

mm), 57°12.1′ N, 152°44.5′ W, 0 m depth, neuston net, 

24 June 1978; UW 32887, 1 (19.2 mm), 58°11.5′ N, 

152°15.2′ W, 90 m depth, Tucker trawl, 28 June 1978; 

UW 141559, 37 (5.2–8.3 mm), 56°18.9′ N, 155°23.1′

W, 0–37 m depth, 23 May 1988; UW 141090, 1 (12.7 

mm), 56°38.4′ N, 156°44.0′ W, 50–75 m depth, Tucker 

trawl, 31 May 1988; UW 141726, 1 (13.1 mm), 55°49.1′

N, 156°40.4′ W, 0–101 m depth, Tucker trawl, 4 June 

1988; UW 27394, 1 (36.5 mm), 60°12.0′ N, 147°50.0′

W, 0–25 m depth, rock dredge, 1 Aug 1989; UW 40411, 

1 (17.0 mm), 60°21.0′ N, 147°49.0′ W, 0–40 m depth, 

beam trawl, 7 August 1989; UW 143886, 6 (7.0–9.8 

mm), 56°49.1′ N, 154°56.2′ W, 0–50 m depth, bongo 

net, 10 May 1993; UW 143071, 4 (12.0–16.4 mm), 

55°0.8′ N, 155°23.3′ W, 0–58 m depth, bongo net, 29 

May 1998; UW 141564, 12 (6.6–20.6 mm), 55°50.7′ N, 

155°50.7′ W, 0–46 m depth, bongo net, 25 June 1998; 

UW 140942, 5 (13.7–20.1 mm), 56°4.2′ N, 155°24.1′ W, 

0–25 m depth, Tucker trawl, 29 June 1998; UW 141548, 

1 (18.2 mm), 55°56.4′ N, 156°1.7′ W, 0–76 m depth, 

Tucker trawl, 29 June 1998; UW 141555, 8 (7.5–11.7 

mm), 57°24.1′ N, 156°14.1′ W, 0–30 m depth, bongo 

net, 1 June 2000; UW 141562, 1 (13.7 mm), 58°57.2′

N, 150°55.9′ W, 0–138 m depth, bongo net, 29 May 

2002; UW 143341, 1 (10.9 mm), 58°29.3′ N, 151°9.1′

W, 0–87 m depth, bongo net, 3 June 2004. Washington 

coast: UW 141720, 1 (18.5 mm), 47°59.6′ N, 124°50.1′

W, 0–49 m depth, bongo net, 3 May 1982. Puget Sound, 

Washington: UW 145147, 16 (6.3–13.8 mm), 47°33.7′

N, 122°32.1′ W, 0 m depth, dip net, 22 May 1987; UW 

145148, 17 (9.2–15.0 mm), 47°33.7′ N, 122°32.1′ W, 

0 m depth, dip net, 5 May 1988; UW 143894, 1 (12.1 

mm), 47°33.7′ N, 122°32.1′ W, 0 m depth, dip net, 5 

May 1989; UW 145149, 1 (13.9 mm), 47°33.7′ N, 

122°32.1′ W, 0 m depth, dip net, 7 June 1989; UW 

145150, 3 (13.7–15.8 mm), 47°33.7′ N, 122°32.1′ W, 

0 m depth, dip net, 5 April 1990; UW 145151, 1 (13.1 

mm), 47°33.7′ N, 122°32.1′ W, 0 m depth, dip net, 20 

April 1991; UW 145152, 2 (13.1–15.6 mm), 47°33.7′

N, 122°32.1′ W, 0 m depth, dip net, 5 May 1992; UW 

145153, 7 (13.0–14.9 mm), 47°33.7′ N, 122°32.1′ W, 

0 m depth, dip net, 5 May 1995; UW 145154, 1 (12.7 

mm), 47°33.7′ N, 122°32.1′ W, 0 m depth, dip net, 12 

May 1995; UW 145155, 10 (11.7–15.9 mm), 47°33.7′

N, 122°32.1′ W, 0 m depth, dip net, 14 May 1995.

