NOAA Technical Memorandum NMFS-AFSC-22

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Alaska Fisheries Science Center

September 1993

by

Mei-Sun Yang

Food Habits of the Commercially

Important Groundfishes in the

Gulf of Alaska in 1990
NOAA Technical Memorandum NMFS

The National Marine Fisheries Service's Alaska Fisheries Science Center uses the

NOAA Technical Memorandum series to issue informal scientific and technical

publications when complete formal review and editorial processing are not

appropriate or feasible. Documents within this series reflect sound professional

work and may be referenced in the formal scientific and technical literature.

The NMFS-AFSC Technical Memorandum series of the Alaska Fisheries Science

Center continues the NMFS-F/NWC series established in 1970 by the Northwest

Fisheries Center. The new NMFS-NWFSC series will be used by the Northwest

Fisheries Science Center.

This document should be cited as follows:

 Yang, M-S. 1993. Food Habits of the Commercially Important 

Groundfishes in the Gulf of Alaska in 1990. U.S. Dep. Commer., 

NOAA Tech. Memo. NMFS-AFSC-22, 150 p.

Reference in this document to trade names does not imply endorsement by the

National Marine Fisheries Service, NOAA.
NOAA Technical Memorandum NMFS-AFSC-22

Food Habits of the Commercially Important

Groundfishes in the Gulf of Alaska in 1990

by

Mei-Sun Yang

Alaska Fisheries Science Center

7600 Sand Point Way N.E., BIN C-15700

Seattle, WA 98115-0070

U.S. DEPARTMENT OF COMMERCE

Ronald H. Brown, Secretary

National Oceanic and Atmospheric Administration

Diana Josephson, Deputy Under Secretary

National Marine Fisheries Service
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iii

ABSTRACT

This report describes the food habits of the main

commercially important groundfish species in the Gulf of Alaska

during the summer of 1990. Commercially important prey,

predator-prey size relationships, and the distribution of the

prey--consumed are discussed.
V

CONTENTS

ABSTRACT ................................................ iii

INTRODUCTION .............................................. 1

METHODS ................................................... 4

RESULTS

Walleye pollock ......................................... 9

Pacific cod ............................................. 2 6

Arrowtooth flounder..................................... 5 1

Pacific halibut ......................................... 6 5

Sablefish ............................................... 8 9

Pacific ocean perch ..................................... 105

Rougheye rockfish ....................................... 113

Northern rockfish ....................................... 120

Dusky rockfish.......................................... 125

Shortspine thornyhead ................................... 131

Shortraker rockfish ..................................... 137

SUMMARY.............. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

Page
INTRODUCTION

There is about 5.2 million metric tons (t) of estimated

biomass of groundfish resources in the Gulf of Alaska (Stark and

Clausen 1993). The predator-prey relationships between these

species form a complex food web within the groundfish community

(Smith et al. 1978). Because of these complex relationships,

there have been many fish food habits studies in the Gulf of

Alaska (Jewett 1978, Smith et al. 1978, Hunter 1979, Frost and

Lowry 1981, Clausen 1983, Albers and Anderson 1985, Best and

St-Pierre 1986, Kendall et al. 1987, Grover and Olla 1990, and

Brodeur et al. 1991). However, in some cases, these studies were

either restricted to a small study area or focused on a single

species. The objective of this study was to describe the food

habits of several commercially important groundfish in the Gulf

of Alaska. Most of the western and central Gulf of Alaska (North

Pacific Fisheries Management Council Regulatory areas) was

sampled for 11 predator species (Pacific cod, Gadus

macroceohalus; walleye pollock, Therasra chalcogramma; Pacific

halibut, Hipposlossus stenolepis; arrowtooth flounder,

Atheresthes stomias; sablefish, Anoplopoma fimbria; Pacific ocean

perch, Sebastes alutus; northern rockfish, Sebastes polyspinis;

dusky rockfish, Sebastes ciliatus; rougheye rockfish, Sebastes

aleutianus; shortraker rockfish, Sebastes borealis; and

shortspine thornyhead, Sebastolobus alascanus).
2

CITATIONS

Albers, W. D., and P. J. Anderson. 1985. Diet of Pacific cod,

Gadus macrocephalus, and predation on the northern pink

shrimp, Pandalus borealis, in Pavlof Bay, Alaska. Fish.

Bull., U.S. 83: 601-610.

Best, E. A., and G. St-Pierre. 1986. Pacific halibut as

predator and prey. Int. Pac. Halibut Comm. Tech. Rep. No.

21.

Brodeur, R. D., K. M. Bailey, and S. Kim. 1991. Cannibalism on

eggs by walleye pollock, Theragra chalcoaramma, in Shelikof

Strait, Gulf of Alaska. Mar. Ecol. Prog. Ser. 71: 207-218.

Clausen, D. M. 1983. Food of walleye pollock, Theragra

chalcogramma, in an embayment of southeastern Alaska. Fish.

Bull., U.S. 81: 637-642.

Frost, K. J., and L. F. Lowry. 1981. Trophic importance of some

marine gadids in northern Alaska and their body-otolith size

relationships. Fish. Bull., U.S. 79: 187-192.

Grover, J. J., and B. L. Olla. 1990. Food habits of larval

sablefish Anoplopoma fimbria from the Bering Sea. Fish.

Bull., U.S. 88: 811-814.

Hunter, M. A. 1979. Food resource partitioning among demersal

fishes in the vicinity of Kodiak Island, Alaska. M.S.

Thesis. Univ. Washington, Seattle, WA, 131 p.
3

Jewett, s. c. 1978. Summer food of the Pacific cod, Gadus

macrocephalus, near Kodiak Island, Alaska. Fish. Bull.,

U.S. 76: 700-706.

Kendall, A. W., Jr., M. E. Clarke, M. M. Yoklavich, and G. W.

Boehlert. 1987. Distribution, feeding, and growth of

larval walleye pollock, Theragra chalcogramma, from Shelikof

Strait, Gulf of Alaska. Fish. Bull., U.S. 85: 499-521.

Smith, R. L., Paulson, A. C., and J. R. Rose. 1978. Food and

feeding relationships in the benthic and demersal fishes of

the Gulf of Alaska and Bering Sea. Final report to NOAA

RU284 OCSEAP. Boulder, CO., p. 33-107., Vol. 1, June 1978.

Stark, J. W. and D. M. Clausen. In prep. 1990 Gulf of Alaska

bottom trawl survey. Alaska Fish. Sci. Cent., 7600 Sand

Point Way NE, BIN C15700, Seattle, WA 98115-0070.
4

METHODS

Study Area

In 1990, the Resource Assessment and Conservation

Engineering (RACE) Division at the Alaska Fisheries Science

Center conducted the third comprehensive triennial survey of

groundfish resources in the western and central Gulf of Alaska

(North Pacific Fishery Management Council Regulatory areas).

This survey covered the area from the Islands of Four Mountains

(170o W long.) to Cape St. Elias (144o 30' W long.). The Food

Habits Program at the Resource Ecology and Fishery Management

(REFM) Division collected fish stomach samples during this

survey. The food habits study area (Fig. 1) was smaller than the

survey area. It covered the area from 161o 35' W long. to 144o

30' W long. and did not include the westernmost portion (from

170° W long. to 161o W long.) of the 1990 Gulf of Alaska survey

area.

Sample Collection

Fish stomach samples were collected by scientists on board

the charter boats Pat San Marie and Green Hope from late June to

early September in 1990. Before excising a stomach, fish were

examined for evidence of regurgitation or net feeding. If a fish

had food in its mouth or around the gills, or if its stomach was

inverted or flaccid, the fish was categorized as having

regurgitated food, and the specimen was discarded. If a predator

had fresh food (usually fish) sticking out of the mouth or the

throat, it was categorized as a net-feeding fish and was also
168 oow 164 0 0 160 0 0 156 0 0 152 0 0 148 0 0 144 0 0

Figure 1. --Sampling locations (+) for the stomachs of groundfish

collected‘ in the Gulf of Alaska in 1990.
6

discarded. When a sampled stomach was retained, it was put in a

cloth stomach bag. A field tag with the species name, fork

length (FL) of the fish, and haul data (vessel, cruise, haul

number, specimen number) was also put in the bag. All of the

samples collected were then preserved in buckets containing a 10%

formalin solution. When the samples arrived at the laboratory,

they were transferred into a 70% ethanol alcohol solution before

the stomach contents were analyzed.

Stomach Contents Analysis

In the laboratory, the stomach contents were first blotted

with a paper towel and the wet weight was then recorded to the

nearest one-tenth of a gram. After obtaining the total weight

for a stomach's contents, the contents were placed on a petri

dish and examined under the microscope. Each prey item was

classified to the lowest practical taxonomic level. The prey

items of all rockfishes, Pacific halibut, and sablefish were

weighed and enumerated. The numbers of non-commercially

important prey were not counted for Pacific cod, walleye pollock,

and arrowtooth flounder; instead the percent volume of these prey

items were visually estimated. Prey weights and numbers of

commercially important crabs and fish were recorded. If pollock

otoliths were found, otolith lengths were measured and the

pollock's standard length (SL) was derived through an otolith

length-fish length regression table. Standard lengths of prey

fish, carapace widths (CW) of Tanner crabs and snow crabs

(Chionoecetes spp.) were also recorded.
7

During this study, discarded fish parts from commercial fish

processing operations were also found quite frequently in the

stomachs of some marine fishes (e.g., sablefish). Fish were

identified as a fishery discard if the parts (usually heads or

tails) had a clean cut.

Data Analysis

The general diet of each species was summarized by showing

the overall percent frequency of occurrence, percent of numbers

(if available), and the percent of the total weight of each prey

item found in the stomach. Change in diet by predator size in

terms of percent by weight of main prey items was shown in each

10 cm FL group. The prey size frequency data of the commercially

important fish and crabs were also summarized by predator size

groups.

The geographic distribution of the commercially important

prey consumed (expressed as percent by weight of the total

stomach contents weight in each haul) by the predators were also

shown. The commercially important prey found in this study

include walleye pollock, Pacific cod, Pacific halibut, yellowfin

sole (Pleuronectes asper), rock sole (Pleuronectes bilineatus),

flathead sole (Hippoglossoides elassodon), rex sole (Errex

zachirus), Dover sole (Microstomus pacificus), arrowtooth

flounder, Greenland turbot (Reinhardtius hippoglossoides),

Pacific herring (Clupea pallasi), coho salmon (Oncorhynchus

kisutch), all rockfish species, Tanner crabs, and pandalid

shrimp. Although capelin (Mallotus villosus), Pacific sand lance

(Ammodytes hexapterus), and eulachon (Thaleichthvs pacificus)
8

were not commercially important fish in the Gulf of Alaska area,

they were food of many commercially important fish and were

economically important in some other areas (e.g., Japan);

therefore, data on these prey fish were also analyzed in this

report.
9

WALLEYE POLLOCK

Walleye pollock (Theraura chalcouramma) ranked second (next

to arrowtooth flounder) in biomass in the groundfish complex in

the Gulf of Alaska in 1990. The exploitable biomass in 1990

estimated from the stock synthesis (SS) model was 1,050,000

metric tons (t) (North Pacific Fishery Management Council 1991).

Walleye pollock feed mainly on euphausiids, majids (in the

megalops stage), and shrimp. With increasing size, pollock also

eat juvenile pollock and other teleosts (Smith et al. 1978,

Clausen 1983). Therefore, it is important to understand the food

habits of the pollock and their possible impact on other

commercially important fishes.

RESULTS

General Diet

A total of 1,059 walleye pollock stomachs were analyzed, of

which 985 (93%) contained food. Pollock size ranged from 8 to

70 cm FL with a mean and standard deviation (hereafter, the

standard deviation will be denoted as SD) of 45.4 + 13.0 cm. The

average depth of the 76 haul locations where pollock stomachs

were collected was 173 + 61 m with a range from 46 to 337 m.

Table l-l lists all of the food items found in the pollock

stomachs, the percentage of the frequency of occurrence of the

prey, and the percentage of the prey weight to the total food

weight. The data presented in Table l-1 suggests that walleye

pollock feed mainly on euphausiids and shrimp (39% and 33% by

weight, respectively). Though copepods, mysids, and amphipods
10

Table l-l.--Prey items (expressed in percent frequency of

occurrence and percent total weight) of Theragra

chalcogramma (walleye pollock) collected in the Gulf of

Alaska in 1990.

Prey Name

% Freq. % Total

occur. weight

Polychaeta (unidentified) 3.40 0.57

Pteropoda (snail) 1.20 0.01

Bivalvia (clam) 0.30 0.02

Cephalopoda (unidentified) 1.70 0.08

Teuthoidea (squid unidentified) 2.30 0.36

Gonatidae (squid) 0.10 1.87

Berrvteuthis masister (squid) 0.10 0.33

Octopoda (octopus unidentified) 0.30 0.19

Calanoida (copepod unidentified) 33.30 0.80

Neocalanus cristatus (copepod) 0.80 <0.01 

Eucalanus bunaii (copepod) 0.20 <0.01

Pseudocalanus sp. (copepod) 0.10 <0.01

Mysidacea (mysid unidentified) 15.50 0.81

Acanthomvsis macronsis (mysid) 0.10 <0.01

Acanthomvsis pseudomacronsis (mysid) 0.30 0.01

Holmesiella anomala (mysid) 0.60 0.05

Metervthrops robusta (mysid) 3.40 0.10

Neomvsis czerniawskii (mysid) 0.50 <0.01

Neomysis rayii 0.40 0.09

Pseudomma truncatum (mysid) 0.60 0.03

Cumacea (cumacean unidentified) 1.60 0.08

Leucon sp. (cumacean) 1.90 0.03

Isopoda (isopod) 0.30 0.06

Gammaridea (unidentified) 16.00 0.80

Ampeliscidae (amphipod) 0.90 0.05

Themisto sp. (amphipod) 42.20 1.73

Euphausiacea (unidentified) 79.30 38.49

Thvsanoessa inermis (euphausiid) 0.40 0.27

Thvsanoessa rachii (euphausiid) 0.10 0.01

Thvsanoessa sninifera (euphausiid) 0.30 0.01

Caridea (unidentified) 27.20 9.55

Pasinhaea pacifica (shrimp) 1.30 2.34

Hippolytidae (unidentified) 0.70 0.10

Spirontocaris arcuata (shrimp) 0.20 0.04

Eualus barbata (shrimp) 0.10 0.03

Eualus avinus (shrimp) 0.40 0.10

Pandalidae (unidentified) 18.10 13.52

Pandalus borealis (northern shrimp) 2.70 3.92

Pandalus goniurus (shrimp) 0.10 0.03

Pandalus iordani (shrimp) 0.60 0.67

Pandaloosis sp. (shrimp) 1.00 0.49

Crangonidae (unidentified) 6.40 1.98
11

Table l-l. --Continued.

