NOAA Technical Memorandum NMFS-AFSC-3

U.S. DEPARTMENT OF COMMERCE

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

Alaska Fisheries Science Center

August 1992

by

Paula A. Johnson, Stanley D. Rice,and Malin M. Babcock (compilers)

Impacts of Oil Pollution and

Prince William Sound Studies:

Bibliography of 1960-91 Publications

and Reports, Auke Bay Laboratory
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:

Johnson, P. A., S. D. Rice, and M. M. Babcock (compilers). 1992. Impacts

of oil pollution and Prince William Sound studies: Bibliography of 1960-91

publications and reports, Auke Bay Laboratory. U.S. Dep. Commer.,

NOAA Tech. Memo. NMFS-AFSC-3, 98 p. 

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

National Marine Fisheries Service, NOAA.
NOAA Technical Memorandum NMFS-AFSC-3

Impacts of Oil Pollution and

Prince William Sound Studies:

Bibliography of 1960-91 Publications

and Reports, Auke Bay Laboratory

Paula A. Johnson, Stanley D. Rice,

and Malin M. Babcock (compilers)

Auke Bay Laboratory

Alaska Fisheries Science Center

11305 Glacier Highway

Juneau, AK 99801-8626

U.S. DEPARTMENT OF COMMERCE

Barbara Hackman Franklin, Secretary

National Oceanic and Atmospheric Administration

John A. Knauss, Administrator

National Marine Fisheries Service

William W. Fox, Jr., Assistant Administrator for Fisheries

August 1992
This document is available to the public through:

National Technical Information Service

U.S. Department of Commerce

5285 Port Royal Road

Springfield, VA 22161
Notice to Users of this Document 

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iii

CONTENTS

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

BIBLIOGRAPHY . . . . . . . . . . . . . . . . . . . . . .

Effects of oil . . . . . . . . . . . . . . . . . . . . .

3

3

Prince William Sound and Related Papers/Reports . . . . 14

1992 Addendum . . . . . . . . . . . . . . . . . . . . . 19

ABSTRACTS ......................... . 21

Effects of Oil ................. . . .. 21

Prince William Sound and Related Papers/Reports ... . 65

1992 Addendum .................... . 88

INDEXES .......................... . 91

Keyword Index .................... . 91

Author Index ..................... . 96
INTRODUCTION

This bibliography lists 137 publications and reports

authored by staff members of the Auke Bay Fisheries Laboratory

(ABL) and was compiled in response to the numerous information

requests following the 1989 Exxon Valdez oil spill. It covers

research done in the early 1960s from the Olsen Bay Field Station

in Prince William Sound (including some post-1964 earthquake

studies), work conducted along the Alaska Peninsula, and

laboratory research on the effects of petroleum hydrocarbons on

many of Alaska's fish and shellfish (primarily temperate or

subarctic species).

The listing is organized into three parts: BIBLIOGRAPHY,

ABSTRACTS, and INDEXES. The BIBLIOGRAPHY and ABSTRACTS sections

are each subdivided into 1) Effects of Oil, 2) Prince William

Sound and Related Papers/Reports, and 3) 1992 Addendum containing

two manuscripts in preparation, then arranged alphabetically by

author and coauthor(s) and chronologically by year. The INDEXES

section is arranged alphabetically under 1)Keyword and 2)Author.

Each bibliographic entry has a corresponding abstract entry

as originally written by the authors of that publication or

report, with the following exceptions: abstracts or summaries

for 11 entries were written specifically for this Technical

Memorandum; and lengthy abstracts were shortened by the

compilers, who accept all responsibility for interpretation.

A limited number of all reports and publications listed in

this bibliography are available from:

Library

National Marine Fisheries Service

Auke Bay Laboratory

11305 Glacier Highway

Juneau, AK 99801-8626.

All the reports and publications (except those in the 1992

Addendum) are also available from:

Oil Spill Public Information Center

645 G Street

Anchorage, AK 99501.
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3

BIBLIOGRAPHY

Effects of Oil

1 Babcock, M. M. 1985. Morphology of olfactory epithelium of

pink salmon, Oncorhynchus gorbuscha, and changes following

exposure to benzene: A scanning electron microscopy study.

In J. S. Gray and M. E. Christiansen (editors), Marine

Biology of Polar Regions and Effects of Stress on Marine

Organisms, p. 259-267. John Wiley & Sons, Chichester,

England.

2 Babcock, M. M., and J. F. Karinen. 1988. Reproductive

success in Tanner (Chionoecetes bairdi) and Dungeness

(Cancer magister) crabs held on oiled sediment.

J. Shellfish Res. 7:109.

3 Benville, P. E., and S. Korn. 1977. The acute toxicity of

six monocyclic aromatic crude oil components to striped bass

(Morone saxatilis) and bay shrimp (Crago franciscorum).

Calif. Fish Game 63:204-209.

4 Brodersen, C. 1987. Rapid narcosis and delayed mortality

in larvae of king crabs and kelp shrimp exposed to the

water-soluble fraction of crude oil. Mar. Environ. Res.

22:233-239.

5 Brodersen, C. C., S. D. Rice, J. W. Short, T. A.

Mecklenburg, and J. F. Karinen. 1977. Sensitivity of

larval and adult Alaskan shrimp and crabs to acute exposures

of the water-soluble fraction of Cook Inlet crude oil. In

1977 Oil Spill Conference (Prevention, Behavior, Control,

Cleanup), p. 575-578. American Petroleum Institute,

Washington, DC.

6 Carls, M. G. 1987. Effects of dietary and water-borne oil

exposure on larval Pacific herring (Clupea harengus

pallasi). Mar. Environ. Res. 22:253-270.

7 Carls, M. G. 1978. Some toxic and behavioral effects of

number 2 fuel oil on the eggs and larvae of Scomber scombrus

L. and Gadus morhua L. M.S. Thesis, Dalhousie Univ.,

Halifax, Nova Scotia, 96 p.

8 Carls, M. G., and S. Korn. 1985. Sensitivity of Arctic

marine amphipods and fish to petroleum hydrocarbons. Can.

Tech. Rep. Fish. Aquat. Sci. 1368:11-26.
4

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11

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17

18

Carls, M. G., and S. D. Rice. 1990. Abnormal development

and growth reductions of pollock Theragra chalcogramma

embryos exposed to water-soluble fractions of oil. Fish.

Bull., U.S. 88:29-37.

Carls, M. G., and S. D. Rice. 1988. Sensitivity

differences between eggs and larvae of walleye pollock

(Theragra chalcogramma) to hydrocarbons. Mar. Environ. Res.

26:285-297.

Carls, M. G., and S. D. Rice. 1984. Toxic contributions of

specific drilling mud components to larval shrimp and crabs.

Mar. Environ. Res. 12:45-62.