Anoplarchus purpurescens. 204 specimens (4.7–37.0 

mm) examined. Bering Sea: UW 48963, 1 (28.0 mm), 

51°32.5′ N, 178°59.0′ W, depth and gear unknown, 

29 May 1956; UW 22417, 1 (17.1 mm), 51°30.0′ N, 

179°2.4′ W, depth and gear unknown, 19 August 1968; 

UW 144877, 2 (6.5–7.1 mm), 54°15.0′ N, 165°49.5′ W, 

0–74 m depth, bongo net, 13 May 2002. Gulf of Alaska: UW 48936, 1 (37.0 mm), 56°53.1′ N, 153°43.3′ W, 

depth and gear unknown, 11 May 1962; UW 27394, 

1 (36.5 mm), 60°12.0′ N, 147°50.0′ W, 25 m depth, 

rock dredge, 1 August 1989; UW 25833, 1 (32.0 mm), 

59°40.0′ N, 151°41.0′ W, depth and gear unknown, 

3 June 1992; UW 144886, 1 (9.9 mm), 57°11.4′ N, 

154°45.3′ W, 0–68 m depth, bongo net, 31 May 1994; 

UW 144884, 1 (14.3 mm), 56°18.3′ N, 155°24.8′ W, 

0–43 m depth, bongo net, 28 May 1998; UW 144885, 

35 (4.7–10.4 mm), 57°24.1′ N, 156°14.1′ W, 0–30 m 

depth, bongo net, 1 June 2000; UW 144878, 1 (9.6 mm), 
96 Professional Paper NMFS 15 

55°4.2′ N, 159°9.2′ W, 0–65 m depth, bongo net, 27 

May 2003; UW 144879, 2 (9.5–12.6 mm), 57°0.3′ N, 

155°52.2′ W, 0–100 m depth, bongo net, 31 May 2003; 

UW 144880, 2 (7.3–8.9 mm), 57°43.5′ N, 152°54.8′ W, 

0–20 m depth, bongo net, 23 May 2004; UW 144883, 

11 (6.4–10.7 mm), 57°40.5′ N, 152°21.0′ W, 0–100 m 

depth, bongo net, 23 May 2004; UW 144881, 1 (6.6 

mm), 56°20.0′ N, 153°43.4′ W, 0–72 m depth, bongo 

net, 24 May 2004; UW 144882, 1 (6.9 mm), 58°27.5′

N, 153°17.0′ W, 0–100 m depth, bongo net, 1 June 

2004. British Columbia, Canada: UW 145873, 8 (7.3–

9.0 mm), 49°21.6′ N, 123°29.3′ W, reared, 11 March 

1976. Puget Sound, Washington: UW 144895, 2 (6.1–9.0 

mm), 47°34.1′ N, 122°12.3′ W, reared, 1 March 1978; 