Prey Name

Crangon communis (shrimp)

Paguridae (hermit crab)

% Freq. % Total

occur. weight

0.80 0.14

13.40 0.72

Chaetognatha (arrow worm)

Larvacean (tunicate)

Teleostei (unidentified fish)

Non-gadoid fish remains

Cluoea harengus pallasi (Pacific herring)

Mallotus villosus (capelin)

Thaleichthys pacificus (eulachon)

Bathylagidae (deepsea smelts)

Chauliodontidae (viperfishes)

Stenobrachius leucoosarus

(northern lampfish)

Gadidae (unidentified gadid)

Gadus macrocephalus (Pacific cod)

Theragra chalcogramma (walleye, pollock)

Zoarcidae (unidentified eelpout)

Scorpaenidae (unidentified)

Cottidae (sculpin)

Cyclopteridae (snailfish)

Stichaeidae (prickleback)

Lumoenus fabricii (slender eelblenny)

Lumoenus maculatus (daubed shanny)

Poroclinus rothrocki

(whitebarred prickleback)

Lyconectes aleutensis (dwarf wrymouth)

Ammodvtes hexaoterus (Pacific sand lance)

Pleuronectidae (unidentified flatfish)

Atheresthes stomias (arrowtooth flounder)

Hipposlossoides elassodon (flathead sole)

Microstomus pacificus (Dover sole)

Reinhardtius hipposlossoides

(Greenland halibut)

Misc. unidentified materials

Total prey weight 2,746 g

Number of stomachs with food 985

Number of empty stomachs 74

1.80 0.01

1.30 0.01

6.80 0.88

3.00 0.46

0.10 0.20

3.20 12.90

0.50 0.04

0.10 <0.01

0.10 0.01

0.50 0.01 

0.50 0.07

0.30 0.01

4.30 2.05

0.60 0.20

0.20 0.03

0.60 0.06

1.90 0.53

0.60 0.12

0.10 0.07

0.50 0.28

0.10 0.03

0.10 0.16

0.50 0.20

0.40 0.02

0.60 0.04

0.40 0.23

0.10 0.54

0.10

0.30

<0.01

0.07
12

were frequently found in pollock stomachs, they were relatively

less important as pollock food. Table l-l also demonstrates the

high diversity of prey fish found in pollock stomachs. Among

which, the commercially important fish, such as Pacific herring

(Clunea oallasi), Pacific cod (Gadus macrocephalus), walleye

pollock, arrowtooth flounder (Atheresthes stomias), flathead

sole (Hippoglossoides elassodon), Dover sole (Microstomus

pacificus), and Greenland halibut (Reinhardtius hippoglossoides)

were found. Other important prey fish like capelin (Mallotus

villosus), eulachon (Thaleichthys pacificus), and Pacific sand

lance (Ammodytes hexapterus) were also found. Overall, capelin

(12.9% by weight of the total stomach contents) was the most

important prey fish of walleye pollock in the Gulf of Alaska.

Variation of Diet Based on Predator Size

Figure l-l illustrates the main prey items of walleye

pollock by predator FL. Calanoids and amphipods were mainly

consumed by smaller-sized (<40 cm) pollock. Euphausiids

comprised the largest portion of pollock food through all size

groups. Shrimp comprised the second largest portion of pollock

food in all but the smallest (<20 cm) size group. Capelin was

important food for pollock larger than 40 cm. It comprised 25%

of the food for the size group greater than or equal to 60 cm.

For all size groups, walleye pollock only consumed very small

amounts (<5%) of prey pollock.

Sizes of the Commercially Important Prey Consumed

Prey size data of walleye pollock were divided into three
13

Figure l-l. --Variations in the main food items of

walleye pollock, by predator size,

in the Gulf of Alaska in 1990.

N = sample size.
14

predator size groups (<40 cm, 40-49 cm, and 150 cm) for analysis.

The size frequencies of the walleye pollock, capelin, and Pacific

sand lance consumed by walleye pollock are shown in Figures l-2

and l-3. Figure l-2 shows that the two larger size groups of

walleye pollock consumed only age-0 walleye pollock

(approximately less than 140 mm SL). These age-0 pollock sizes

had a mean and SD of 43.7 + 24.6 mm SL and ranged from 14 to

110 mm SL. No measurable juvenile walleye pollock were found in

pollock less than 40 cm FL. Figure l-4 shows the size

relationship between walleye pollock as predator and as prey.

Capelin were primarily consumed by the two larger size groups

(40-49 cm and >50 cm) of pollock (Fig. l-3). They had a mean SL

of 95.2 + 23.7 mm and a range from 43 to 140 mm SL. Pacific sand

lance were only consumed by pollock greater than or equal to

40 cm (Fig. l-3), and they had a mean SL of 40.8 + 21.7 mm and a

range from 28 to 118 mm SL. Walleye pollock also consumed

Pacific cod, arrowtooth flounder, Greenland halibut (Reinhardtius

hippoglossoides), flathead sole (Hippoqlossoides elassodon),

Dover sole (Microstomus pacificus), snailfish (Cyclopteridae),

and daubed shanny (Lumpenus maculatus). The number, mean SL, and

the SD of these prey fish are listed in Table l-2.

Geographic Distributions of the Prey Consumed

Figures l-5 to l-7 show the percentage by weight of prey

pollock, capelin, and pandalid shrimp consumed by walleye pollock

in different locations. Figure l-5 shows that small amounts of

pollock were consumed northeast of Kodiak Island. Large amounts

of capelin were primarily consumed northeast of Kodiak Island
15

Figure l-2. --Size frequency distributions of prey

walleye pollock consumed by two size

groups of walleye pollock in the

Gulf of Alaska in 1990.
Figure 1-3 .--Size frequency distributions of capelin and Pacific

sand lance consumed by different size groups of

walleye pollock in the Gulf of Alaska in 1990.
Figure l-4. --Relationship between the predator walleye pollock fork

length (cm) and the prey walleye pollock standard

length (mm) in the Gulf of Alaska in 1990.
18

Table l-2. --Mean standard length and standard deviation of the

miscellaneous prey fish consumed by walleye pollock

in the Gulf of Alaska in 1990.

Prey name No. measured Mean (mm) SD (mm)

Pacific cod 3 16.4 2.5

Arrowtooth flounder 5 26.5 9.3

Greenland halibut 1 23.0 0.0

Flathead sole 4 50.5 19.8

Dover sole 1 114.8 0.0

Snailfish 78 20.3 5.3

Daubed shanny 12 62.1 17.5

20

(Fig. 1-6). Pandalid shrimp were consumed both northeast and

west of Kodiak Island (Fig. l-7).

DISCUSSION

In my analysis, I compared the diets of walleye pollock from

the Gulf of Alaska to those reported by Livingston (1991) for

pollock from the eastern Bering Sea. Table l-3 lists the

percentage by weight of the major prey categories for pollock

from the two study areas. This data shows that prey pollock,

(comprised 41% by weight) was the main food of pollock in the

Bering Sea area, whereas pollock comprised only 2% of the total

food in the Gulf of Alaska area. On the other hand, capelin were

consumed by pollock in the Gulf of Alaska area (about 13% by

weight), whereas no capelin were found in pollock stomachs in the

eastern Bering Sea area. Another difference is that pollock

consumed more miscellaneous fish species (n=22) in the Gulf of

Alaska area than in the eastern Bering Sea (n=7). As for the

crustacean consumed, euphausiids comprised a high percentage of

the stomach contents in both the eastern Bering Sea (25%) and in

the Gulf of Alaska (39%). However, pollock consumed more

pandalids in the Gulf of Alaska area (19%) than in the eastern

Bering Sea (4%).

Smith et al. (1978) did a food habits study of benthic and

demersal fishes in the northern Gulf of Alaska and Clausen (1983)

did a pollock food habits study in Southeast Alaska. Their

studies also showed that large amounts of euphausiids (87% in

Smith's study and 17% in Clausen's study) and shrimp (14% in
Figure l-6. --Geographic distribution of capelin consumed by

walleye pollock in the Gulf of Alaska in 1990.
Figure l-7. --Geographic distribution of Pandalid shrimp consumed

by walleye pollock in the Gulf of Alaska in 1990.
23

Table l-3. --Percent by weight of the major prey categories of

walleye pollock in the eastern Bering Sea and in the

Gulf of Alaska. EBS, eastern Bering Sea (from

Livingston 1991); GOA, Gulf of Alaska (from this

study).

Prey name EBS GOA

Copepoda 6.3 0.8

Mysid 1.4 1.1

Amphipod 1.0 2.6

Euphausiid 24.6 38.8

Pandalid 3.8 18.6

Crangonid 0.9 2.1

Pasiphaeid 0.0 2.3

Chionoecetes sp. 0.1 0.0

Capelin 0.0 12.9

Pollock 41.0 2.1

Miscellaneous fish 5.6 15.0

Unidentified fish 7.7 0.9
2 4

Clausen's study) were consumed by pollock. However, in these

studies, 'the authors either reported a large amount of

unidentified digested materials (26% in Clausen's study) or the

prey fish consumed (less than 10% in both studies) by pollock

were only listed as being teleost.
25

CITATIONS

Clausen, D. M. 1983. Food of walleye pollock, Theragra

chalcogramma in an embayment of southeastern Alaska. Fish.

Bull., U.S. 81:637-642.

North Pacific Fishery Management Council. 1991. Introduction,

p. l-27. In Stock assessment and fishery evaluation report

for the 1992 Gulf of Alaska groundfish fishery. November

1991, prepared by the Gulf of Alaska Groundfish Plan Team,

North Pacific Fishery Management Council, P.O. Box 103136,

Anchorage, AK 99510.

Livingston, P. A. 1991. Walleye pollock, p. 9-30. In P. A.

Livingston (ed.), Groundfish food habits and predation on

commercially important prey species in the eastern Bering Sea

from 1984 to 1986. U.S. Dep. Commer., NOAA Tech. Memo. NMFS

F/NWC-207.

Smith, R. L., A. C. Paulson, and J. R. Rose. 1978. Food and

feeding relationships in the benthic and demersal fishes of

the Gulf of Alaska and Bering Sea. In Environmental

assessment of the Alaskan Continental Shelf. Final Rep.,

Biol. Stud. 1:33-107. Available Arctic Environ. Assess.

Cent., 222 W. 8th Ave. No. 56, Anchorage, AK 99513.
26

PACIFIC COD

Pacific cod (Gadus macrocephalus), with an exploitable

biomass of 387,658 t (Zenger and Thompson 1991), ranks fourth in

abundance in the Gulf of Alaska groundfish community. The

landings of Pacific cod have been increasing in recent years (up

to about 66,000 t in 1990). Pacific cod feed both in the water

column and in benthic areas; hence, they have a high variety of

prey in their diets, including several commercially important

fish and crabs.

RESULTS

General Diets

A total of 916 Pacific cod stomachs were analyzed, of which

892 (97%) contained food. Pacific cod length ranged from 7 to

90 cm FL with a mean and SD of 54.7 cm and + 12.1 cm,

respectively. The average depth of the 78 haul locations was

127 + 39 m with a range from 20 to 205 m. The long list of prey

items presented in Table 2-l shows that Pacific cod is an

opportunistic feeder. They ate many different invertebrates and

various fish (54% and 33% of the total stomach content weight,

respectively). The remaining 13% of the Pacific cod diet was

made up of fishery discards (processed fish carcasses).

Polychaetes were the most frequently found invertebrate in the

stomachs but they were not very important food of Pacific cod in

terms of the percentage of the total stomach content weight.

Octopus and pandalid shrimp were relatively more important.

Pacific cod also consumed decorator crabs (Oregonia spp.), and
27

Table 2-1. --Prey items (expressed in percent frequency of

occurrence, and percent total weight) of Gadus

macrocephalus (Pacific cod) collected in the Gulf of

Alaska in 1990.

Prey Name

% Freq.

occur.

% Total

weight
28

Table 2-1. --Continued.

% Freq. % Total

Prey Name occur. weight

Eualus stoneyi (shrimp) 0.10 <0.01

Eualus avinus (shrimp) 3.00 0.04

Heptacarpus sp. 0.10 0.01

Pandalidae (unidentified) 38.00 6.34

Pandalus borealis (northern shrimp) 6.10 1.63

Pandalus goniurus (shrimp) 1.20 0.37

Pandalus jordani (shrimp) 0.10 <0.01

Pandalus montasui tridens (shrimp) 1.30 0.31

Pandaloosis sp. (shrimp) 3.30 0.22

Pandaloosis dispar (shrimp) 0.10 0.19

Crangonidae (unidentified) 39.50 2.56

Cranson alaskensis (shrimp) 0.10 0.03

Cranson stvlirostris (shrimp) 0.10 0.01

Crangon dalli (shrimp) 0.30 <0.01

Crangon communis (shrimp) 16.70 1.68

Rhvnocranson alata (shrimp) 0.40 0.01

Arsis sp. (shrimp) 5.30 0.29

Arais lar (shrimp) 0.70 0.13

Argis dentata (shrimp) 0.20 0.02

Argis ovifer (shrimp) 0.20 0.02

Argis alaskensis (shrimp) 0.10 <0.01

Metacrangon munita (shrimp) 0.40 0.02

Anomura (unidentified) 0.60 0.03

Paguridae (hermit crab) 20.40 3.72

Pacrurus aleuticus (hermit crab) 0.20 0.03

Pasurus rathbuni (hermit crab) 0.30 0.03

Elassorhirus sp. (hermit crab) 0.10 0.01

Elassachirus tenuimanus (hermit crab) 0.30 0.02

Acantholithodes hisnidus (fussy crab) 0.10 0.22

Lopholithodes foraminatus (box crab) 0.40 1.68

Munida auadrispina (pinch bug) 1.90 0.16

Majidae (unidentified) 2.80 0.23

Oresonia sp. (decorator crab) 0.40 0.01

Oreqonia sracilis (decorator crab) 1.60 0.16

Oresonia bifurca (decorator crab) 0.10 <0.01

Hyas sp. (lyre crab) 3.60 1.07

Hyas lyratus (lyre crab) 3.10 0.94

Hyas coarctatus (lyre crab) 0.20 0.01

Chionoecetes sp. (unidentified) 6.20 0.84

Chionoecetes opilio (snow crab) 0.30 0.05

Chionoecetes bairdi (Tanner crab) 34.60 10.77

Erimacrus isenbeckii (crab) 0.10 0.01

Cancer sp. (crab) 0.30 0.01

Cancer oresonensis (pygmy cancer crab) 2.10 0.10

Pinnotheridae (pea crab) 5.40 0.33
29

Table 2-l. --Continued.