Carls, M. G., and S. D. Rice. 1981. Toxicity of oil-well

drilling muds to Alaskan larval shrimp and crabs. In

Environmental Assessment of the Alaskan Continental Shelf.

Final Reports of Principal Investigators, Vol. 11

Biological Studies, p. 1-35. U.S. Dep. Commer., NOAA.

Available Arctic Environmental Assessment Center, 222 W. 8th

Ave., No. 56, Anchorage, AK 99513.

Collodi, P. 1983. Effect of petroleum hydrocarbons on

hepatic aryl hydrocarbon hydroxylase activity in coho salmon

(Oncorhynchus kisutch). M.S. Thesis, Univ. Alaska, Juneau,

120 p.

Collodi, P., M. S. Stekoll, and S. D. Rice. 1984. Hepatic

aryl hydrocarbon hydroxylase activities in coho salmon

(Oncorhynchus kisutch) exposed to petroleum hydrocarbons.

Comp. Biochem. Physiol. C Comp. Pharmacol. 79:337-341.

Evans, D. R., and S. D. Rice. 1974. Effects of oil on

marine ecosystems: A review for administrators and policy

makers. Fish. Bull., U.S. 72:625-638.

Gharrett, J. A. 1986. Effects of a hydrocarbon -

contaminated diet on survival, feeding, growth, and molting

of juvenile red king crab, Paralithodes camtschatica

(Tilesius). M.S. Thesis, Univ. Alaska, Juneau, 178 p.

Gharrett, J. A., and S. D. Rice. 1987. Influence of

simulated tidal cycles on aromatic hydrocarbon uptake and

elimination by the shore crab Hemigrapsus nudus. Mar. Biol.

95:365-370.

Gharrett, J. A., S. D. Rice, and M. S. Stekoll. 1985.

Influence of an oil-contaminated diet on feeding rates,

growth, and molting success of juvenile red king crabs,

Paralithodes camtschatica. In Proceedings of the
5

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20

21

22

23

24

25

26

International King Crab Symposium, January 1985, Anchorage,

Alaska, p. 371-375. Univ. Alaska, Fairbanks. Alaska Sea

Grant Coll. Program, Alaska Sea Grant Rep. 85-12.

Karinen, J. F. 1988. Sublethal effects of petroleum on

biota. In D. G. Shaw and M. J. Hameedi (editors),

Environmental Studies in Port Valdez, Alaska: A Basis for

Management, p. 293-328. Springer-Verlag, Heidelberg.

Karinen, J. F. 1985. Occurrence of juvenile king crabs,

Paralithodes camtschatica (Tilesius), in Auke Bay, Alaska,

on sediments with relatively high concentrations of aromatic

hydrocarbons. In Proceedings of the International King Crab

Symposium, January 1985, Anchorage, Alaska, p. 377-387.

Univ. Alaska, Fairbanks. Alaska Sea Grant Coll. Program,

Alaska Sea Grant Rep. 85-12.

Karinen, J. F. 1980. Petroleum in the deep sea

environment: Potential for damage to biota. Environ. Int.

3:135-144.

Karinen, J. F. 1977. Assessing oil impacts with laboratory

data applications, limitations, and needs. In B. Melteff

(editor), Oil and Aquatic Ecosystems, Tanker Safety and Oil

Pollution Liability. Proceedings of the Cordova Fisheries

Institute, April 1-3, 1977, Cordova, Alaska, p. 99-110.

Univ. Alaska, Fairbanks, Alaska Sea Grant Rep. 77-8.

Karinen, J. F., and S. D. Rice. 1974. Effects of Prudhoe

Bay crude oil on molting Tanner crabs, Chionoecetes bairdi.

Mar. Fish. Rev. 36(7):31-37.

Karinen, J. F., S. D. Rice, and M. M. Babcock. 1990.

Reproductive success in Dungeness crab (Cancer magister)

during long-term exposures to oil-contaminated sediments.

In Outer Continental Shelf Assessment Program, Final Reports

of Principal Investigators, Vol. 67, p. 435-461. U.S. Dep.

Commer., NOAA. Available Arctic Environmental Assessment

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

Korn, S., N. Hirsch, and J. W. Struhsaker. 1977 The

uptake, distribution, and depuration of 14C benzene and 14C

toluene in Pacific herring, Clupea harengus pallasi. Fish.

Bull., U.S. 75:633-636.

Korn, S., N. Hirsch, and J. W. Struhsaker. 1976. Uptake,

distribution, and depuration of 14C-benzene in northern

anchovy, Engraulis mordax, and striped bass, Morone

saxatilis. Fish. Bull., U.S. 74:545-551.
6

27 Korn, S., D. A. Moles, and S. D. Rice. 1979. Effects of

temperature on the median tolerance limit of pink salmon and

shrimp exposed to toluene, naphthalene, and Cook Inlet crude

oil. Bull. Environ. Contam. Toxicol. 21:521-525.

28 Korn, S., and S. Rice. 1981. Sensitivity to, and

accumulation and depuration of, aromatic petroleum

components by early life stages of coho salmon (Oncorhynchus

kisutch). Rapp. P.-V. Reun. Cons. Int. Explor. Mer

178:87-22.

29 Korn, S., S. D. Rice, D. L. Cheatham, and D. W. Brown.

1985. Contribution of phenol and p-cresol to the toxicity

of crude oil to pink salmon (Oncorhynchus gorbuscha) fry and

kelp shrimp (Eualus suckleyi). In F. J. Vernberg, F. P.

Thurberg, A. Calabrese, and W. B. Vernberg (editors), Marine

Pollution and Physiology: Recent Advances, p. 447-458.

Univ. South Carolina Press, Columbia, SC.

30 Korn, S., J. W. Struhsaker, and P. Benville. 1976. Effects

of benzene on growth, fat content, and caloric content of

striped bass, Morone saxatilis. Fish. Bull., U.S.

74:694-698.

31 Lauren, D. J., and S. Rice. 1985. Significance of active

and passive depuration in the clearance of naphthalene from

the tissues of Hemigrapsus nudus (Crustacea: Decapoda).

Mar. Biol. 88:135-142.

32 Mecklenburg, T. A., S. D. Rice, and J. F. Karinen. 1977.

Molting and survival of king crab (Paralithodes

camtschatica) and coonstripe shrimp (Pandalus hypsinotus)

larvae exposed to Cook Inlet crude oil water-soluble

fraction. In D. A. Wolfe (editor), Fate and Effects of

Petroleum Hydrocarbons in Marine Ecosystems and Organisms,

p. 221-228. Pergamon Press, New York.

33 Merrell, T. R., Jr. 1988. Fisheries resources. In D. G.

Shaw and M. J. Hameedi (editors), Environmental Studies in

Port Valdez, Alaska: A Basis for Management, p. 203-224.

Springer-Verlag, New York.

34 Moles, A. 1980. Sensitivity of parasitized coho salmon fry

to crude oil, toluene, and naphthalene. Trans. Am. Fish.

soc. 109:293-297.