UW 144891, 46 (5.2–8.0 mm), 47°20.5′ N, 122°19.4′ W, 

0–10 m depth, 0.5-m net, 11 April 1989; UW 145895, 

1 (13.2 mm), 48°20.0′ N, 123°8.9′ W, 0 m depth, dip 

net, 11 May 1989; UW 144888, 1 (14.5 mm), 48°20.0′

N, 123°8.9′ W, 0 m depth, dip net, 12 May 1989; UW 

144870, 1 (13.1 mm), 47°20.5′ N, 122°19.4′ W, 0 m 

depth, dip net, 7 June 1989; UW 144896, 1 (6.0 mm), 

48°56.0′ N, 122°24.1′ W, depth and gear unknown, 25 

February 1992; UW 144892, 4 (5.8–13.2 mm), 47°20.5′

N, 122°19.4′ W, 0 m depth, dip net, 5 May 1992; UW 

145872, 1 (14.3 mm), 47°20.5′ N, 122°19.4′ W, 0 

m depth, dip net, 12 May 1995; UW 144894, 1 (16.8 

mm), 47°20.5′ N, 122°19.4′ W, 0 m depth, dip net, 14 

May 1995; UW 144890, 5 (10.3–14.6 mm), 47°20.5′ N, 

122°19.4′ W, 0 m depth, dip net, 21 April 1998; UW 

144893, 3 (12.6–14.3 mm), 47°20.5′ N, 122°19.4′ W, 0 

m depth, dip net, 26 April 1998; UW 144889, 68 (6.4–

11.2 mm), 48°19.4′ N, 123°0.3′ W, 0 m depth, dip net, 

5 April 2000. Oregon coast: UW 144875, 1 (20.0 mm), 

45°39.9′ N, 124°1.8′ W, 0–49 m depth, bongo net, 27 

April 1985.

Cebidichthys violaceus. 10 specimens (6.2–14.4 mm) 

examined (depths unknown). California coast: UW 

144868, 1 (13.6 mm), 35°12.0′ N, 120°50.5′ W, gear 

unknown, 9 June 1998; UW 144863, 3 (6.2–6.7 mm), 

35°12.0′ N, 120°50.5′ W, bongo net, 22 May 1999; 

UW 144865, 3 (9.0–10.7 mm), 35°12.0′ N, 120°50.5′

W, bongo net, 22 May 1999; UW 144866, 1 (8.6 mm), 

35°12.0′ N, 120°50.5′ W, bongo net, 22 May 1999; UW 

144867, 1 (10.5 mm), 35°12.0′ N, 120°50.5′ W, bongo 

net, 22 May 1999; UW 144869, 1 (14.4 mm), 35°12.0′

N, 120°50.5′ W, bongo net, 25 May 1999. 

Other material (illustrated specimens only)

Eumesogrammus praecisus (tentative). Beaufort Sea: 

UW 145179, 1 (10.6 mm), 71°10.2′ N, 156°8.0′ W, 45 m 

depth, bongo net, 13 August 2008.

Chirolophis cf. snyderi. 2 specimens (18.5–21.0 mm) 

examined. Gulf of Alaska: UW 144864, 1 (21.0 mm), 

57°12.1′ N, 152°44.5′ W, 0 m depth, neuston net, 24 

June 1978; UW 145178, 1 (18.5 mm), 56°37.1′ N, 

156°10.8′ W, 0–101 m depth, bongo net, 13 May 1993.

Acantholumpenus mackayi. Norton Sound, Alaska: 

UW 145177, 1 (36.5 mm), 64°26.4′ N, 163°39.5′ W, 16 

m depth, bottom trawl, 26 July 2002.

Lumpenopsis hypochroma (tentative). Lambert Channel, British Columbia, Canada: UW 145180, 1 (10.3 

mm), 49°30.6′ N, 124°43.5′ W, reared (Vancouver Public 

Aquarium [VPA]), 22 April 1990.

Alectridium aurantiacum (tentative). Bering Sea: 

UW 145871 (specimen lost), 1 (21.0 mm), 56°42.5′ N, 

152°35.9′ W, 25–64 m depth, Tucker trawl, 10 July 1997.

Xiphister atropurpureus. 2 specimens (6.3–16.0 mm) 

examined. British Columbia, Canada: UW 145176, 1 

(16.0 mm), 49°21.6′ N, 123°29.3′ W, reared (VPA), 16 

May 1973. Washington coast: UW 144848, 1 (6.3 mm), 

48°15.9′ N, 124°15.2′ W, reared (VPA), 15 March 1991.

Phytichthys chirus (tentative). Puget Sound, Washington: UW 145181, 1 (19.0 mm), 47°33.7′ N, 122°32.1′ W, 

0 m depth, dip net, 5 May 1988.