Prey Name

% Freq. % Total

occur. weight

Pinnixa sp. (pea crab)

Sipuncula (peanut worm)

Echiura (marine worm)

Asteroidea (starfish)

Ophiurida (brittle star)

Teleostei (unidentified fish)

Non-gadoid fish remains

Cluoea pallasi (Pacific herring)

Mallotus villosus (capelin)

Thaleichthvs oacificus (eulachon)

Gadidae (unidentified gadid)

Therasra chalcosramma (walleye pollock)

Zoarcidae (unidentified eelpout)

Lvcodes brevioes (shortfin eelpout)

Cottidae (sculpin)

Hemilenidotus sp. (sculpin)

Dasvcottus setiser (sculpin)

Icelinus borealis (sculpin)

Agonidae (poacher)

Asterotheca alascana (poacher)

Asterotheca oentacanthus (poacher)

Sarritor frenatus (poacher)

Cyclopteridae (unidentified snailfish)

Eumicrotremus orbis (snailfish)

Bathymaster siqnatus (searcher)

Stichaeidae (unidentified)

Lumpenus fabricii (slender eelblenny)

Lumpenus maculatus (daubed shanny)

Poroclinus rothrocki

(whitebarred prickleback)

Lyconectes aleutensis (dwarf wrymouth)

Ammodvtes hexapterus (Pacific sand lance)

Pleuronectidae (unidentified flatfish) 1

Atheresthes stomias (arrowtooth flounder)

Errex zachirus (rex sole)

Hipooqlossoides elassodon (flathead sole)

Pleuronectes bilineatus (rock sole)

Hippoalossus stenolepis (Pacific halibut)

Fishery discards

Misc. unidentified materials

5.30 0.27

0.70 0.08

5.00 0.65

0.80 0.04

0.90 0.02

18.00 6.81

9.90 0.55

0.30 0.37

5.00 1.91

0.20 0.26

1.80 0.15

5.00 7.43

1.60 1.34

1.20 0.51

9.40 0.97

0.10 0.55

0.20 0.86

0.10 <0.0l

2.60 0.07

0.10 <0.01

0.10 0.02

0.80 0.06

0.60 <0.01

0.10 0.02

0.80 0.73

8.40 0.74

0.10 0.01

1.00 0.09

0.60

1.70

1.20

4.80

2.00

0.10

2.70

0.30

0.20

2.90

0.90

Total prey weight 20,872 g

Number of stomachs with food 892

Number of empty stomachs 24

0.29

0.34

0.34

1.25

5.80

0.06

1.74

0.33

0.01

12.51

0.03
30

lyre crabs (Hyas spp.). However, Tanner crab (Chionoecetes

bairdi) comprised the highest proportion (11% of the total

stomach content weight) of the invertebrates consumed. Hermit

crabs, euphausiids, amphipods, and marine worms were also

invertebrates consumed by Pacific cod. Of the fish consumed by

Pacific cod, walleye pollock (Theragra chalcogramma) comprised

the highest proportion (7%) of the total stomach contents weight.

Other commercially important prey fish include arrowtooth

flounder (Atheresthes stomias),. capelin (Mallotus villosus),

flathead sole (Hippoglossoides elassodon), rock sole

(Pleuronectes bilineatus), Pacific halibut (Hippoglossus

stenolepis), Pacific herring (Clupea Dallasi), eulachon

(Thaleichthys pacificus), Pacific cod, Pacific sand lance

(Ammodytes hexapterus), and rex sole (Errex zachirus). Pacific

cod also consumed many non-commercially important prey fish, such

as zoarcids, cottids, agonids, cyclopterids, bathymasterids, and

stichaeids.

Variation of Diet Based on Predator Size

Figure 2-l shows that Pacific cod ate more prey fish with

increasing cod size (FL), especially for cod greater than or

equal to 70 cm FL. The fish consumed (including fish and fishery

discards) comprised about 75% of the total stomach contents in

this size group. Only this group of larger Pacific cod (270 cm

FL) consumed a significant amount (17%) of walleye pollock. All

other smaller size groups consumed very small amounts of pollock

(<5%). All but the largest size groups consumed a fair amount of

shrimp (>15%). Tanner crabs were consumed by Pacific cod between
31

Figure 2-1. --Variations in the main food items of

Pacific cod, by predator size, in

the Gulf of Alaska in 1990.

N = sample size.
32

40 and 70 cm FL. The smallest size group (<20 cm FL) of cod ate

a large amount (45%) of miscellaneous prey (mainly amphipods and

mysids).

Sizes of the Commercially Important Prey Consumed

The commercially important prey consumed by Pacific cod were

analyzed by three predator size groups (<30 cm, 30-59 cm, and

>60 cm FL). Because of insufficient 'measurable' prey data in the

size group less than 30 cm, this group was not included in the

analysis. Examination of Figure 2-2 suggests that the walleye

pollock consumed by 30-59 cm Pacific cod were mainly age-0 fish

plus some age-l fish, whereas Pacific cod greater than or equal

to 60 cm consumed age-O, 1, 2, and 3+ walleye pollock. The body

size relationship between Pacific cod and the pollock they

consumed is shown in Figure 2-3. The mean SL of pollock consumed

by Pacific cod was 161.5 + 133.0 mm with a range from 33 to

468 mm. Figure 2-2 also shows that Pacific sand lance were

primarily consumed by Pacific cod 30-59 cm long. The mean SL of

all Pacific sand lance consumed by Pacific cod was 94.4 +. 32.7 mm

with a range from 46 to 165 mm. The mean SL of the capelin

consumed was 78.9 + 14.0 mm with a range from 57 to 125 mm.

Figure 2-4 shows that capelin were mainly consumed by Pacific cod

30-59 cm long. The length distributions of flathead sole and

arrowtooth flounder consumed by Pacific cod are shown in Figure

2-5. Larger predators fed on larger prey. The mean SL of

flathead sole consumed by Pacific cod was 80.5 + 46.2 mm, with a

range from 17 to 220 mm, The mean SL of arrowtooth flounder

consumed was 153.2 + 135.3 mm with a range between 33 to 390 mm.
33

Figure 2-2.--Size frequency distributions of walleye pollock and

Pacific sand lance consumed by two size groups of

Pacific cod in the Gulf of Alaska in 1990.
Figure 2-3. --Scatterplot of walleye pollock prey size versus

Pacific cod in the Gulf of Alaska in 1990.
35

Figure 2-4. --Size frequency distributions of Tanner crab and

capelin consumed by two size groups of Pacific

cod in the Gulf of Alaska in 1990.
36

Figure 2-5 .--Size frequency distributions of flathead sole and

arrowtooth flounder consumed by two size groups of

Pacific cod in the Gulf of Alaska in 1990.
3 7

In general, Pacific cod greater than or equal to 60 cm fed on

larger Tanner crabs than cod 30-59 cm in length (Fig. 2-4). The

average carapace width (CW) of the Tanner crabs consumed was 19.8

+ 9.4 mm with a range of 3 to 99 mm. Most of the Tanner crabs

measured from Pacific cod stomach contents were age-l crab (9-

34 mm cw). Some commercially important prey fish were not

analyzed in detail because of insufficient data. For example,

only two Pacific halibut were consumed (21 and 60 mm SL) by

Pacific cod in this study. Pacific cod also consumed one rex

sole (121 mm SL), two eulachon (132 and 180 mm SL), and three

rock sole (95, 140, and 150 mm SL).

Geographic Distributions of the Prey Consumed

Figures 2-6 to 2-15 illustrate the geographic distributions

of the commercially important prey (percent by weight) consumed

by Pacific cod in each haul where stomach samples were collected.

Consumption of walleye pollock by Pacific cod was not

widespread (26 out of 78 hauls) (Fig. 2-6). In addition, the

proportion of walleye pollock in the stomach content weights at

each station was not high. Only in the area southwest of Kodiak

Island did pollock make up a higher percentage (>75%) of prey

found in the stomach content samples.

The consumption of arrowtooth flounder by Pacific cod was

also found mainly in the area southwest of Kodiak Island

(Fig. 2-7).

Figures 2-8, 2-9, and 2-10 illustrate that the consumption

of Pacific halibut, rock sole, and rex sole by Pacific cod was

not significant in the Gulf of Alaska area. Compared to the
Figure 2-6. --Geographic distribution of walleye pollock consumed

by Pacific cod in the Gulf of Alaska in 1990.




43

three flatfish mentioned above, flathead sole were eaten more

frequently by the Pacific cod and were found in the area

southwest of Kodiak Island (Fig. 2-11).

Capelin seemed to be consumed evenly throughout the sampling

area. There was only one station where capelin comprised more

than 75% of the stomach contents of Pacific cod (Fig. 2-12).

Pacific sand lance were consumed primarily in the area

southwest of Kodiak Island, although there was one station to the

north of Kodiak Island where Pacific sand lance comprised more

than 50% of the stomach contents (Fig. 2-13).

Tanner crab consumption was widespread in all the areas

sampled, although the higher percent by weight of Tanner crab in

stomach contents seemed to be focused only on the east side of

Kodiak Island (Fig. 2-14).

Fishery discards were consumed by Pacific cod mainly in the

area southwest of Kodiak Island (Fig. 2-15).

DISCUSSION

Previous studies indicated that fish, crab, and shrimp were

the main food components of Pacific cod found in the Gulf of

Alaska (Jewett 1978, Clausen 1981). Jewett found that, in the

Kodiak area, walleye pollock was the most common (3.9% frequency

of occurrence) prey fish eaten by cod, followed by some

unidentified flatfish and Pacific sand lance. Jewett also

reported that Tanner crabs occurred in 36.7% and pandalid shrimp

occurred in 4.7% of the Pacific cod stomachs sampled. In

Clausen's (1981) study, Pacific herring and pollock (9.6 and 8.7%
165 OOW 158 OOW 151 oow J44 oow 137 oow

Figure 2-11. --Geographic distribution of flathead sole consumed by

Pacific cod in the Gulf of Alaska in 1990.


Figure 2-14.--Geographic distribution of Tanner crab consumed by

Pacific cod in the Gulf of Alaska in 1990.

49

frequency of occurrence, respectively) were the most common prey

fish found in cod stomachs and Tanner crabs (26.2%) were the most

common prey of Pacific cod in Southeast Alaskan waters. Pandalid

shrimp were found in 25.4% of the cod stomachs sampled. The main

difference between the present study and earlier studies in the

Gulf of Alaska area is that fishery discards now comprise a

significant proportion (13%) of the total stomach content weights

in this study, whereas no fishery discards were reported as food

in the earlier studies. Another significant difference is that

in both Jewett (1978) and Clausen's (1981) studies, reported a

high percentage (35%) of unidentified fish, whereas this

unidentified category was only 7% in this study.

In another comparison, walleye pollock comprised 40% by

weight of the total stomach contents of Pacific cod in the Bering

Sea (Livingston 1991), whereas only 7% of the total stomach

contents were pollock from this study. More prey fish species

were consumed by Pacific cod in the Gulf of Alaska area compared

with the Bering Sea. In addition, king crabs (Paralithodes spp.)

were consumed in the Bering Sea area, but they were not found in

the stomachs of Pacific cod in the Gulf of Alaska area.
50

CITATIONS

Clausen, D. M. 1981. Summer food of Pacific cod, Gadus

macrocephalus, in coastal waters of southeastern Alaska.

Fish. Bull., U.S. 78:968-973.

Jewett, S. C. 1978. Summer food of the Pacific cod, Gadus

macrocephalus, near Kodiak Island, Alaska. Fish.

Bull., U.S. 76:700-706.

Livingston, P. A. 1991. Pacific cod, p. 31-88. In P. A.

Livingston (ed.), Groundfish food habits and predation on

commercially important prey species in the eastern Bering

Sea from 1984 to 1986. U.S. Dep. Commer., NOAA Tech. Memo.

NMFS F/NWC-207.

Zenger, H. H., and G. G. Thompson. 1991. Pacific cod. In Stock

assessment and fishery evaluation report for the 1992 Gulf

of Alaska groundfish fishery. November 1991, prepared by

the Gulf of Alaska Groundfish Plan Team, North Pacific

Fishery Management Council, P.B. Box 103136, Anchorage, AK

99510.
51

ARROWTOOTH FLOUNDER

Arrowtooth flounder (Atheresthes stomias), with an

exploitable biomass of 1,144,242 t in 1990, ranked first in the

total groundfish biomass (North Pacific Fishery Management

Council 1991). However, because of the species' low commercial

value, catches have been low (Wilderbuer and Brown 1990).

Arrow-tooth flounder is a large flatfish with a symmetrical mouth

and sharp teeth that feeds on commercially important fish like

walleye pollock (Theragra chalcogramma), capelin (Mallotus

villosus), and other fish, as well as shrimp and other

crustaceans (Smith et al. 1978). Therefore, with the feeding

behavior and the high abundance of arrowtooth flounder in the

Gulf of Alaska area, it is important to study its food habits and

its impact on commercially important prey.

RESULTS

General Diets

A total of 1,144 arrowtooth flounder stomachs were analyzed.

Of this total, 489 were empty and 655 (57%) contained food.

Arrowtooth flounder sizes ranged from 12 to 80 cm FL with a mean

and SD of 42.1 and +. 10.1 cm, respectively. The average depth of

the 62 hauls where stomachs were collected was 164 + 69 m with a

range from 66 to 432 m.

Table 3-l displays all the prey items found in arrowtooth

flounder stomachs. Even though many invertebrate species like

euphausiids (34.8% by frequency of occurrence), pandalid shrimp,

squid, mysids, and amphipods were found in the diet of arrowtooth
52

Table 3-l. --Prey items (expressed in percent frequency of

occurrence, and percent total weight) of Atheresthes

stomias (arrowtooth flounder) collected in the Gulf

of Alaska in 1990.

Prey Name

% Freq. % Total

occur. weight

Polychaeta (unidentified)

Bivalvia (clam)

Teuthoidea (squid unidentified)

Gonatidae (squid)

Calanoida (copepod)

Mysidae (mysid)

Holmesiella anomala (mysid)

Neomvsis czerniawskii (mysid)

Neomvsis ravii

Cumacea (cumacean)

Isopoda (isopod)

Gammaridea (unidentified)

Ampeliscidae (amphipod)

Themisto sp.