35 Moles, D. A. 1977. Effect of parasitism by glochidia of

Anodonta oregonensis Lea on the sensitivity of coho salmon

fry, Oncorhynchus kisutch Walbaum, to the water-soluble

fraction of Prudhoe Bay crude oil and the effect of crude

oil on the development of the glochidium. M.S. Thesis,

Univ. Alaska, Juneau, 50 p.
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44

Moles, A., M. M. Babcock, and S. D. Rice. 1987. Effects of

oil exposure on pink salmon, Oncorhynchus gorbuscha, alevins

in a simulated intertidal environment. Mar. Environ. Res.

21:49-58.

Moles, A., S. Bates, S. D. Rice, and S. Korn. 1981.

Reduced growth of coho salmon fry exposed to two petroleum

components, toluene and naphthalene, in fresh water. Trans.

Am. Fish. Soc. 110:430-436.

Moles, A., and S. D. Rice. 1983. Effects of crude oil and

naphthalene on growth, caloric content, and fat content of

pink salmon juveniles in seawater. Trans. Am. Fish. Soc.

112:205-211.

Moles, A., S. D. Rice, and S. Andrews. 1985.

Continuous-flow devices for exposing marine organisms to the

water-soluble fraction of crude oil and its components.

Can. Tech. Rep. Fish. Aquat. Sci. 1368:53-61.

Moles, A., S. D. Rice, and S. Korn. 1979. Sensitivity of

Alaskan freshwater and anadromous fishes to Prudhoe Bay

crude oil and benzene. Trans. Am. Fish. Soc. 108:408-414.

Myren, R. T., and J. J. Pella. 1977. Natural variability

in distribution of an intertidal population of Macoma

balthica subject to potential oil pollution at Port Valdez,

Alaska. Mar. Biol. 41:371-382.

O'Clair, C. E., J. L. Hanson, J. S. MacKinnon, J. A.

Gharrett, N. I. Calvin, T. R. Merrell, Jr. 1978. Baseline/

reconnaissance characterization littoral biota, Gulf of

Alaska and Bering Sea. In Environmental Assessment of the

Alaskan Continental Shelf. Annual Reports of Principal

Investigators for the Year Ending March 1978. Volume 4,

Receptors--Fish, Littoral, Benthos, p. 256-415. U.S. Dep.

Commer., NOAA. Available Arctic Environmental Assessment

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

O'Clair, C. E., and S. D. Rice. 1985. Depression of

feeding and growth rates of the seastar Evasterias

troschelii during long-term exposure to the water-soluble

fraction of crude oil. Mar. Biol. 84:331-340.

Pelto, M. J., R. W. Williamson, and J. Knull. 1978.

Dispersion of drogues: An estimation of pollutant dispersal

in Port Valdez, Alaska. Northwest and Alaska Fisheries

Center Processed Report, 10 p. Northwest and Alaska Fish.

Cent., Natl. Mar. Fish. Serv., NOAA, 2725 Montlake Blvd. E.,

Seattle, WA 98112.
8

45 Rice, S. D. 1985. Effects of oil on fish. In F. R.

Engelhardt (editor), Petroleum Effects in the Arctic

Environment, p. 157-182. Elsevier Applied Science

Publishers, New York.

46 Rice, S. D. 1985. Fisheries and oil: Facts and figures.

Alaska Fish and Game 18[17](6):35-37.

47 Rice, S. D. 1977. A review of oil toxicity studies

conducted at the Auke Bay Laboratory. In B. Melteff

(editor), Oil and Aquatic Ecosystems, Tanker Safety and Oil

Pollution Liability, Proceedings of the Cordova Fisheries

Institute, April 1-3, 1977, Cordova, Alaska, p. 111-113.

Univ. Alaska, Fairbanks, Alaska Sea Grant Rep. 77-8.

48 Rice, S. D. 1974. Toxicity and avoidance tests with

Prudhoe Bay oil and pink salmon fry. In 1973 Proceedings of

the Joint Conference on Prevention and Control of Oil

spills, March 13-15, 1973, Washington, DC, p. 667-670.

American Petroleum Institute, Washington, DC.

49 Rice, S. D., and M. M. Babcock. 1989. Effects of habitat

and environmental variables on red king crabs, and settling

of glaucothoe. In L. E. Jarvela and L. K. Thorsteinson

(editors), Proceedings of the Gulf of Alaska, Cook Inlet,

and North Aleutian Basin Information Update Meeting,

February 7-8, 1989, Anchorage, Alaska, p. 49-53. Available

National Marine Fisheries Service, Auke Bay Laboratory.

50 Rice, S. D., M. M. Babcock, C. C. Brodersen, M. G. Carls,

J. A. Gharrett, S. Korn, A. Moles, and J. W. Short. 1987.

Lethal and sublethal effects of the water-soluble fraction

of Cook Inlet crude oil on Pacific herring (Clupea harengus

pallasi) reproduction. U.S. Dep. Commer., NOAA Tech. Memo.

NMFS F/NWC-111, 63 p.

51 Rice, S. D., M. M. Babcock, C. C. Brodersen, J. A. Gharrett,

and S. Korn. 1987. Uptake and depuration of aromatic

hydrocarbons by reproductively ripe Pacific herring and the

subsequent effect of residues on egg hatching and survival.

In W. B. Vernberg, A. Calabrese, F. P. Thurberg, and F. J.

Vernberg (editors), Pollution Physiology of Estuarine

Organisms, p. 139-154. Univ. South Carolina Press,

Columbia, SC.

52 Rice, S. D. and J. F. Karinen. 1976. Acute and chronic

toxicity, uptake and depuration, and sublethal metabolic

response of Alaska marine organisms to petroleum

hydrocarbons. In Environmental Assessment of the Alaskan

Continental Shelf. Annual Reports of Principal

Investigators for the Year Ending March 1976, Vol. 8,

Effects of contaminants, p. 25-118. U.S. Dep. Commer.,
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NOM. Available Arctic Environmental Assessment Center,

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

Rice, S. D., J. F. Karinen, and C. C. Brodersen. 1985.

Effects of oiled sediment on juvenile king crab. In Outer

Continental Shelf Assessment Program, Final Reports of

Principal Investigators, Vol. 29, p. 287-310. U.S. Dep.

Commer., NOAA. Available Arctic Environmental Assessment

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

Rice, S. D., J. F. Karinen, and S. Korn. 1978. Acute and

chronic toxicity, uptake and depuration, and sublethal

response of Alaska marine organisms to petroleum

hydrocarbons. In D. A. Wolfe (editor), Marine Biological

Effects of OCS Petroleum Development, p. 11-24. U.S. Dep.

Commer., NOM Tech. Memo. ERL-OCSEAP-1.

Rice, S. D., S. Korn, C. C. Brodersen, S. A. Lindsay, and

S. A. Andrews. 1981. Toxicity of ballast-water treatment

effluent to marine organisms at Port Valdez, Alaska. In

Proceedings 1981 Oil Spill Conference, Atlanta, Georgia,

March 2-5, p. 55-61. American Petroleum Institute,

Washington, DC.

Rice, S. D., S. Korn, and J. F. Karinen. 1981. Lethal and

sublethal effects on selected Alaskan marine species after

acute and long-term exposure to oil and oil components. In

Environmental Assessment of the Alaskan Continental Shelf.

Annual Reports of Principal Investigators for the Year

Ending March 1981, Vol. 4, Effects of contaminants,

p. 61-78. U.S. Dep. Commer., NOAA. Available Arctic

Environmental Assessment Center, 222 W. 8th Ave., No. 56,

Anchorage, AK 99513.

Rice S. D., S. Korn, and J. F. Karinen. 1980. Lethal and

sublethal effects on selected Alaskan marine species after

acute and long-term exposure to oil and oil components. In

Environmental Assessment of the Alaskan Continental Shelf.

Annual Reports of Principal Investigators for the Year

Ending March 1980, Vol. 3, Effects, Contaminant Baselines,

p. 1-12. U.S. Dep. Commer., NOAA. Available Arctic

Environmental Assessment Center, 222 W. 8th Ave., No. 56,

Anchorage, AK 99513.

Rice, S. D., S. Korn, and J. F. Karinen. 1978. Lethal and

sublethal effects on selected Alaskan marine species after

acute and long-term exposure to oil and oil components. In

Environmental Assessment of the Alaskan Continental Shelf.

Final Reports of Principal Investigators, Vol. 1, Biological

Studies, p. 1-32. U.S. Dep. Commer., NOM. Available

Arctic Environmental Assessment Center, 222 W. 8th Ave.,

No. 56, Anchorage, AK 99513.
10

59 Rice, S. D., S. Korn, and J. F. Karinen. 1977. Lethal and

sublethal effects on selected Alaskan marine species after

acute and long-term exposure to oil and oil components. In

Environmental Assessment of the Alaskan Continental Shelf.

Annual Reports of Principal Investigators for the Year

Ending March 1977, Vol. 12, Effects, p. 23-124. U.S. Dep.

Commer., NOAA. Available Arctic Environmental Assessment

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

60 Rice, S. D., and D. A. Moles. 1991. Humpies and

hydrocarbons: Relating laboratory effects studies to

potential effects from the Exxon Valdez oil spill.

Unpublished manuscript. National Marine Fisheries Service,

Auke Bay Laboratory.

61 Rice, S. D., D. A. Moles, J. F. Karinen, S. Korn, M. G.

Carls, C. C. Brodersen, J. A. Gharrett, and M. M. Babcock.

1984. Effects of petroleum hydrocarbons on Alaskan aquatic

organisms: A comprehensive review of all oil-effects

research on Alaskan fish and invertebrates conducted by the

Auke Bay Laboratory, 1970-81. U.S. Dep. Commer., NOAA Tech.

Memo. NMFS F/NWC-67, 128 p.

62 Rice, S. D., A. Moles, and J. W. Short. 1975. The effect

of Prudhoe Bay crude oil on survival and growth of eggs,

alevins, and fry of pink salmon, Oncorhynchus gorbuscha. In

Proceedings of 1975 Conference on Prevention and Control of

Oil Pollution, p. 503-507. American Petroleum Institute,

Washington, DC.

63 Rice, S. D., D. A. Moles, T. T. Taylor, and J. F. Karinen.

1979. Sensitivity of 39 Alaskan marine species to Cook

Inlet crude oil and No. 2 fuel oil. In Proceedings of 1979

Oil Spill Conference (Prevention, Behavior, Control,

Cleanup), p. 549-554. American Petroleum Institute,

Washington, DC.

64 Rice, S. D., J. W. Short, C. C. Brodersen, T. A.

Mecklenburg, D. A. Moles, C. J. Misch, D. L. Cheatham, and

J. F. Karinen. 1976. Acute toxicity and uptake-depuration

studies with Cook Inlet crude oil, Prudhoe Bay crude oil,

No. 2 fuel oil and several subarctic marine organisms.

Northwest Fisheries Center Processed Report, 90 p.

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

NOAA, 2725 Montlake Blvd. E., Seattle, WA 98112.

65 Rice, S. D., J. W. Short, and J. F. Karinen. 1977.

Comparative oil toxicity and comparative animal sensitivity.

In D. A. Wolfe (editor), Fate and Effects of Petroleum

Hydrocarbons in Marine Ecosystems and Organisms, p. 78-94.

Pergamon Press, New York.
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Rice, S. D., J. W. Short, and J. F. Karinen. 1976.

Toxicity of Cook Inlet crude oil and No. 2 fuel oil to

several Alaskan marine fishes and invertebrates. In

Sources, Effects, and Sinks of Hydrocarbons in the Aquatic

Environment, p. 395-406. American Institute of Biological

Sciences, Washington, DC.

Rice, S. D., and R. E. Thomas. 1989. Effect of

pre-treatment exposures of toluene or naphthalene on the

tolerance of pink salmon (Oncorhynchus gorbuscha) and kelp

shrimp (Eualis [sic] suckleyi). Comp. Biochem. Physiol. C

Comp. Pharmacol. 94:289-293.

Rice, S. D., R. E. Thomas, and J. W. Short. 1977. Effect

of petroleum hydrocarbons on breathing and coughing rates

and hydrocarbon uptake-depuration in pink salmon fry. In F.

J. Vernberg, A. Calabrese, F. P. Thurberg, and W. B.

Vernberg (editors), Physiological Responses of Marine Biota

to Pollutants, p. 259-277. Academic Press, New York.

Schwartz, J. P. 1985. Effect of oil-contaminated prey on

the feeding and growth rate of pink salmon fry (Oncorhynchus

gorbuscha). In F. J. Vernberg, A. Calabrese, and W.

Vernberg (editors), 'Marine Pollution and Physiology: Recent

Advances, p. 459-476. Univ. South Carolina Press, Columbia,

SC.

Schwartz, J. P. 1984. The effects of oil-contaminated prey

on the feeding, growth, and related energetics of pink

salmon, Oncorhynchus gorbuscha Walbaum, fry. Ph.D. Thesis,

Univ. Alaska, Fairbanks, 99 p.

Short, J. W., S. D. Rice, and D. L. Cheatham. 1976.

Comparison of two methods for oil and grease determination.

In D. W. Hood and D. C. Burrell (editors.) Assessment of the

Arctic Marine Environment: Selected Topics, p. 451-465.

Univ. Alaska, Fairbanks.

Stickle, W. B., M. A. Kapper, T. C. Shirley, M. G. Carls,

and S. D. Rice. 1987. Bioenergetics and tolerance of the

pink shrimp (Pandalus borealis) during long-term exposure to

the water-soluble fraction and oiled sediment from Cook

Inlet crude oil. In W. B. Vernberg, A. Calabrese, F. P.