Caprellidea (amphipod)

Euphausiacea (unidentified)

Thvsanoessa inermis (euphausiid)

Thvsanoessa soinifera (euphausiid)

Caridea (unidentified)

Pasinhaea oacifica (shrimp)

Hippolytidae (unidentified)

Eualus avinus (shrimp)

Pandalidae (unidentified)

Pandalus borealis (northern shrimp)

Pandalus iordani (shrimp)

Pandalus montasui tridens (shrimp)

Pandalus olatvceros (shrimp)

Crangonidae (unidentified)

Cranson communis (shrimp)

lar Arsis

Arsis ovifer (shrimp)

Paguridae (hermit crab)

Echiuridae (marine worm)

Chaetognatha (arrow worm)

Teleostei (unidentified fish)

Non-gadoid fish remains

0.80 0.02

0.20 co.01

0.80 0.21

0.20 1.80

0.80 co.01

0.60 0.01

0.20 co.01

0.20 co.01

0.20 co.01

0.30 co.01

0.20 0.02

1.40 0.02

0.30 0.01

0.60 co.01

0.20 co.01

34.80 2.84

4.90 0.15

0.20 co.01

1.40 0.05

0.50 0.11

0.20 0.01

0.70 0.02

12.00 1.67

7.00 1.80

0.80 0.27

0.60 0.15

0.20 0.01

1.40 0.08

1.80 0.12

0.60 0.06

0.20 co.01

0.20 co.01

0.20 0.03

0.60 co.01

2.70 0.06

1.50 0.38

Cluoea harensus oallasi (Pacific herring) 4.40 9.15

Coresonus sp. (salmonidae) 0.20 0.12

Mallotus villosus (capelin) 19.70 8.18

Thaleichthvs pacificus (eulachon) 0.30 0.78

Gadidae (unidentified gadid) 5.50 1.73

Therasra chalcoqramma (walleye pollock) 15.30 . 66.43
53

Table 3-l. --Continued.

Prey Name

% Freq. % Total

occur. weight

Zoarcidae (unidentified eelpout) 0.60

Scorpaenidae (unidentified) 0.40

Pleurosrammus monopteryaius

(Atka mackerel) 0.20

Stichaeidae (prickleback) 1.20

Poroclinus rothrocki

(whitebarred prickleback) 0.30

Ammodytes hexapterus (Pacific sand lance) 0.30

Pleuronectidae (unidentified flatfish) 0.90

Atheresthes stomias (arrowtooth flounder) 0.20

Hippoglossoides elassodon (flathead sole) 0.20

Fishery discard 0.30

Total prey weight

Number of stomachs with food

Number of empty stomachs

6,501 g

655

489

0.36

0.26

0.50

0.33

0.16

0.02

1.21

<0.01

0.26

0.56
54

flounder, they comprised only 10% by weight of the total stomach

contents. The remaining 90% of stomach content weight was made

up of fish. Walleye pollock was the most important prey of

arrowtooth flounder; they comprised 66% by weight of the total

stomach contents. Pacific herring (Clupea pallasi) and capelin

comprised 9% and 8% of the total stomach contents by weight,

respectively. Arrowtooth flounder also consumed some eulachon

(Thaleichthys pacificus), Atka mackerel (Pleurogrammus

monopterygius), Pacific sand lance (Ammodytes hexapterus),

arrowtooth flounder, flathead sole (Hipposlossoides elassodon),

fishery discard, and some non-commercially important species

(zoarcids and stichaeids).

Variation of Diet Based on Predator Size

Figure 3-l shows the percentage by weight of the main prey

items for different arrowtooth flounder size groups. With the

exception of the smallest size group (<20 cm FL), all size groups

had at least 70% fish by weight in their stomach contents.

Walleye pollock were the predominant prey fish (60 to 90%) for

arrowtooth flounder greater than or equal to 40 cm, whereas

capelin predominate in the diets of fish between 20 and 40 cm

long. Pacific herring comprised about 20% of the food for fish

between 30 and 50 cm. Shrimp were the predominant food of fish

less than 20 cm.

Sizes of the Commercially Important Prey Consumed

The commercially important prey consumed by arrowtooth

flounder were analyzed by three different size groups: less than
5 5

Figure 3-1. --Variations in the main food items of

arrowtooth flounder, by predator size,

in the Gulf of Alaska in 1990.

N = sample size.
56

20 cm, 20-39 cm, and greater than or equal to 40 cm FL (Figs. 3-2

and 3-3). Figure 3-2 (left side) shows that pollock were not

consumed by arrowtooth flounder less than 20 cm long. The

medium-sized fish (20-39 cm) consumed some age-0 and age-l

pollock. The larger-sized (>40 cm) arrowtooth flounder contained

many age-O, age-l, and age-2 pollock (approximately <300 mm), and

some age 3 and older pollock. The average SL (mean + SD) of the

pollock consumed by arrowtooth flounder was 196.1 + 89.2 mm with

a range of 21 to 450 mm. The relationship between the arrow-tooth

flounder size and the size of the pollock that they consumed is

shown in Figure 3-4. Figure 3-2 (right side) displays the

consumption of Pacific herring by arrowtooth flounder. It

illustrates that no Pacific herring were found in arrowtooth

flounder smaller than 20 cm long. And the medium-sized and the

larger-sized fish consumed only a few Pacific herring. The mean

length of the 11 Pacific herring consumed was 159.9 + 23.0 mm SL

with a range from 127 to 195 mm SL. Figure 3-3 shows that many

capelin were consumed by arrowtooth flounder larger than 20 cm

long. The average SL of capelin consumed by arrowtooth flounder

was 85.3 + 18.1 mm with a range from 43 to 130 mm SL. Arrowtooth

flounder also consumed one flathead sole (130 mm SL), one Atka

mackerel (128 mm SL), two eulachons (130 and 154 mm SL), and one

Pacific sand lance (70 mm SL).

Geographic Distributions of the Prey Consumed

Figures 3-5, 3-6, and 3-7 describe the geographic

distributions of the pollock, capelin, and Pacific herring

consumed by arrowtooth flounder. It shows that pollock were
57

Figure 3-2.--Size frequency distributions of walleye

pollock and Pacific herring consumed by

two size groups of arrow-tooth flounder

in the Gulf of Alaska in 1990.
Figure 3-3. --Size frequency distributions of capelin

consumed by three size groups of arrowtooth

flounder in the Gulf of Alaska in 1990.
Figure 3-4.-- scatterplot of walleye pollock prey size versus

arrowtooth flounder in the Gulf of Alaska in 1990.
60

primarily consumed around Kodiak Island (Fig. 3-5). Figure 3-6

shows that most capelin consumed by arrowtooth flounder were on

the east and southwest of Kodiak Island. Most Pacific herring

consumed by arrowtooth flounder were located east of the Kenai

Peninsula (Fig. 3-7).

DISCUSSION

Smith et al. (1978) found that similar prey items (pollock,

euphausiids, shrimp, and miscellaneous fish) were consumed by

arrowtooth flounder in the Gulf of Alaska area, although

euphausiids comprised a higher percentage by volume in their

study compared with this study. This discrepancy is probably due

to the fact that 85% of the arrowtooth flounder in their study

were less than or equal to 35 cm long, whereas most fish in our

study were greater than or equal to 40 cm. In the Bering Sea

area, arrowtooth flounder also consumed a high percentage (56% by

weight) of pollock (Yang 1991). The other prey items consumed in

the Bering Sea area were similar to those in the Gulf of Alaska

area. The main difference is that capelin made up 8% by weight

of the diet in the Gulf of Alaska area, whereas less than 1% of

the diet of arrowtooth flounder in the Bering Sea area was

capelin.



64

CITATIONS

North Pacific Fishery Management Council. 1991. Introduction,

P. l-27. In Stock assessment and fishery evaluation report

for the 1992 Gulf of Alaska Groundfish Fishery, prepared by

Gulf of Alaska Groundfish Plan Team, North Pacific Fishery

Management Council. P. 0. Box 103136, Anchorage, AK 99510.

Smith, R. L., A. C. Paulson, and J. R. Rose. 1978. Food and

feeding relationships in the benthic and demersal fishes of

the Gulf of Alaska and Bering Sea. In Environmental

assessment of the Alaskan Continental Shelf. Final Rep.,

Biol. Stud. 1:33-107. Available Arctic Environ. Assess.

Cent., 222 W. 8th Ave. No. 56, Anchorage, AK 99513.

Wilderbuer, T. K. and E. S. Brown. 1990. Flatfish. In Stock

assessment and fishery evaluation report for the 1991 Gulf

of Alaska groundfish fishery. North Pacific Fishery

Management Council, P. 0. Box 103136, Anchorage, AK 99510.

Yang, M-S. 1991. Arrowtooth flounder, p. 143-162. In P. A.

Livingston (ed.), Groundfish food habits and predation on

commercially important prey species in the eastern Bering

Sea from 1984 to 1986. U.S. Dep. Commer., NOAA Tech. Memo.

NMFS F/NWC-207.
65

PACIFIC HALIBUT

Pacific halibut (Hippoglossus stenolenis) had an exploitable

biomass of 216,888 t in the Gulf of Alaska (3A, 3B, and 4A North

Pacific Fishery Management Council Regulatory areas) in 1990 (P.

J. Sullivan, International Pacific Halibut Commission, P.O. Box

95009, Seattle, WA 98145-2009. Pers. commun., September 1993).

The commercial catch in the Gulf of Alaska area in 1990 was

23,836 t (data compiled from International Pacific Halibut

Commission). Because the Pacific halibut feed on fish, crab and

squid, it is important to study their food habits and their

impact on other commercially important species in the Gulf of

Alaska area.

RESULTS

General Diets

A total of 467 Pacific halibut stomachs were analyzed, of

which 79 were empty and 388 (83%) contained food. The size range

of the Pacific halibut was between 11 to 151 cm FL with a mean

and SD of 71.5 + 25.9 cm. The average depth of the 71 haul

locations was 109 + 48 m with a range from 20 to 234 m.

Table 4-l lists the percent of frequency of occurrence, the

percent by number, and the percent by weight of the prey found in

Pacific halibut stomachs. Nearly 80% of the total stomach

contents weight were fish and invertebrates comprised only 20% by

weight. Walleye pollock (Theragra chalcogramma) was the. most

important prey item. They were the most frequently occurring

prey in the stomachs (28%). In terms of weight, they comprised
66

Table 4-l. --Prey items (expressed in percent frequency of

occurrence, numerical percentage, and percent total

weight) of Pacific halibut (Hippoglossus stenolepis)

collected in the Gulf of Alaska in 1990.

'Prey Name

% Freq. % Total

occur. % Number weight
6 7

Table 4-l. --Continued.

Prey Name

% Freq. % Total

occur. % Number weight
68

Table 4-l. --Continued.

Prey Name

% Freq. % Total

occur. % Number weight 

Hippoglossoides elassodon (flathead sole) 0.10 0.12 0.56

Pleuronectes bilineatus (rock sole) 0.30 0.03 0.13

Pleuronectes asper (yellowfin sole) 0.30 0.06 0.01

Microstomus pacificus (Dover sole) 0.30 0.03 0.03

Hippoglossoides stenolepis (Pacific halibut) 0.80 0.09 <0.01

Fishery discards 2.10 0.47 7.07

Total prey count

Total prey weight

Number of stomachs with food

Number of empty stomachs

3,383

50,417 g

388

79
69

57% of the total stomach contents. Coho salmon (Oncorhynchus

kisutch) made up 5% of the total stomach contents by weight, but

they did not often occur in the Pacific halibut sampled. Capelin

(Mallotus villosus) were consumed quite frequently (14%) but they

comprised less than 1% of the total stomach contents weight.

Other commercially important fish consumed by Pacific halibut

include Pacific cod (Gadus macroceDhalus), eulachon (Thaleichthvs

pacificus), Pacific sand lance (Ammodvtes hexapterus), Pacific

ocean perch (Sebastes alutus), arrowtooth flounder (Atheresthes

stomias), flathead sole (Hippoqlossides elassodon), rock sole

(Pleuronectes bilineatus), yellowfin sole (Pleuronectes asper),

Dover sole (Microstomus pacificus), and young Pacific halibut.

Pacific halibut also consumed 7%, by weight, of fishery discards

(processed fish carcasses). Many non-commercially important fish

(cottids, agonids, cyclopterids, bathymasterids, and stichaeids)

were also consumed by Pacific halibut; however, they played a

minor role as food of the Pacific halibut.

Tanner crabs (Chionoecetes bairdi) were the most important

invertebrate prey of Pacific halibut; they comprised about 6% of

the total stomach contents weight. Less important invertebrate

prey included octopus (4.8%), lyre crab (Hyas lyratus) (2.7%),

cancer crab (Cancer oreqonensis) (2%), hermit crab (1.3%), and

decorator crab (Oresonia gracilis) (1%). All of the other

invertebrates consumed (polychaetes, gastropods, clams, squid,

shrimp, and other crustaceans) were found less frequently and

were considered to be relatively less important as food of

Pacific halibut.
70

Variation of Diet Based on Predator Size

The diet of Pacific halibut varies greatly by predator size

(Fig. 4-l). Miscellaneous prey fish (mainly capelin, Pacific

sand lance, cottids, zoarcids, stichaeids, and flatfish) were

important food for Pacific halibut between 30 and 70 cm FL long,

whereas walleye pollock were mainly consumed by Pacific halibut

greater than or equal to 80 cm FL. Crabs (Tanner crabs,

decorator crabs, lyre crabs, and cancer crabs) were the main food

of Pacific halibut between 50 and 80 cm FL (they comprised more

than 40% by weight of the total stomach contents weight in this

size group). Hermit crabs (pagurids) were important food (225%

by weight) for Pacific halibut less than 60 cm FL, especially for

fish less than 30 cm long. Cephalopods (mainly octopus) were

important food for fish less than 40 cm FL.

Sizes of the Commercially Important Prey Consumed

The prey length data of the Pacific halibut were divided by

predator size into two groups (<80 cm and 280 cm FL) for analysis

(Figs. 4-2 to 4-4). Figure 4-2 (left) shows that walleye pollock

consumed by Pacific halibut less than 80 cm were mainly age-0

(<140 mm SL) fish plus a few age-l to age-3 fish. The

larger-sized Pacific halibut (<80 cm) consumed many different

sizes of walleye pollock including both pre-recruits (<300 mm SL)

and recruits (>300 mm SL). Figure 4-2 (top left) shows that most

pre-recruit pollock consumed were age-l and age-2 fish, whereas

most recruited pollock consumed were age-3 and age-4 fish. A few

large pollock (250 cm SL) were consumed by some Pacific halibut

larger than 100 cm FL; one 151 cm female Pacific halibut consumed
71

Figure 4-1. --Variation in the main food items of

Pacific halibut, by predator size,

in the Gulf of Alaska in 1990.