Thurberg, and F. J. Vernberg (editors), Pollution Physiology

of Estuarine Organisms, p. 87-106. Univ. South Carolina

Press, Columbia, SC.

Stickle, W. B., S. D. Rice, and A. Moles. 1984.

Bioenergetics and survival of the marine snail Thais lima

during long-term oil exposure. Mar. Biol. 80:281-289.
12

74 Stickle, W. B., Jr., S. D. Rice, C. Villars, and W. Metcalf.

1985. Bioenergetics and survival of the marine mussel,

Mytilus edulis L., during long-term exposure to the

water-soluble fraction of Cook Inlet crude oil. In F. J.

Vernberg, F. P. Thurberg, A. Calabrese, and W. B. Vernberg

(editors), Marine Pollution and Physiology: Recent

Advances, p. 427-446. Univ. South Carolina Press, Columbia,

SC.

75 Stickle, W. B., T. D. Sabourin, and S. D. Rice. 1982.

Sensitivity and osmoregulation of coho salmon, Oncorhynchus

kisutch, exposed to toluene and naphthalene at different

salinities. In W. B. Vernberg, A. Calabrese, F. P.

Thurberg, and F. J. Vernberg (editors), Physiological

Mechanisms of Marine Pollutant Toxicity, p. 331-348.

Academic Press, New York.

76 Taylor, T. L., and J. F. Karinen. 1977. Response of the

clam, Macoma balthica (Linnaeus), exposed to Prudhoe Bay

crude oil as unmixed oil, water-soluble fraction, and

oil-contaminated sediment in the laboratory. In D. A. Wolfe

(editor), Fate and Effects of Petroleum Hydrocarbons in the

Marine Ecosystems and Organisms, p. 229-237. Pergamon

Press, New York.

77 Thomas, R. E., and S. D. Rice. 1987. Effect of

water-soluble fraction of Cook Inlet crude oil on swimming

performance and plasma cortisol in juvenile coho salmon

(Oncorhynchus kisutch). Comp. Biochem. Physiol. C Comp.

Pharmacol. Toxicol. 87:177-180.

78 Thomas, R. E., and S. D. Rice. 1986. The effects of

salinity on uptake and metabolism of toluene and naphthalene

by Dolly Varden, Salvelinus malma. Mar. Environ. Res.

18:203-214.

79 Thomas, R. E., and S. D. Rice. 1986. Effect of temperature

on uptake and metabolism of toluene and naphthalene by Dolly

Varden char, Salvelinus malma. Comp. Biochem. Physiol. C

Comp. Pharmacol. Toxicol. 84:83-86.

80 Thomas, R. E., and S. D. Rice. 1985. Effect of

pretreatment exposure to toluene and naphthalene on the

subsequent metabolism of dietary toluene and naphthalene by

Dolly Varden, Salvelinus malma. In F. J. Vernberg, F. P.

Thurberg, A. Calabrese and W. A. Vernberg (editors), Marine

Pollution and Physiology: Recent advances, p. 505-520.

Univ. South Carolina Press, Columbia, SC.
13

81 Thomas, R. E., and S. D. Rice. 1982. Metabolism and

clearance of phenolic and mono-, di-, and polynuclear

aromatic hydrocarbons by Dolly Varden char. In W. B.

Vernberg, A. Calabrese, F. P. Thurberg, and F. J. Vernberg

(editors), Physiological Mechanisms of Marine Pollutant

Toxicity, p. 161-176. Academic Press, New York.

82 Thomas, R. E., and S. D. Rice. 1981. Excretion of aromatic

hydrocarbons and their metabolites by freshwater and

seawater Dolly Varden char. In F. J. Vernberg,

A. Calabrese, F. P. Thurberg, and W. B. Vernberg (editors),

Biological Monitoring of Marine Pollutants, p. 425-448.

Academic Press, New York.

83 Thomas, R. E., and S. D. Rice. 1979. The effect of

exposure temperature on oxygen consumption and opercular

breathing rates of pink salmon fry exposed to toluene,

naphthalene, and water-soluble fractions of Cook Inlet crude

oil and No. 2 fuel oil. In W. B. Vernberg, A. Calabrese, F.

P. Thurberg, and F. J. Vernberg (editors), Marine Pollution:

Functional Responses, p. 39-52. Academic Press, New York.

84 Thomas, R. E., and S. D. Rice. 1975. Increased opercular

rates of pink salmon (Oncorhynchus gorbuscha) fry after

exposure to the water-soluble fraction of Prudhoe Bay crude

oil. J. Fish. Res. Board Can. 32:2221-2224.

85 Thomas, R. E., S. D. Rice, M. M. Babcock, and A. Moles.

1989. Differences in hydrocarbon uptake and mixed function

oxidase activity between juvenile and spawning adult coho

salmon (Oncorhynchus kisutch) exposed to Cook Inlet crude

oil. Comp. Biochem. Physiol. C Comp. Pharmacol.

93:155-159.

86 Thomas, R. E., S. D. Rice, M. M. Babcock, and A. Moles.

1988. Differences in hydrocarbon uptake and mixed function

oxidase activity between juvenile and spawning adult coho

salmon exposed to Cook Inlet crude oil. Am. Zool.

28(4):104A. (Abstract only).

87 Thomas, R. E., S. D. Rice, and S. Korn. 1987. Reduced

swimming performance of juvenile coho salmon (Oncorphynchus

[sic] kisutch) exposed to the water-soluble fraction of Cook

Inlet crude oil. In W. B. Vernberg, A. Calabrese, F. P.

Thurberg, and F. J. Vernberg (editors), Pollution Physiology

of Estuarine Organisms, p. 127-137. Univ. South Carolina

Press, Columbia, SC.
14

Prince William Sound and Related Papers/Reports

88

89

90

91

92

93

94

95

96

97

98

99

Barr, L. 1970. Alaska's fishery resources: The shrimps.

U.S. Fish Wildl. Serv. Fish. Leafl. 631, 10 p.

Barr, L. 1970. Diel vertical migration of Pandalus

borealis in Kachemak Bay, Alaska. J. Fish. Res. Board. Can.

27:669-676.

Calvin, N. I., and R. J. Ellis. 1978. Quantitative and

qualitative observations of Laminaria dentigera and other

subtidal kelps of southern Kodiak Island, Alaska. Mar.

Biol. 47:331-336.

Calvin, N. I., and S. C. Lindstrom. 1980. Intertidal algae

of Port Valdez, Alaska: Species and distribution with

annotations. Bot. Mar. 23:791-797.

Dahlberg, M. L. 1979. History of fishery and summary

statistics of the sockeye salmon, Oncorhynchus nerka, runs

to the Chignik Lakes, Alaska, 1888-1966. NOM (Natl.

Oceanic Atmos. Adm.) Tech. Rep. NMFS (Natl. Mar. Fish.

Serv.) SSRF (Spec. Sci. Rep. Fish.) 735, 16 p.

Dahlberg, M. L. 1973. Stock-and-recruitment relationships

and optimum escapements of sockeye salmon stocks of the

Chignik Lakes, Alaska. Rapp. P.-V. Reun. Cons. Perm. Int.

Explor. Mer 164:98-105.

Dahlberg, M. L. 1968. Analysis of the dynamics of sockeye

salmon returns to the Chignik Lakes, Alaska. Ph.D. Thesis,

Univ. Washington, Seattle, 337 p.

Frame, G. W. 1976. Alaska's fishing black bears. Pac.

Discovery 29(3):19-25.

Frame, G. W. 1974. Black bear predation on salmon at Olsen

Creek, Alaska. Z. Tierpsychol. 35:23-38.

Harry, G. Y. 1964. The shrimp fishery of Alaska. Proc.

Gulf Caribb. Fish. Inst. 16:64-71.

Haynes, E. 1983. Distribution and abundance of larvae of

king crab, Paralithodes camtschatica, and Pandalid shrimp in

the Kachemak Bay area, Alaska, 1972 and 1976. NOAA (Natl.

Oceanic Atmos. Adm.) Tech. Rep. NMFS (Natl. Mar. Fish.

Serv.) SSRF (Spec. Sci. Rep. Fish.) 765, 64 p.

Haynes, E. 1981. Description of stage II zoeae of snow

crab, Chionoecetes bairdi, (Oxyrhyncha, Majidae) from

plankton of lower Cook Inlet, Alaska. Fish. Bull., U.S.

79:177-182.
15

Haynes, E. 1980. Stage I zoeae of a crangonid shrimp,

Crangon franciscorum angustimana, hatched from ovigerous

females collected in Kachemak Bay, Alaska. Fish. Bull.,

U.S. 77:991-995.

100

101

102

103

104

105

106

107

108

109

Haynes, E. 1979. Description of larvae of the northern

shrimp, Pandalus borealis, reared in situ in Kachemak Bay,

Alaska. Fish. Bull., U.S. 77:157-173.

Haynes, E. 1970. Age and growth of giant Pacific sea

scallops in the Gulf of Alaska. Proc. Natl. Shellfish.

Assoc. 60:14. (Abstract only).

Haynes, E. B., and J. C. McMullen. 1970. Relation between

meat weight and shell height of the giant Pacific sea

scallop, Patinopecten caurinus, from the Gulf of Alaska.

Proc. Natl. Shellfish. Assoc. 60:50-53.

Heard, W. R., R. L. Wallace, and W. L. Hartman. 1969.

Distributions of fishes in fresh water of Katmai National

Monument, Alaska, and their zoogeographical implications.

U. S. Fish Wildl. Serv. Spec. Sci. Rep. Fish. 590, 20 p.

Helle, J. H. 1989. Relation between size-at-maturity and

survival of progeny in chum salmon, Oncorhynchus keta

(Walbaum). J. Fish Biol. 35(A):99-107.

Helle, J. H. 1984. Age and size at maturity of some

populations of chum salmon in North America. In P. A.

Moiseev (editor),

Conference,

Proceedings of the Pacific Salmon Biology

Yuzhno-Sakhalinsk, USSR, 3-13 October 1978,

p. 126-143. Pacific Scientific Institute of Fisheries and

Oceanography (TINRO),

Vladivostok, USSR.

4 Shevchenko Alley 690600,

Helle, J. H. 1982. Some effects of the marine environment

on age at maturity and growth of chum salmon in Prince

William Sound, Alaska. In B. R. Melteff and R. A. Neve

(editors), Proceedings of the North Pacific Aquaculture

Symposium, p. 91.

Fairbanks,

(Abstract only.) Univ. Alaska,

Sea Grant Rep. 82-2.

Helle, J. H. 1981. Significance of the stock concept in

artificial propagation of salmonids in Alaska. Can. J.

Fish. Aquat. Sci. 38:1665-1671.

Helle, J. H. 1979. Influence of marine environment on age

and size at maturity, growth, and abundance of chum salmon,

Oncorhynchus keta (Walbaum), from Olsen Creek, Prince

William Sound, Alaska. Ph.D. Thesis,

118 p.

Oregon State Univ.,
16

110

111

112

113

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119

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121

Helle, J. H. 1970. Biological characteristics of

intertidal and fresh-water-spawning pink salmon at Olsen

Creek, Prince William Sound, Alaska, 1962-63. U.S. Fish

Wildl. Serv. Spec. Sci. Rep. Fish. 602, 19 p.

Helle, J. H. 1966. Behavior of displaced adult pink

salmon. Trans. Am. Fish. Soc. 95:188-195.

Helle, J. H. 1960. Characteristics and structure of early

and late spawning runs of chum salmon, Oncorhynchus keta

(Walbaum), in streams of Prince William Sound, Alaska. M.S.

Thesis, Univ. Idaho, 53 p.

Helle, J. H., R. S. Williamson, and J. E. Bailey. 1964.

Intertidal ecology and life history of pink salmon at Olsen

Creek, Prince William Sound, Alaska. U.S. Fish Wildl. Serv.

Spec. Sci. Rep. Fish. 483, 26 p.

Hubbard, J. D. 1971. Distribution and abundance of

intertidal invertebrates at Olsen Bay in Prince William

Sound, Alaska, one year after the 1964 earthquake. In The

Great Alaska Earthquake of 1964: Biology, p. 137-157.

National Academy of Sciences, Washington, DC.

Kirkwood, J. B. 1962. Inshore-marine and freshwater life

history phases of the pink salmon, Oncorhynchus gorbuscha

(Walbaum) and the chum salmon O. keta (Walbaum) in Prince

William Sound, Alaska. Ph.D. Thesis, Univ. Louisville,

Louisville, KY, 300 p.

Kirkwood, J. B., and R. M. Yancey. 1965. Effects of the

March 27 earthquake upon the shellfish resources of Alaska.

Proc. Alaska Sci. Conf. 15:162.

Leatherwood, S., A. E. Bowles, E. Krygier, J. D. Hall, and

S. Ignell. 1984. Killer whales (Orcinus orca) in southeast

Alaska, Prince William Sound, and Shelikof Strait; A review

of available information. Rep. Int. Whaling Comm.

34:521-530.

Lindstrom, S. C., and N. I. Calvin. 1975. New records of

marine red algae from the Gulf of Alaska. Syesis 8:405-406.

Meehan, W. R. 1966. Growth and survival of sockeye salmon

introduced into Ruth Lake after removal of resident fish

populations. U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish.

532, 18 p.

Meyer, R. M. 1968.

Kodiak, Alaska.