N = sample size.
7 2

Figure 4-2 .--Size frequency distributions of walleye

pollock and Tanner crabs consumed by two

size groups of Pacific halibut in the

Gulf of Alaska in 1990.
73

a 670 mm walleye pollock. The average (mean + SD) standard

length of pollock consumed by Pacific halibut was 315.9 +

155.0 mm with a range from 34 to 670 mm long. The relationship

between the Pacific halibut size and the size of the pollock they

consumed is shown in Figure 4-5.

The size of Chionoecetes bairdi consumed by Pacific halibut

less than 80 cm (Fig. 4-2, top right) were mainly age-0 (<9 mm

CW) and age-l (9-34 mm CW) crabs, whereas large Pacific halibut

(180 cm) consumed mainly age-l C. bairdi and some age-2 and older

crabs (Fig. 4-2, bottom right). The mean CW of the C. bairdi

consumed 24.7 + 18.0 mm with a range from 3 to 97 mm.

Figure 4-3 shows that large Pacific halibut consumed more

larger-sized arrowtooth flounder. Consumption of capelin

occurred mainly in smaller (<80 cm FL) Pacific halibut, and only

a few were found in large (280 cm) Pacific halibut (Fig. 4-4,

left). The mean SL of the capelin consumed was 78.8 + 15.3 mm

with a range from 46 to 108 mm. Figure 4-4 (right) shows that

the Pacific sand lance consumed by Pacific halibut were mainly

between 100 and 150 mm SL. They had a mean SL of 136.5 + 26.5 mm

with a range from 55 to 187 mm.

Pacific halibut also consumed two coho salmon (450 and

500 mm SL, respectively), one Pacific cod (380 mm SL), one

eulachon (120 nun SL), one Pacific ocean perch (208 mm SL), four

flathead sole (58, 101, 147, and 280 mm SL), one rock sole

(160 mm SL), two yellowfin sole (61, 64 mm SL), one Dover sole

(105 mm SL.), and two Pacific halibut larvae (21 and 30 mm SL).
74

Figure 4-3.--Size frequency distributions of arrowtooth

flounder consumed by two size groups of

Pacific halibut in the Gulf of Alaska in 1990.
7 5

Figure 4-4. --Size frequency distributions of capelin

and Pacific sand lance consumed by two

size groups of Pacific halibut in the

Gulf of Alaska in 1990.
Figure 4-5. --Relationship between the Pacific halibut fork length

(cm) and the walleye pollock standard length (mm) in

the Gulf of Alaska in 1990.
77

Geographic Distributions of the Prey Consumed

The distributions of the commercially important prey

consumed by Pacific halibut are shown in Figures 4-6 to 4-14.

Figure 4-6 illustrates that walleye pollock were consumed by

Pacific halibut at most of the survey stations, and the percent

by weight of the pollock in most of the stations was high (>75%).

Capelin were consumed by Pacific halibut primarily in the areas

to east and southeast of Kodiak Island (Fig. 4-7). The amount of

capelin consumed in each station was low (<25%). Tanner crabs

were also consumed in the Kodiak area (Fig. 4-8). Pacific sand

lance were only found in samples from areas southwest of Kodiak

Island (Fig. 4-9). Figure. 4-10 shows that the arrowtooth

flounder consumed by Pacific halibut were found mainly around the

Kodiak area. The locations where Pacific halibut consumed coho

salmon, Pacific cod, and Dover sole are shown in Figures 4-11 to

4-13, respectively. Figure 4-14 shows the locations where

fishery discards (mainly fish carcasses) were consumed by Pacific

halibut.

DISCUSSION

The earlier food habits studies of Pacific halibut showed

that the diet was primarily composed of fish, crustaceans, and

mollusks (Novikov 1968). Novikov found that, in the Bering Sea

area, the frequency of occurrence and the species composition of

the diet depended on the locations inhabited and the size of the

Pacific halibut. The smaller fish (<30 cm) fed mainly on

crustaceans. The medium-sized fish (30-60 cm) switched to feed
Figure 4-6. --Geographic

by Pacific

distribution of walleye pollock consumed

halibut in the Gulf of Alaska in 1990.
Figure 4-7 .--Geographic distribution of capelin consumed by

Pacific halibut in the Gulf of Alaska in 1990.
165 OOW 158 OOW I51 oow 144 oow 137 oow

Figure 4-0. --Geographic distribution of Tanner crabs

(Chionoecetes bairdi) consumed by Pacific halibut in

the Gulf of Alaska in 1990.






87

on larger crustaceans and fish. Pacific halibut larger than

60 cm fed mainly on fish. Novikov also pointed out the

importance of some flatfish as prey of Pacific halibut,

especially the connection between the yellowfin sole and the

Pacific halibut. Livingston et al. (1993) also reported that, in

addition to the large amount of walleye pollock, Tanner crabs,

and snow crabs (Chionoecetes opilio) consumed, Pacific halibut

also consumed relatively high proportions of Pacific cod,

yellowfin sole, and flathead sole in some areas in the eastern

Bering Sea.

Best and Pierre (1986) analyzed the stomach contents of

Pacific halibut collected in the Gulf of Alaska and characterized

the species as opportunistic feeders. When Pacific halibut are

young, they feed mainly on crustaceans. The Pacific halibut's

diet changes to primarily fish as they grow and increase in size.

They found that walleye pollock, Pacific sand lance, octopus, and

Tanner crabs were the main food of the Pacific halibut.
88

CITATIONS

Best, E. A., and G. St-Pierre. 1986. Pacific halibut as

predator and prey. Technical Report No. 21. International

Pacific Halibut Commission, P.O. Box 95009, Seattle, WA

98145-2009.

Livingston, P. A., A. Ward, G. M. Lang, and M-S. Yang. 1993.

Groundfish food habits and predation on commercially

important prey species in the eastern Bering Sea from 1987

to 1989. U.S. Dep. Commer., NOAA Tech. Memo. NMFS-AFSC-11,

192 p.

Novikov, N. P. 1968. Basic elements of the biology of the

Pacific halibut (Hippoglossoides Hippoglossoides stenolepis

Schmidt) in the Bering Sea. In: Sov. Fish. Inv. N. Pac. Pt.

II, 175-219. Isr. Prog. Sci. Transl. TT67-51204. Available

from the Department of Commerce, Clearinghouse for Federal

Scientific and Technical Information, Springfield, VA 22151.
89

SABLEFISH

Sablefish, Anoplonoma fimbria, ranked sixth in total

groundfish biomass in the Gulf of Alaska in 1990 with an

exploitable biomass of 226,000 t (North Pacific Fishery

Management Council 1991). Annual catches of sablefish from 1986

to 1990 ranged from 20,000 to 30,000 t (Fujioka 1990). Sablefish

has been found to feed opportunistically on fish, crustaceans,

and worms (Grinols and Gill 1968). Because sablefish is an

important commercial species and it probably feeds on some other

commercially important fish, it is included in this study on food

habits and potential impacts on the other marine fishes in the

Gulf of Alaska.

RESULTS

General Diets

A total of 331 sablefish stomachs were analyzed, of which 92

were empty and 239 (72%) contained food. Sablefish ranged in

size from 40 cm to 80 cm FL with a mean and of 60.3 and + 9.1 cm,

respectively. The average depth of the 40 haul locations sampled

was 201 + 63 m with a range from 101 to 432 m. Table 5-l lists

the percent frequency of occurrence, the percent by number, and

the percent by weight of the prey items found in sablefish

stomachs. Approximately 74% of the weight of the total stomach

contents was fish, with invertebrates making up only one-quarter

of the total stomach contents weight. Walleye pollock was the

predominate fish-prey consumed by sablefish. Pollock comprised

about 24% by weight of the total sablefish stomach contents.
90

Table 5-l. --Prey items (expressed in percent frequency of

occurrence, numerical percentage, and percent total

weight) of sablefish (Anonloooma fimbria) collected in

the Gulf of Alaska in 1990.

Prey Name

% Freq. % Total

occur. % Number weight
91

Table 5-l. --Continued.

Prey Name

% Freq. % Total

occur. % Number weight

Hemitripterus bolini (bigmouth sculpin) 0.80 0.02 0.05

Cyclopteridae (snailfish) 1.30 0.05 0.03

Stichaeidae (prickleback) 0.80 0.02 0.02

Ammodytes hexapterus (Pacific sand lance) 0.40 0.01 0.08

Pleuronectidae (flatfish) 0.40 0.01 0.05

Fishery discards 12.10 0.38 29.08

Total prey count 10,493

Total prey weight 4,354 g

Number of stomachs with food 239

Number of empty stomachs 92
9 3

Figure 5-l. --Variations in the main food items of

sablefish, by predator size, In the

Gulf of Alaska in 1990.

N = sample size.
94

40 cm FL were available. Figure 5-1 illustrates that the diet

did not vary much by size of sablefish for the fish examined in

this study. Cephalopods (14%) were primarily consumed by the

60-69 cm FL size group and more euphausiids and decapods were

consumed by the two smaller-sized groups (40-49 cm and 50-59 cm

FL). The miscellaneous fish category comprised higher

percentages in both the smallest size group (40-49 cm FL) and

largest size group (<70 cm FL). Sablefish 50-59 cm FL consumed a

high percentage (56%) of fishery discards, though all other sizegroups also consumed a fairly large amount (about 20% by weight)

of fishery discard. Walleye pollock were consumed more by the

two larger size groups (60-69 cm and >70 cm FL ) than the two

smaller size groups.

Sizes of the Commercially Important Prey Consumed

The prey length data of the sablefish were divided by

predator size into less than 55 cm and greater than or equal to

55 cm size groups for analysis (Fig. 5-2). Figure 5-2 (left)

shows that walleye pollock consumed by both size groups were

mainly the pre-recruits (<300 mm SL), though the larger size

group also consumed a few pollock greater than or equal to

400 mm. The average (mean + SD) SL of pollock consumed by

sablefish was 234.9 1 124.9 mm with a range from 56 to 445 mm.

The relationship between the size of the sablefish and the size

of the pollock that they consumed is shown in Figure 5-3.

The size of the eulachon consumed by sablefish is shown in

Figure 5-2 (right). It shows that sablefish less than 55 cm FL

only consumed smaller eulachon (<l00 mm SL), whereas larger
95

Figure 5-2.--Size frequency distributions of walleye

pollock and eulachon consumed by two

size groups of sablefish in the Gulf

of Alaska in 1990.
Figure 5-3. --Relationship between the sablefish fork length (cm)

and the walleye pollock standard length (mm) in the

Gulf of Alaska in 1990.
97

sablefish (>55 cm FL) also consumed some larger eulachon (about

150 mm SL). The average length of the eulachon consumed was

134.1 +. 41.0 mm SL with a range from 68 to 165 mm SL.

Sablefish also consumed three capelin (82, 96, 106 mm SL),

two Pacific herring (154 and 170 mm SL), one Pacific sand lance

(97 mm SL), and three Tanner crabs (5, 21, 54 mm CW).

Geographic Distributions of the Prey Consumed

The distributions of the commercially important prey

consumed by sablefish are shown in Figures 5-4 to 5-7.

Figure 5-4 shows that walleye pollock were consumed by sablefish

at many of the sampled stations, and that the percent by weight

in many stations was high (>50%). Figure 5-5 illustrates that

capelin were consumed by sablefish in quite a few stations even

though they usually did not comprise a high proportion of the

stomach contents at any station. The distribution of the

consumption of Tanner crabs by sablefish is shown in Figure 5-6.

Even though Tanner crabs were consumed at only two stations,

their high percentages of occurrence in stomach contents

indicates that they were heavily targeted by the sablefish in

those areas. The consumption of fishery discard by sablefish was

primarily distributed around Kodiak Island (Fig. 5-7).

DISCUSSION

Shubnikov (1963) found that fish (including pollock, capelin

and herring) made up the major part of the Bering Sea sablefish

diet in spring and autumn; during the summer months the sablefish

switch to a diet including shrimp, jellyfish, and some benthic
Figure 5-4. -,-Geographic distribution of walleye pollock consumed

by sablefish in the Gulf of Alaska in 1990.


165 OOW 158 OOW I51 oow 144 oow I37 oow

Figure 5-T.--Geographic distribution of fishery discard consumed

by sablefish in the Gulf of Alaska in 1990.
102

invertebrates. Mito (1974) found that pollock was the

predominate food of sablefish in the eastern Bering Sea (in some

areas pollock made up as much as 97% of the diet of sablefish).

Brodeur and Livingston (1988) found gadid fish (mainly pollock)

accounted for about 50% of the total stomach content weight.

They also reported that 8.5% of the sablefish diet was fishery

discards.

Comparing Mito's (1974) study, Brodeur and Livingston's

(1988) study, and this study, I found that sablefish consumed

more pollock in the eastern Bering Sea than in the Gulf of

Alaska. On the other hand, fishery discards comprised a higher

percentage (29%) of the food of sablefish in the Gulf of Alaska

area than in the eastern Bering Sea (8.5%).
103

CITATIONS

Brodeur, R. D., and P. A. Livingston. 1988. Food habits and diet

overlap of various eastern Bering Sea fishes. U.S. Dep.

Commer., NOAA Tech. Memo. NMFS F/NWC-127, 76 p.

Fujioka, J. T. 1990. Sablefish, p. 125-139. In Stock assessment

and fishery evaluation report for the 1991 Gulf of Alaska

groundfish fishery. Gulf of Alaska Groundfish Plan Team,

North Pacific Fishery Management Council, P.O. Box 103136,

Anchorage, AK 99510.

Grinols, R. B., and C. D. Gill. 1968. Feeding behavior of three

oceanic fishes (Oncorhynchus kisutch, Trachurus symmetricus,

and Anoplopoma fimbria)from the northeastern Pacific. J.

Fish. Res. Board. Can. 25(4): 825-827.

Mito, K. I. 1974. Food relationships among benthic fish

populations in the Bering Sea. M.S. Thesis, Hokkaido Univ.,

Hokkaido, Japan, 135 p.

North Pacific Fishery Management Council. 1991. Introduction,

P- l-27. In Stock assessment and fishery evaluation report

for the 1992 Gulf of Alaska Groundfish Fishery, prepared by

Gulf of Alaska Groundfish Plan Team, North Pacific Fishery

Management Council. P. 0. Box 103136, Anchorage, AK 99510.
104

Shubnikov, D. A. 1963. Nekotorye dannye po biologii ugol'noi ryby

Beringova morya (Data on the biology of sablefish of the

Bering sea). Tr. Vses. Nauchno Issled. Inst. Morsk. Rybn.

Khoz. Okeanogr. 48 (Izv. Tikhookean. Nauchno-Issled. Inst.

Rybn. Khoz. Okeanogr. 50):271-295. In Russ. (Transl. by Isr.

Program Sci. Transl., 1968, p. 287-296, &I P. A. Moiseev

(ea.), Soviet fisheries investigations in the northeast

Pacific, Part 1, available U.S. Dep. Commer., Natl. Tech. Inf.

Serv., Springfield, VA, as TT 67-51203).
105

PACIFIC OCEAN PERCH

The estimated 1990 exploitable biomass of Pacific ocean

perch (Sebastes alutus) in the Gulf of Alaska area was 129,734 t

(Heifetz and Clausen 1991). It ranked eighth (first of all of

the rockfishes) in total biomass of the Gulf of Alaska groundfish

complex. Pacific ocean perch is a commercially important fish in

the Gulf of Alaska area. Trawl fisheries by the U.S.S.R. and

Japan for this species began in the early 1960s and overexploited

the stock (Heifetz and Clausen 1991). Since the mid-1960s the

catches have fluctuated. In 1990 the total catch' was about

21,114 t (Heifetz and Clausen 1991). The objective of this study

was to understand the food habits of Pacific ocean perch and

their relation to other marine fishes in the Gulf of Alaska area.

RESULTS

A total of 143 Pacific ocean perch stomachs were analyzed,

of which 41 were empty and 102 (71%) contained food. Pacific

ocean perch sizes ranged from 14 to 48 cm FL with a mean and SD

of 32.3 + 7.2 cm. Figure 6-l shows the haul locations for the

collected stomach samples of the Pacific ocean perch in the Gulf

of Alaska in 1990. The average depth of the 16 haul locations

was 183 + 71 m with a range from 101 to 346 m.

l/ The total catch included the 18 slope rockfish species,

in which Pacific ocean perch was the dominant species.

107

Table 6-1 lists the percent frequency of occurrence, the

percent by number, and the percent total weight of the prey items

of Pacific ocean perch. Fish comprised only 2% by weight of the

diet of Pacific ocean perch in which arrowtooth flounder

(Atheresthes stomias) was the only major groundfish species

consumed. The SLs of the two measurable arrowtooth flounder were

12.1 mm and 22.0 mm. Other fish consumed by Pacific ocean perch

included myctophids, snailfish, and stichaeids. Euphausiids

(mainly Thysanoessa inermis) were the most important prey of

Pacific ocean perch; they comprised about 87% by weight of the

total stomach contents. Calanoid copepods, amphipods, and arrow

worms were found frequently in Pacific ocean perch stomachs.

However, they comprised very low percentages by weight of the

total diet of Pacific ocean perch. Pacific ocean perch stomachs

also contained 0.5% by weight of Tanner crabs (Chionoectes.

bairdi). However, no Tanner crabs were measurable. Shrimp,

including pasiphaeids, pandalids, and crangonids comprised 2.5%

by weight of the diet of Pacific ocean perch.

Figure 6-2 illustrates that there were no large diet

variations between the different size groups of the Pacific ocean

perch except the smallest size group ((25 cm FL) consumed more

euphausiids.

DISCUSSION

Because of the wide range distributions (from southern

California to the Bering Sea) of the Pacific ocean perch, the

diets of this species have been documented in many areas
108

Table 6-l. --Prey items (expressed in percent frequency of

occurrence, numerical percentage, and percent total

weight) of Pacific ocean perch (Sebastes alutus)

collected in the Gulf of Alaska in 1990.
109

Figure 6-2. --Variations in the main food items of

Pacific ocean perch, by predator size,

in the Gulf of Alaska in 1990.

N = sample size.
110

(Lyubimova 1963 and Somerton et al. 1978 in the Gulf of Alaska;

Skalkin 1964 and Mito 1974 in the Bering Sea; Brodeur and Pearcy

1984 along the West Coast). All of these previous studies showed

similar results to this study. That is, Pacific ocean perch is

mainly planktivorous; the smaller fish eat a higher percent of

canlanoid copepods and the larger fish eat a higher percent

(usually more than 85%) of euphausiids and some shrimp. The

structure of the gill rakers also supports the belief that

Pacific ocean perch is planktivorous. The outer rakers (about 35

to 39 counts based on my study) on the first gill arch are

slender and long, the inner rakers on the first gill arch

interlock with the outer rakers on the second gill arch, and so

on for the rest of the gill arches. In this way the gill arches

form a screen to filter planktonic organisms.
111

CITATIONS

Brodeur, R. D., and W. G. Pearcy. 1984. Food habits and dietary

overlap of some shelf rockfishes (Genus Sebastes) from the

northeastern Pacific Ocean. Fish. Bull., U.S. 82:269-293.

Heifetz, J., and D. M. Clausen. 1991. Slope rockfish. In Stock

assessment and fishery evaluation report for the 1992 Gulf

of Alaska groundfish fishery. North Pacific Fishery

Management Council, P.O. Box 103136, Anchorage, AK 99510.

Lyubimova, T. G. 1963. Osnovnye cherty biologii raspredeleniya

Tikhookeanskogo morskogo okunya (Sebastodes alutus Gilbert)

v zalive Alyaska (Basic aspects of the biology and

distribution of Pacific rockfish (Sebastodes alutus Gilbert)

in the Gulf of Alaska). Tr. Vses. Nauchno-Issled. Inst.

Morsk. Rybn. Khoz. Okeanogr. 48 (Izv. Tikhookean. NauchnoIssled. Inst. Rybn. Khoz. Okeanogr. 50):293-303. In Russ.

(Transl. by Isr. Program Sci. Transl., 1968, p. 308-318, In

P. A. Moiseev (ed.), Soviet fisheries investigations in

the northeastern Pacific, Part 1, available U.S. Dep.

Commer., Natl. Tech. Inf. Serv., Springfield, VA, as TT

67-51203).

Mito, K. I. 1974. Food relationships among benthic fish

populations in the Bering Sea. M.S. Thesis, Hokkaido Univ.,

Hokkaido, Japan, 135 p.
112

Skalkin, V. A. 1964. Pitanie morskikh okunei v Beringovum more

(Diet of rockfish in the Bering Sea). Tr. Vses. NauchnoIssled. Inst. Morsk. Rybn. Khoz. Okeanogr. 49 (Izv.

Tikhookean. Nauchno-Issled. Inst. Rybn. Khoz. Okeanogr.

51): 151-166. In Russ. (Transl. by Isr. Program. Sci.

Transl., 1968, p. 159-174. In P. A. Moiseev (ed.), Soviet

fisheries investigations in the northeast Pacific, Part 2,

available U.S. Dep. Commer., Natl. Tech. Inf. Serv.,

Springfield, VA, as TT 67-51204).

Somerton, D., F. Funk, K. Mesmer, L. J. Bledsoe, and K.

Thornburgh. 1978. A comparative study of the diets of

Pacific ocean perch (Sebastes alutus) and walleye pollock

(Theragra chalcoaranuna) in the Gulf of Alaska. NORFISH

NPB8, 25 p. Washington Sea Grant Communications Program,

University of Washington, Division of Marine Resources, 3716

Brooklyn Ave. N.E., Seattle, WA 98195.
113

ROUGHEYE ROCKFISH

The rougheye rockfish (Sebastes aleutianus) had an

exploitable biomass of 36,794 t in the Gulf of Alaska in 1990

(Heifetz and Clausen 1991). It ranked third in abundance of the

rockfish complex in the Gulf of Alaska. The catch of the

rougheye rockfish was not available since it was combined with 17

other rockfish species of the slope rockfish assemblage. Slope

rockfish are defined as those Sebastes species that inhabit the

Gulf of Alaska area at depths greater than 150-200 m (Heifetz and

Clausen 1991).

RESULTS

A total of 141 rougheye rockfish stomachs were analyzed, of

which 57 were empty and 84 stomachs (60%) contained food. The

average size of the rougheye rockfish was 36.4 cm FL with a SD of

5.9 cm. Fish size ranged from 21 to 56 cm FL. The average depth

of the 17 haul locations was 234 + 82 m with a range from 148 to

368 m.

Table 7-l shows the percent frequency of occurrence, percent

by number, and percent by weight of the prey items of rougheye

rockfish. It shows that shrimp, including Pandalus borealis,

Pandalus montaaui tridens, hippolytids, and crangonid, are the

main food (more than 50% of the total stomach contents weight) of

rougheye rockfish. The area and amount (percent by weight) of

pandalid shrimp consumed by rougheye rockfish is shown in Figure

7-l. Squid also comprised a high proportion (21%) of the stomach

contents weight. This was mostly due to one large rougheye
114

Table 7-l. --Prey items (expressed in percent frequency of

occurrence, numerical percentage, and percent total

weight) of rougheye rockfish (Sebastes aleutianus)

collected in the Gulf of Alaska in 1990.

Prey Name

% Freq. % Total

occur. % Number weight

116

rockfish (56 cm) that ate one squid weighing 43.1 g; however,

squid were not the common food of rougheye rockfish (only 2.4% of

the frequency of occurrence). Other food items included

amphipods, mysids, euphausiids, cumaceans, isopods, and

polychaetes. Prey fish consumed by rougheye rockfish included

non-gadoid fish, snailfish, flatfish, and some unidentifiable

fish. They comprised about 7% of the total stomach contents

weight. The commercially important Tanner crab (Chionoecetes

bairdi) was also consumed by rougheye rockfish, it comprised

about 2% of the total stomach contents weight. One measurable

Tanner crab, 26.5 mm CW, was consumed by one 39 cm rougheye

rockfish.

Figure 7-2 shows the variations of the main food items of

different size groups of rougheye rockfish. It shows that all

size groups consumed a large amount of shrimp (>50% by weight).

The smallest size group (<30 cm FL) consumed more amphipods

whereas the largest size group (40-49 cm FL) consumed more fish

than the other size groups. The medium-sized (30-39 cm FL)

rougheye rockfish consumed relatively more crab compared to other

groups. Diet of rougheye rockfish larger than 50 cm FL (only

one) was not included in Figure 7-2 because of the small sample

size.
Figure 7-2.--Variations in the main food items of

rougheye rockfish, by predator size,

in the Gulf of Alaska in 1990.

N = sample size.
118

DISCUSSION

Feder (1980) found that shrimp were the most important food

of rougheye rockfish (occurred in 75% of the stomachs) in the

southeastern Gulf of Alaska. He also reported that fish, squid,

and mysids were found occasionally in rougheye stomachs. Feder's

(1980) results are similar to this study.
119

CITATIONS

Feder, H. M. 1980. Distribution, abundance, community structure,

and trophic relationships of the benthos of the northeastern

Gulf of Alaska from Yakutat Bay to Cross Sound. In

Environmental Assessment of the Alaskan Continental Shelf,

Annual Reports 1:597-648. U. S. Dep. Commer., Natl. Oceanic

Atmos. Admin., Off. Mar. Pollut. Assess., Juneau, Alaska.

Heifetz, J., and D. M. Clausen. 1991. Slope rockfish. In Stock

assessment and fishery evaluation report for the 1992 Gulf

of Alaska groundfish fishery. North Pacific Fishery

Management Council, P.O. Box 103136, Anchorage, AK 99510.
120

NORTHERN ROCKFISH

Northern rockfish (Sebastes polyspinis) had an exploitable

biomass of 74,626 t in the Gulf of Alaska in 1990 (Heifetz and

Clausen 1991). It was the second largest group of rockfish (next

to Pacific ocean perch) in the Gulf of Alaska in terms of

biomass. Along with Pacific ocean perch (S. alutus), rougheye

rockfish (S. aleutianus), shortraker rockfish (S. borealis), and

the northern rockfish were categorized as members of the Pacific

ocean perch (POP) complex between 1979 and 1987. Since 1988,

however, northern rockfish has been classified and managed as one

of the 18 Sebastes species within the slope rockfish assemblage.

Slope rockfish are defined as those species of Sebastes that

inhabit depths greater than 150-200 m (Heifetz and Clausen 1991).

RESULTS

A total of 57 northern rockfish stomachs were analyzed, of

which 18 stomachs were empty and 39 stomachs (68%) contained

food. The average size of the northern rockfish was 35.1 cm FL

with a SD of 2.3 cm. Fish size ranged from 30 to 41 cm FL. The

average depth of the seven haul locations was 116 + 9 m with a

range between 102 and 126 m.

Table 8-l shows the percent frequency of occurrence, the

percent by number, and the percent by weight of the prey items

consumed by northern rockfish. It shows that euphausiids (mainly

Thysanoessa inermis) were the most important (95% by weight) food

of northern rockfish. Other food included calanoid copepods,

cephalopods, gastropods, amphipods, Chionoecetes sp., pagurid
121

Table 8-l. --Prey items (expressed in percent frequency of

occurrence, numerical percentage, and percent total

weight) of northern rockfish (Sebastes polyspinis)

collected in the Gulf of Alaska in 1990.

Prey Name

% Freq. % Total

occur. % Number weight

Gastropods (snail) 2.60 0.01 0.01

Cephalopoda (unidentified) 7.70 0.03 0.30

Teuthoidea oegopsida (squid) 5.10 0.01 0.01

Calanoida (unidentified) 10.30 0.06 0.02

Neocalanus cristatus (copepod) 28.20 3.82 2.43

Gammaridea (unidentified) 12.80 0.05 0.08

Themisto sp. (amphipod) 35.90 0.21 0.05

Euphausiacea (unidentified) 89.70 94.65 95.41

Thvsanoessa inermis (euphausiid) 10.30 0.50 0.82

Thvsanoessa soinifera (euphausiid) 2.60 0.01 0.05

Caridea (unidentified shrimp) 2.60 0.01 0.01

Paguridae (hermit crab) 5.10 0.01 0.01

Chionoecetes sp. (snow and Tanner crab) 5.10 0.01 0.01

Sasitta sp. (arrow worm) 35.90 0.60 0.41

Non-gadoid fish remains 2.60 0.01 0.32

Atheresthes stomias (arrowtooth flounder) 7.70 0.03 0.07

Total prey count 14,772

Tota- prey weight 142 g

Number of stomachs with food 39

Number of empty stomachs 18
122

crabs, and arrow worms. Prey fish found in northern rockfish

stomachs were arrowtooth flounder (Atheresthes stomias) and some

non-gadoid fish remains. The four arrowtooth flounder consumed

had a mean and SD of 12.8 + 2.0 mm SL with a range from 11.5 to

15.7 mm. The locations where the arrowtooth flounder were

consumed are shown in Figure 8-l. Because of the small sample

size and the small size range (30 to 41 cm FL) of the northern

rockfish collected, variation of the diet of different size

groups is not available.