The Dungeness crab fishery around

comm. Fish. Rev. 30(8-9):44-47.

Moyle, Peter. 1966. Feeding behavior of the glaucous

winged gull on an Alaska salmon stream. Wilson Bull.

78:175-190.
17

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O'Clair, C. E., and S. T. Zimmerman. 1987. Biogeography

and ecology of intertidal and shallow subtidal communities.

In D. W. Hood and S. T. Zimmerman (editors), The Gulf of

Alaska: Physical Environment and Biological Resources,

p. 305-343. NOM, U.S. Dep. Commer. Available

Superintendent of Documents, U.S. Gov. Print. Office,

Washington, DC 20402.

Olsen, J. C. 1973. Pandalid shrimp life history research

at Kachemak Bay, Alaska. Mar. Fish. Rev. 35(3-4):62-64.

Orsi, J. A., R. K. Gish, and B. L. Wing. 1991. Northern

range extensions of four nearshore marine fishes in Alaska.

Can. Field-Nat. 105:82-86.

Phinney, D. E. 1970. Spawning ground catalog of the

Chignik River system, Alaska. U. S. Fish Wildl. Serv. Data

Rep. 41, 147 p.

Quast, J. C., and E. L. Hall. 1972. List of fishes of

Alaska and adjacent waters with a guide to some of their

literature. NOAA (Natl. Oceanic Atmos. Admin.) Tech. Rep.

NMFS (Natl. Mar. Fish. Serv.) SSRF (Spec. Sci. Rep. Fish.)

658, 47 p.

Reid, G. M. 1971. Age composition, weight, length, and sex

of herring, Clupea pallasii, used for reduction in Alaska,

1929-66. NOM (Natl. Oceanic Atmos. Admin.) Tech. Rep. NMFS

(Natl. Mar. Fish. Serv.) SSRF (Spec. Sci. Rep. Fish.) 634,

25 p.

Sears, H. S., and S. T. Zimmerman. 1977. Alaska intertidal

survey atlas, 402 p. National Marine Fisheries Service,

Auke Bay Laboratory.

Skud, B. E., H. M. Sakuda, and G. M. Reid. 1960.

Statistics of the Alaska herring fishery 1878-1956. U.S.

Fish Wildl. Serv. Bur. Comm. Fish. Stat. Dig. 48, 21 p.

Tait, H. D., and J. B. Kirkwood. 1962. Estimating

abundance of pink and chum salmon fry in Prince William

Sound, 1957. U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish.

429, 21 p.

Thorsteinson, F. V. 1965. Effects of the Alaska earthquake

on pink and chum salmon runs in Prince William Sound. Proc.

Alaska Sci. Conf. 15:267-280.

Thorsteinson, F. V. 1965. Aftermaths of the Alaska earthquake in Prince William Sound. Pac. Fisherman 63(6):10-11.
18

133 Thorsteinson, F. V., J. H. Helle, and D. G. Birkholz. 1971.

Salmon survival in intertidal zones of Prince William Sound

in uplifted and subsided areas. In The Great Alaska

Earthquake of 1964: Biology, p. 194-219. National Academy

of Sciences, Washington, DC.

134 Thorsteinson, F. V., W. H. Noerenberg, and H. D. Smith.

1963. The length, age, and sex ratio of chum salmon in the

Alaska Peninsula, Kodiak Island, and Prince William Sound

area of Alaska. U. S. Fish Wildl. Serv. Spec. Sci. Rep.

Fish. 430, 84 p.

135 Zimmerman, S. T., J. L. Hanson, J. T. Fujioka, N. I. Calvin,

J. A. Gharrett, and J. S. MacKinnon. 1978. Intertidal

biota and subtidal kelp communities of the Kodiak Island

area. In Environmental Assessment of the Alaskan

Continental Shelf. Final Reports of Principal

Investigators, Vol. 4, Biological studies, p. 316-508. U.S.

Dep. Commer., NOAA. Available Arctic Environmental

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99513.
19

1992 Addendum

These two manuscripts represent research conducted before the

Exxon Valdez oil spill; both manuscripts are currently in draft

form and nearing completion.

136 Karinen, J. F., M. M. Babcock, D. W. Brown, W. D. MacLeod,

Jr., L. S. Ramos, and J. W. Short. In Preparation.

Hydrocarbons in intertidal sediments and mussels from Prince

William Sound, Alaska, 1977-1980: Characterization and

probable sources. National Marine Fisheries Service, Auke

Bay Laboratory.

137 Myren, R. T., G. Perkins, and T. R. Merrell. In

Preparation. Reduced abundance of Macoma balthica near an

oil tanker terminal in Port Valdez, Alaska, 1971-1984.

National Marine Fisheries Service, Auke Bay Laboratory.
THIS PAGE INTENTIONALLY LEFT BLANK
21

Effects of Oil

1 Babcock, M. M. 1985. Morphology of olfactory epithelium of

pink salmon, Oncorhynchus gorbuscha, and changes following

exposure to benzene: A scanning electron microscopy study.

In J. S. Gray and M. E. Christiansen (editors), Marine

Biology of Polar Regions and Effects of Stress on Marine

Organisms, p. 259-267. John Wiley & Sons, Chichester,

England.

The pink salmon fishery is the most valuable fishery in Prince,

William Sound, Alaska (USA), and this resource may be damaged by

oil pollution from tankers or discharges from a ballast-water

treatment plant near Valdez. Because juvenile pink salmon school

along shallow estuarine shorelines of Prince William Sound

several weeks before migrating to oceanic feeding grounds, they

are vulnerable to oil pollutants from these sources. Benzene, a

major component of crude oil and the effluent from the treatment

plant, is water soluble and toxic to fish. To determine the

histopathological effects of benzene on olfactory rosettes of

pink salmon, I exposed juveniles to sublethal concentrations of

benzene in seawater. Fish were exposed in seawater to 4.3 ppm

benzene for 12 d or to one of four concentrations ranging from

0.15 to 4.40 ppm benzene for 29 d. (The 96-h LC50=8.47 ppm).

Olfactory rosettes from these fish were examined with scanning

electron microscopy. Rosettes from fry exposed to >0.15 ppm

benzene had exhausted mucous cells. Olfactory lamellae of fry

exposed to ? ? ?? ??ppm benzene had altered distribution of cilia.

Lamellae of fish exposed to 4.3 ppm benzene 12 d had patchy

losses of cilia; lamellae of fish exposed 29 d to concentrations

of benzene ? ? ?? ??ppm had a generalized loss of cilia. Exhausted

mucous cells and loss of cilia on the olfactory lamellar surfaces

could change circulation of water through the olfactory rosettes

or otherwise interfere with normal chemosensory reception and

consequently affect homing,

feeding activity,

traditional migratory patterns,

and avoidance of predators.

2 Babcock, M. M., and J. F. Karinen. 1988. Reproductive

success in Tanner (Chionoecetes bairdi) and Dungeness

(Cancer magister) crabs held on oiled sediment.