'DISCUSSION

Skalkin (1964) found that northern rockfish in the Bering

Sea area consumed large amounts of euphausiids, as well as

cephalopods, copepods, arrow worms, and larvaceans. Mito (1974)

also found a similar diet: euphausiids, hermit crabs, northern

shrimp Pandalus borealis, and squid in northern rockfish stomachs

in the Bering Sea. Skalkin (1964), Mito (1974), and this study

indicate that northern rockfish is mainly planktivorous. This

fact is supported by the similarities noted in the gill arch

structures of northern rockfish to the Pacific ocean perch gill

arch structures described earlier in the food habits section for

that species.
Figure 8-l. --Geographic distribution of the arrowtooth flounder

consumed by northern rockfish in the Gulf of Alaska

in 1990.
124

CITATIONS

Heifetz, J., and D. M. Clausen. 1991. Slope rockfish. In Stock

assessment and fishery evaluation report for the 1992 Gulf

of Alaska groundfish fishery. North Pacific Fishery

Management Council, P.O. Box 103136, Anchorage, AK 99510.

Mito, K. I. 1974. Food relationships among benthic fish

populations in the Bering Sea. M.S. Thesis, Hokkaido Univ.,

Hokkaido, Japan, 135 p.

Skalkin, V. A. 1964. Pitanie morskikh okunei v Beringovum more

(Diet of rockfish in the Bering Sea). Tr. Vses. NauchnoIssled. Inst. Morsk. Rybn. Khoz. Okeanogr. 49 (Izv.

Tikhookean. Nauchno-Issled. Inst. Rybn. Khoz. Okeanogr.

51): 151-166. In Russ. (Transl. by Isr. Program. Sci.

Transl., 1968, p. 159-174. In P. A. Moiseev (ed.), Soviet

fisheries investigations in the northeast Pacific, Part 2,

available U.S. Dep. Commer., Natl. Tech. Inf. Serv.,

Springfield, VA, as TT 67-51204).
125

DUSKY ROCKFISH

Dusky rockfish (Sebastes ciliatus) had an exploitable

biomass of 24,141 t in the Gulf of Alaska in 1990 (Clausen and

Heifetz 1991). It ranked fifth in abundance among the rockfish

complex in the Gulf of Alaska in 1990. Along with black rockfish

(S. melanops), widow rockfish (S. entomelas), blue rockfish (S.

mystinus), and yellowtail rockfish (S. flavidus), dusky rockfish

is one of the five Sebastes spp. in the pelagic shelf rockfish

assemblage. This assemblage is defined as those species of

Sebastes that inhabit waters of the continental shelf of the Gulf

of Alaska and exhibit a midwater schooling behavior (Clausen and

Heifetz 1991). The catch of this assemblage in 1990 (1,647 t)

was much less than the quota (8,200 t) assigned by the North

Pacific Fishery Management Council, 'indicating this assemblage

was underutilized by the commercial fishery.

RESULTS

A total of 41 dusky rockfish stomachs were analyzed, of

which 19 were empty and 22 (54%) contained food. The average

size of the dusky rockfish was 39.7 cm FL with a SD of 7.7 cm.

Fish size ranged from 16 to 47 cm FL. The average depth of the

five haul locations (Fig. 9-1) was 99 + 27 m with a range between

51 and 117 m.

Table 9-l lists the percent frequency of occurrence, the

percent by number, and the percent by weight of the prey items

found in dusky rockfish stomachs. It shows that euphausiids

(mainly Thysanoessa inermis) were the most important food (67% by

127

Table 9-l. --Prey items (expressed in percent frequency of

occurrence, numerical percentage, and percent total

weight) of dusky' rockfish (Sebastes ciliatus)

collected in the Gulf of Alaska in 1990.

Prey Name

% Freq. % Total

occur. % Number weight

Gastropods (snail) 4.50 0.05 0.08

Cephalopoda (unidentified) 22.70 0.83 6.07

Teuthoidea oegopsida (squid) 4.50 0.05 0.07

Calanoida (unidentified) 18.20 0.68 0.11

Neocalanus cristatus (copepod) 9.10 2.43 1.82

Gammaridea (unidentified) 4.50 0.05 0.13

Ampeliscidae (amphipod) 9.10 0.10 0.20

Euphausiacea (unidentified) 59.10 26.19 55.89

Thvsanoessa inermis (euphausiid) 31.80 4.47 10.92

Thvsanoessa sninifera (euphausiid) 4.50 0.24 2.04

Caridea (unidentified shrimp) 9.10 1.55 0.71

Hippolytidae (shrimp) 4.50 0.05 0.01

Pandalidae (shrimp) 9.10 0.15 3.46

Paguridae (hermit crab) 18.20 8.41 3.10

Cancer sp. (crab) 4.50 0.05 0.02

Sasitta sp. (arrow worm) 36.40 8.21 3.34

Larvacea (tunicate) 22.70 45.29 11.64

Total prey count 2,058

Total prey weight 24 g

Number of stomachs with food 22

Number of empty stomachs 19
128

weight) of dusky rockfish. Larvaceans were the second most

important prey of dusky rockfish; they comprised 12% of the total

stomach contents weight. Cephalopods, calanoid copepods, arrow

worms, and hermit crabs were also found-quite frequently;

however, they all comprised less than 10% by weight of the

stomach contents. Dusky rockfish also consumed small amounts of

gammarid amphipods, pandalids, hippolytids, and snails. Cancer

sp. (one megalops larvae) was the only commercially important

prey consumed by dusky rockfish. It comprised less than 1% of

the total stomach contents weight.

Diet variations between different size groups were not

available because of the small sample size (n=41) in this study.

DISCUSSION

Simenstad et al. (1977) reported that mysids, amphipods,

copepods, and sand lance were the main food of dusky rockfish

near Amchitka Island in the Aleutian Islands. Blackburn et al.

(1981) found that crab megalops, chaetognaths, and calanoid

copepods were important as food items for dusky rockfish in the

Cook Inlet estuary. Rosenthal et al. (1988) found that dusky

rockfish consumed large amounts (25.4% by volume of the total

stomach content) of brachyuran crab larvae in the southeastern

Gulf of Alaska. Calanoid copepods, amphipods, jellyfish, and

pteropods were also consumed. Rosenthal et al. (1988) also

reported that some gadid larvae and Pacific sand lance were found

in dusky rockfish stomachs.
129

Based on this study and similar studies, dusky rockfish,

like Pacific ocean perch and northern rockfish, is mainly a

planktivore. The compositions of the prey might vary depending

on different study areas and different sampling seasons.
130

CITATIONS

Blackburn, J.E., K. Anderson, C.I. Hamilton, and S. J. Starr.

1981. Pelagic and demersal fish assessment in the lower Cook

Inlet estuary system. In Environmental Assessment of the

Alaskan Continental shelf, Final Reports, Biological Studies,

l2: 259-602. U. S. Dep. Commer., Natl. Oceanic Atmos. Admin.,

Off. Mar. Pollut. Assess., Juneau, Alaska.

Clausen, D. M., and J.

Stock assessment

Heifetz. 1991. Pelagic shelf rockfish. In

and fishery evaluation report for the 1992

Gulf of Alaska groundfish fishery. North Pacific Fishery

Management Council, P.0. Box 103136, Anchorage, AK 99510.

Rosenthal, R. J., V. M. O'Connell, and M. C. Murphy. 1988.

Feeding ecology of ten species of rockfishes (Scorpaenidae)

from the Gulf of Alaska. Calif. Fish Game 74(l): 16-37.

Simenstad, C. A., J. S. Isakson, and R. E. Nakatani. 1977. Marine

fish communities of Amchitka Island, Alaska. In M. L. Merritt

and R. G. Fuller (ed.), The Environment of Amchitka Island,

Alaska. U.S. Energy Research and Development Administration.

TID 267-12:451-492.
131

SHORTSPINE THORNYHEAD

Shortspine thornyhead (Sebastolobus alascanus) had an

exploitable biomass of 25,697 t in the Gulf of Alaska in 1990

(Dawson 1991). It ranked seventh in abundance of the total

rockfish complex in the Gulf of Alaska. Since 1980, the

shortspine thornyhead resource has been managed as a unit in the

Gulf of Alaska. In the past, thornyheads were not the primary

target of the rockfish fleet. Today, thornyheads are one of the

most commercially valuable rockfish species. In 1990, the catch

of this species in the Gulf of Alaska area was 1,646 t (Dawson

1991).

RESULTS

A total of 120 shortspine thornyhead stomachs were analyzed,

of which 39 were empty and 81 (68%) contained food. The average

size of the shortspine thornyhead was 29.7 cm FL with a SD of

6.6 cm. Fish size ranged from 13 to 48 cm FL. The average depth

of the 11 haul locations was 310 + 77 m with a range from 212 to

439 m.

Table 10-l lists the percent frequency of occurrence, the

percent by number, and the percent by weight of the prey items

found in the stomachs of shortspine thornyhead. It shows that

shrimp (mainly pandalids) were the most important food of the

thornyhead (67% by weight). The geographic distribution and the

amount of the pandalids (including Pandalus borealis, p. jordani,

p. montagui tridens) is shown in Figure 10-l. Tanner crabs

(Chionoecetes bairdi) comprised about 1% (by weight) of the food
132

Table 10-l. --Prey items (expressed in percent frequency of

occurrence, numerical percentage, and percent total

weight) of shortspine thornyhead (Sebastolobus

alascanus) collected in the Gulf of Alaska in 1990.

Prey Name

% Freq. % Total

occur. % Number weight

Polychaeta (worm)

Gastropoda (snail)

Teuthoidea (squid)

Calanoida (copepod)

Mysidacea (unidentified mysid)

Holmesiella anomala (mysid)

Metervthrops robusta (mysid)

Pseudomma truncatum (mysid)

Gammaridea (unidentified amphipod)

Ampeliscidae (amphipod)

Lysianassidae (amphipod)

Caridea (unidentified shrimp)

Pasiphaeidae (shrimp)

Hippolytidae (unidentified shrimp)

Eualus barbata (shrimp)

Eualus biunquis (shrimp)

Eualus avinus (shrimp)

Heptacarpus sp. (shrimp)

Pandalidae (unidentified shrimp)

Pandalus borealis (northern shrimp)

Pandalus jordani (shrimp)

Pandalus montaqui tridens (shrimp)

Crangonidae (unidentified shrimp)

Cranaon communis (shrimp)

Arais ovifer (shrimp)

Paguridae (hermit crab)

Lithodidae (crab)

Oresonia sracilis (decorator crab)

Chionoecetes bairdi (Tanner crab)

Cherilia lonqioes (decorator crab)

Pinnotheridae (pea crab)

Teleostei (unidentified fish)

Non-gadoid fish remains

Mallotus villosus (capelin)

Theraqra chalcoqramma (walleye pollock)

Zoarcidae (eelpout)

Icelinus borealis (northern sculpin)

Total prey count

Total prey weight

Number of stomachs with food

Number of empty stomachs

6.20

1.20

3.70

2.50

7.40

8.60

1.20

3.70

13.60

3.70

1.20

4.90

1.20

13.60

2.50

3.70

4.90

2.50

33.30

4.90

3.70

2.50

7.40

9.90

1.20

2.50

1.20

2.50

3.70

3.70

1.20

2.50

7.40

1.20

1.20

1.20

2.50

2.37

0.95

1.42

0.95

3.32

8.06

0.47

1.42

9.00

3.32

0.47

1.90

0.47

8.06

1.42

1.42

2.84

0.95

23.69

1.90

2.37

0.95

3.32

5.21

0.47

0.95

0.47

0.95

1.90

1.42

0.47

0.95

2.84

0.47

0.47

0.47

0.95

211

147 g

81

39

1.18

1.54

0.72

0.01

0.17

0.41

0.03

0.07

0.84

0.11

0.25

0.38

1.04

3.46

0.52

0.64

0.59

0.38

27.86

7.38

18.39

0.81

1.22

4.12

0.11

2.04

1.12

2.19

1.16

3.96

0.10

1.30

13.75

0.72

0.85

0.04

0.51
Figure l0-l.--Geographic distribution of pandalid shrimp consumed

by shortspine thornyhead in the Gulf of Alaska in

1990.
134

of thornyheads. The sizes of the four Tanner crabs consumed were

between 13 and 18 mm carapace width with a mean and standard

deviation of 16.4 + 2.5 mm. Fish comprised about 17% (by weight)

of the food of the thornyhead. Although the thornyhead stomach

did not contain a large amount of prey fish, they had the highest

proportion of prey fish as food between all the rockfish species.

Among the prey fish consumed, one 49 mm SL walleye pollock

(Theragra chalcogramma) was found in a 37 cm thornyhead stomach.

One capelin (Mallotus villosus) was also found in a thornyhead

stomach; however, the fish size was not measurable. Shortspine

thornyhead also consumed a zoarcid, sculpin, and some

unidentifiable fish.

DISCUSSION

Feder (1980) found that shrimp (occurred in 90% stomachs)

were the most important food for shortspine thornyhead in the

southeastern Gulf of Alaska area, followed by fish (15% frequency

of occurrence). His results basically agreed with this study.

However, in the Bering Sea area, Mito (1974) found that fish,

euphausiids, Chionoecetes opilio, and macrura were important

food, whereas shrimp were not. This difference was probably

because of the differences of the prey abundances and prey

availabilities in the different areas.

Compared to most other rockfish, shortspine thornyhead has a

more elongated body and a relatively large terminal mouth. It

has 18 to 23 gill rakers on the first gill arch (fewer than most

of the other rockfishes), and the gill rakers are stout and not
135

very long. All of these contributed to their feeding mainly on

epibenthic shrimp and fish and not much on the planktons.
Dawson, P. 1991. Shortspine thornyhead. In Stock assessment

and fishery evaluation report for the 1992 Gulf of Alaska

groundfish fishery. North Pacific Fishery Management

Council, P.O. Box 103136, Anchorage, AK 99510.

Feder, H. M. 1980. Distribution, abundance, community structure,

and trophic relationships of the benthos of the northeastern

Gulf of Alaska from Yakutat Bay to Cross Sound. In

Environmental Assessment of the Alaskan Continental Shelf,

Annual Reports 1:597-648. U. S. Dep. Commer., Natl. Oceanic

Atmos. Admin., Off. Mar. Pollut. Assess., Juneau, Alaska.

Mito, 'K. I. 1974. Food relationships among benthic fish

populations in the Bering Sea. M.S. Thesis, Hokkaido Univ.,

Hokkaido, Japan, 135 p.
137

SHORTRAKER ROCKFISH

Shortraker rockfish (Sebastes borealis) had an exploitable

biomass of 9,449 t the Gulf of Alaska in 1990 (Heifetz and

Clausen 1991). It ranked 10th in abundance of the rockfish

complex in-the Gulf of Alaska in 1990. This species has been

classified and managed as one of the 18 slope assemblage

rockfishes since 1988 (Heifetz and Clausen 1991). Because of

their special morphological characteristics (large mouth, short

gill rakers, and low gill raker number), they are a potential

predator of fish, cephalopods, and other commercially important

species. Therefore, it is important to study the food habits of

the shortraker rockfish.

RESULTS

Ten stomachs of shortraker rockfish collected from one haul

(361 m deep) in the Gulf of Alaska were analyzed, of which seven

were empty, and three contained food. The average size of the

shortraker rockfish was 57.3 cm FL with a SD of 5.4 cm. Fish

length ranged between 50 and 68 cm FL. The diet of the

shortraker rockfish was comprised of 82% (by weight) of squid and

18% myctophids. Since the sample size and the size range of the

fish were too small, the data was insufficient to describe the

food habits of shortraker rockfish. More samples should be

collected for the future study of this species.
138

CITATIONS

Heifetz, J., and D. M. Clausen. 1991. Slope rockfish. In Stock

assessment and fishery evaluation report for the 1992 Gulf

of Alaska groundfish fishery. North Pacific Fishery

Management Council, P.O. Box 103136, Anchorage, AK 99510.
139

SUMMARY

A total of 4,429 stomachs from 11 species were analyzed to

describe the food habits of the major groundfish species in the

Gulf of Alaska. The analysis emphasized predation on

commercially important fish, crab, and shrimp. The predator

sizes and the number of the stomachs collected for each species

are summarized in table 1. Although juveniles of some species

(walleye pollock, Pacific cod, Pacific halibut, and arrow-tooth

flounder) were sampled, the main focus was on the food habits of

the adult fish.

A Table 2 lists the percent by weight of the commercially

important fish, crab, shrimp, and other major prey or prey groups

consumed by groundfish. It shows that arrowtooth flounder,

Pacific halibut, sablefish, Pacific cod, and pollock are the main

predators that consume fish. The main predators that feed on

Tanner crabs are Pacific halibut and Pacific cod. Pollock,

shortspine thornyhead, and rougheye rockfish are the main

consumers of pandalid shrimp. Pacific ocean perch, northern

rockfish, and dusky rockfish did not have too much impact on the

commercially important species in the Gulf of Alaska since they

feed mainly on zooplankton (mainly euphausiids). Although there

was not enough data (only three stomachs with food) to describe

the diet of the shortraker rockfish, based on their morphological

characteristics (large mouth, short gillrakers, and less

gillraker numbers), they probably also consume some fish and crab

in addition to the cephalopods found in this study.
140

Table 1 .--Number of stomachs and fish size analyzed in the Gulf

of Alaska in 1990. F, stomachs with food; E, empty

stomachs; T, total; SD, standard deviation.
141

Table 2. --Percent by weight of the important prey or prey group

consumed by the groundfish in the Gulf of Alaska in

1990. "-" means less than 1%. PLK, pollock; HER,

herring; CAP, capelin; ATF, arrowtooth flounder; FHS,

flathead sole; GT, Greenland turbot; RS, rock sole;

REX, rex sole; YFS, yellowfin sole; PH, Pacific

halibut; DOV, Dover sole; SAN, Pacific sand lance; EUL,

eulachon; COD, Pacific cod; COH, coho salmon; POP,

Pacific ocean perch; ATK, Atka mackerel; TAN, Tanner

crab; PAN, pandalids; CEP, cephalopods; SST shortspine

thornyhead; ROU, rougheye rockfish; NOR, northern

rockfish; SHR, shortraker rockfish; DUS, dusky

rockfish; MSF, miscellaneous fish; FSD, fishery

discard; EUP, euphausiids; PAG, paguid; OCR, other

crabs; OSH, other shrimps; JFH, jellyfish; POL,

polychaete; CAL, calanoid; MSP, miscellaneous prey.'
142

The data in Table 2 indicates that pollock were the dominant

prey fish and were consumed mainly by arrowtooth flounder (66%),

Pacific halibut (57%), sablefish (24%), and Pacific cod (7%).

Pollock cannibalism, which accounted for only 2% of the total

stomach contents weight, was not an important phenomenon in the

Gulf of Alaska compared with the Bering Sea (Livingston 1991).

Pacific herring and capelin can be categorized as the next most

important prey fishes. Arrowtooth flounder was the main predator

of Pacific herring (herring comprised 9% of their total stomach

contents weight), whereas arrow-tooth flounder and pollock both

consumed relatively large amounts of capelin (8% and 13 % by

weight, respectively). The flatfish consumed by the groundfish

include arrowtooth flounder, flathead sole, Greenland turbot,

rock sole, rex sole, yellowfin sole, Pacific halibut, and Dover

sole. Arrowtooth flounder and flathead sole were consumed mostly

by Pacific halibut and Pacific cod (Table 2). Pacific sand lance

were consumed by the main predator species (arrow-tooth flounder,

Pacific halibut, sablefish, Pacific cod, and pollock) but

comprised less than 1% of the stomach content weight of each

species. Although eulachon comprised 6% of the stomach content

weight of sablefish, some might be due to undetected net feeding

because we found evidence of net feeding by sablefish at some

stations. Pacific cod (1 % each) were consumed by Pacific halibut

and sablefish. Coho salmon (5%) and Pacific ocean perch (less

than 1%) were also found in Pacific halibut stomachs. Atka

mackerel (1%) were only found in arrowtooth flounder stomachs.

The commercially important Tanner crabs were mainly consumed by
143

Pacific cod and Pacific halibut (12% and 6% of the total stomach

contents weight, respectively), though they were also consumed by

rougheye rockfish (2%) and shortspine thornyhead (1%). Table 2

shows that almost all predator species consumed a certain amount

of cephalopods (squid and octopus), of which rougheye rockfish

and shortraker rockfish were the two main predators of the

cephalopods. All predators, except northern rockfish and

shortraker rockfish, preyed on pandalid shrimp, which includes

all the Pandalus and Pandalopsis species. Among them, shortspine

thornyhead (54%), rougheye rockfish (51%), pollock (19%), and

Pacific cod (9%) were the main predators of the pandalids.

By using the proportions of the prey items in the stomachs

(values in Table 2), I calculated the Schoener's (1970) index to

show the diet overlap between groundfish species in the Gulf of

Alaska (Fig. 1). Schoener's index (Cxy) is calculated as

cxy = 1 - 0.5 ( Px,i - Py,i ),

where Px,i and Py,i are the proportions by weight of prey i in

the diets of species x and y, respectively.

The upper diagonal-section of Figure 1 shows the Schoener's

indices (percentage) between different species. For example, the

value 64 in column 3 and row 2 means the diet overlap between

arrowtooth flounder (ATF) and Pacific halibut (PH) was 64% in

term of the Schoener's index. The lower diagonal section of

Figure 1 shows the diet overlap between species by categorizing

the Schoener's indices into low (<30%), medium (30-60%), and high

(>60%) values. These values are denoted as blank cells
144

Figure 1. --Schoener's index (%) of dietary overlap of groundfish

species in the Gulf of Alaska. ATF, arrowtooth

flounder: PH, Pacific halibut: SAB, sablefish; COD,

Pacific cod; PLK, pollock; SST, shortspine thornyhead;

ROU, rougheye rockfish; SHR, shortraker rockfish; DUS,

dusky rockfish: POP, Pacific ocean perch; NOR,

northern rockfish.
145

(for <30% overlap), gray cells (for 30-60% overlap) and black

cells (for >60% overlap). For example, in -column 2, it shows

that arrowtooth flounder has high diet overlap (black cell) with

Pacific halibut, medium diet overlap (gray cell) with sablefish

and low diet overlap (blank cells) with all other species in this

study. Overall, Figure 1 shows that two flatfish, arrowtooth

flounder, and Pacific halibut that feed mainly on pollock had a

high (>60% in this study) dietary overlap value of 64%. The

overlap values between the Pacific halibut, sablefish, and

Pacific cod were medium (30-60%) since they all fed on different

proportions of shrimp, crab, and cephalopods even though they fed

mainly on fish. It is very obvious from Figure 1 that pollock

holds a very special position in the food web of the Gulf of

Alaska. It had minimal (from 19 to 31%) overlap values with the

primary piscivorous fish, arrowtooth flounder, Pacific halibut,

sablefish, and Pacific cod. Pollock also had relatively high (39

to 49%) overlap values with all but the shortraker rockfish.

This can be easily explained since the pollock's diet included

not only large amounts (39%) of euphausiids (like many

rockfishes) and shrimp (33%), but also some fish (e.g., capelin

and pollock). The combination of euphausiids, shrimp, and fish

as prey made pollock the most competitive (overlap diet with most

of other species) groundfish species in the Gulf of Alaska.

Figure 1 also shows that shortspine thornyhead and rougheye

rockfish had a high dietary overlap value (73%) because they both

fed on large amounts (54 and 51%, respectively) of pandalids.

Dusky rockfish, Pacific ocean perch, and northern rockfish also
146

had high overlap values since they all fed largely (269%) on

euphausiids.

Table 3 lists the size of the commercially important prey

consumed by each predator species. While details of the prey

size of each of the predators were presented in the different

sections describing each species, this summary compares the prey

size of the same prey species consumed by different predators.

Table 3 shows that prey pollock size varies from species to

species. The average size (316 + 155 mm SL) of the prey pollock

consumed by Pacific halibut was the highest, whereas predator

pollock only consume smaller prey pollock (44 + 25 mm). The

herring consumed were mainly age-l or age-2 fish. The mean

length of the capelin consumed by different predator ranged from

79 to 95 mm SL. Most of the flatfish consumed were less than

200 mm. The mean size of Pacific sand lance consumed varied

greatly from 41 + 22 mm SL consumed by pollock, to 94 +. 33 mm SL

consumed by Pacific cod, to 137 + 27 mm SL consumed by Pacific

halibut. The mean lengths of eulachon consumed by the predator

species ranged from about 120 to 150 mm SL. The mean carapace

widths of the Tanner crabs ranged from 16 + 3 mm (shortspine

thornyhead) to 27 + 25 mm (sablefish). Prey Pacific cod included

larvae 14-18 mm in length consumed by pollock and the 380 mm

adult consumed by the Pacific halibut. Coho salmon (450, 500 mm

SL) and Pacific ocean perch (208 mm SL) were only found in

Pacific halibut stomachs. The only prey fish found in Pacific

ocean perch and northern rockfish were larval arrowtooth flounder

(about 20 mm SL).
147

Table 3. --The standard length (mm) of the commercially important

prey consumed by the groundfish in the Gulf of Alaska

in 1990. "+" indicates prey not measurable. "*"

indicates no data. PLK, pollock; HER, Pacific herring;

CAP, capelin; ATF, arrow-tooth flounder; FHS, flathead

sole; GT, Greenland turbot; RS, rock sole; REX, rex

sole; YFS, yellowfin sole; PH, Pacific halibut; DOV,

Dover sole; EUL, eulachon; COD, Pacific cod, COH, coho

salmon; POP, Pacific ocean perch; ATK, Atka mackerel;

TAN, Tanner crab.
148
149

Table 3. --Continued.
150

CITATIONS

Livingston, P. A. 1991. Walleye pollock, p. 9-30. In P. A.

Livingston (ed.), Groundfish food habits and predation on

commercially important prey species in the eastern Bering

Sea from 1984 to 1986. U.S. Dep. Commer., NOAA Tech. Memo.

NMFS F/NWC-207.

Schoener, T. W. 1970. Non-synchronous spatial overlap of

lizards in patchy habitats. Ecology 51:408-418.
RECENT TECHNICAL MEMORANDUMS

Copies of this and other NOAA Technical Memorandums are available from the

National Technical Information Service, 5285 Port Royal Road, Springfield, VA 22167.

Paper copies vary in price. Microfiche copies cost $3.50.

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KINOSHITA, R. K., and J. M. TERRY. 1993. Oregon, Washington, and Alaska exports of edible

fishery products, 1992, 52 p. NTIS No. PB93-226652.

REEVES, J. E. 1993. Use of lower minimum size limits to reduce discards in the Bristol Bay red

king crab (Paralithodes camtschaticus) fishery, 16 p. NTIS number pending.

SYRJAlA, S. E. 1993. Species-specific stratification and the estimate of groundfish biomass in

the Eastern Bering Sea, 20 p. NTIS number pending.

PELIA, J., M. HOFFMAN, S. HOFFMAN, M. MASUDA, S. NELSON, and L. TALLEY. 1993. Adult

sockeye and pink salmon tagging experiments for separating stocks in northern British Columbia

and southern Southeast Alaska, 1982-1985, 134 p. NTIS No. PB93-226660.

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and ship-based surveys of Steller sea lions (Eumetopias jubatus) in Southeast Alaska, the Gulf of

Alaska, and Aleutian Islands during June and July 1992, 57 p. NTIS No. PB93-226025.

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KINOSHITA, R. K., A. GREIG, L. E. QUEIROLO, and J. M. TERRY. 1993. Economic status of

the groundfish fisheries off Alaska, 1991, 94 p. NTIS No. PB93-197861.

PEREZ, M. A., and W. B. MCALISTER. 1993. Estimates of foodconsumption by marine mammals

in the Eastern Bering Sea, 36 p. NTIS No. PB93-191195.

BERGER, J. D. 1993. Comparisons between observed and reported catches of retained and

discarded groundfish in the Bering Sea and the Gulf of Alaska, 89 p. NTIS No. PB93;184711.

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LIVINGSTON, P. A., A. WARD, G. M. LANG, and M-S. YANG. 1993. Groundfish food habits and

predation on commercially important prey species in the eastern Bering Sea from 1987 to 1989,

192 p. NTIS No. PB93-184703.

KINOSHITA, R. K., and J. M. TERRY. 1993. Oregon, Washington, and Alaska exports of edible

fishery products, 1991, 47 p. NTIS No. PB93-159101.

KARINEN, J. F., M. M. BABCOCK, D. W. BROWN, W. D. MACLEOD, JR., L. S. RAMOS, and J.W.

SHORT. 1993. Hydrocarbons in intertidal sediments and mussels from Prince William Sound,

Alaska, 1977-1980: Characterization and probable sources, 69 p. NTIS No. PB93-159093.

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Sea and continental shelf, 190 p. NTIS No. PB93-156677.