J. Shellfish Res. 7:109.

Gravid female Tanner (Chionoecetes bairdi) and Dungeness (Cancer

magister) crabs were held on one of several concentrations

(0-8.9 μ1 Cook Inlet crude oil/gm sediment) of oiled sediment

through one complete reproductive cycle. These two species of

commercially important crabs exhibit different behavioral

characteristics while carrying eggs. Brooding Dungeness crab

females consistently bury in the sediments while gravid Tanner

crab females rarely bury. Because of differences in behavior, we

predicted dissimilar responses to oiled sediments.
22

Dungeness crabs held on all doses of oiled sediments

produced significantly fewer numbers of larvae than did control

crabs. Larvae from crabs in high-dose tanks survived

significantly shorter periods than did larvae from the control,

low- and mid-dose tanks. Eggs from crabs in high-dose tanks had

significantly elevated levels of aromatic and aliphatic

hydrocarbons, compared with eggs from control crabs.

Production of larvae and viability of larvae of Tanner crabs

held on oiled sediments showed no differences from that of

control crabs. Likewise, there was no significant uptake of

hydrocarbons in eggs over control levels.

Intimate contact of gravid Dungeness crabs with

sediment-absorbed oil and oil present in interstitial waters in

polluted habitat could reduce reproductive success in this

species.

3 Benville, P. E., and S. Korn. 1977. The acute toxicity of

six monocyclic aromatic crude oil components to striped bass

(Morone saxatilis) and bay shrimp (Crago franciscorum).

Calif. Fish Game 63:204-209.

The acute toxicities of benzene, toluene, ethylbenzene, p-xylene,

m-xylene, and o-xylene were determined for striped bass and bay

shrimp by static bioassay. The 96-h LC50, ranged from 2.0 to

11 μ1/1 (ppm) for striped bass and from 0.49 to 20 μ1/1 (ppm) for

bay shrimp. Solubilities of these aromatics were determined by

gas chromatography in 16°C (61°F) seawater with a salinity of 25%

as part of our procedure for dosing the animals. The

solubilities were 1400, 330, 180, 180, 210, and 230 μ1/1 (ppm),

respectively, which is high enough to be lethal to striped bass

and bay shrimp. The toxic effect of the aromatics was more

latent in shrimp than in fish as demonstrated by the difference

in the 24- and 96-h tests.

4 Brodersen, C. 1987. Rapid narcosis and delayed mortality

in larvae of king crabs and kelp shrimp exposed to the

water-soluble fraction of crude oil. Mar. Environ. Res.

22:233-239.

Sensitivity of larval Paralithodes camtschatica and Eualus

suckleyi exposed 20 min-96 h to the water-soluble fraction (WSF)

of crude oil was determined. Swimming cessation in half the

animals occurred within 20 min exposure to 2 ppm WSF for shrimp

larvae and only 0.5 ppm for crab larvae. Half the larvae of both

species died after 6 h exposure to 8 ppm or 24 h exposed to

2-4 ppm, but deaths did not occur until several days after

exposure ended. In the field, most non-swimming larvae would

probably die as well.
23

5 Brodersen, C. C., S. D. Rice, J. W. Short, T. A.

Mecklenburg, and J. F. Karinen. 1977. Sensitivity of

larval and adult Alaskan shrimp and crabs to acute exposures

of the water-soluble fraction of Cook Inlet crude oil. In

1977 Oil Spill Conference (Prevention, Behavior, Control,

Cleanup), p. 575-578. American Petroleum Institute,

Washington, DC.

The sensitivity of adult and larval Alaskan shrimp and crabs to

the water-soluble fraction (WSF) of Cook Inlet crude oil was

measured by tests using 96-h static bioassays at the water

temperatures that these animals normally encounter. Larval

crustaceans were found to die more slowly than adults, making it

necessary to measure sensitivity in terms of concentrations

causing moribundity (death imminent) instead of in terms of

concentrations causing death during exposure. Cessation of

motion and reaction indicated moribundity in adults, and

cessation of swimming indicated moribundity in larvae exposed for

96 h. Ninety-six hour LC50' s for moribundity for stage I larvae

ranged from 0.95 to 1.8 ppm depending on species, while 96-h

LC50' s for adults ranged from 1.9 to 4.2 ppm oil. Sensitivities

for stage I-VI larvae of coonstripe shrimp ranged between

0.24 ppm to 1.9 ppm.

Larvae were more sensitive to oil than adults. The

sensitivity of larvae depended on species and developmental

stage. Larvae are probably more vulnerable than adult to oil

exposure because of greater sensitivity to oil and greater

susceptibility to predation. Cold-water species may be

particularly vulnerable because of increased time spent as

developing larvae.

6 Carls, M. G. 1987. Effects of dietary and water-borne oil

exposure on larval Pacific herring (Clupea harengus

pallasi). Mar. Environ. Res. 22:253-270.

Pacific herring (Clupea harengus pallasi) larvae were exposed

either directly to the water-soluble fraction (WSF) of crude oil

or indirectly via oil-contaminated prey (OCP) (Artemia salina)

for ? ?28 days, to determine the relative effects of diet and

water as routes of contamination. Larvae were affected rapidly

by 0.9 ppm WSF, which caused high larval mortality, reduced

swimming ability and rapid reduction in feeding rates. Larval

length was significantly reduced by 0.7 ppm WSF in 7 days and by

0.3 ppm WSF within 14 days.

reductions in larval weights.

The WSF exposure also caused similar

Highly contaminated prey (6 ppm

prey exposure) caused significant mortality, but surviving larvae

appeared robust. The OCP did not affect swimming, feeding, or

growth. Exposure of larvae was not significantly extended by

OCP, which rapidly depurated WSF in clean water (98% in 1 day).

Therefore, OCP is probably not an important source of low

molecular weight petroleum contamination to larval fish in the

marine environment.
24

7 Carls, M. G. 1978. Some toxic and behavioral effects of

number 2 fuel oil on the eggs and larvae of Scomber scombrus

L. and Gadus morhua L. M.S. Thesis, Dalhousie Univ.,

Halifax, Nova Scotia, 96 p.

Acute toxic and behavioral effects of No. 2 fuel oil on mackerel

and cod eggs and larvae were studied. Abbreviated ranges of

compounds obtained from the fuel oil were also tested. Maximum

exposure to the oil was 139+30 h for mackerel and 278+60 h for

cod. Oil loss (determined fluorimetrically) was exponential or

nearly exponential. Mean exposure concentrations ranged from

0.00 to 16.22 ppm oil.

Based on mortality, effective oil concentrations (EC's) for

mackerel larvae were 0.28 after 96 h of exposure, but dropped to

0.10 ppm after 480 h (predicted). EC's for cod larvae at 5.9°C

were not statistically different from those for the mackerel

larvae (0.36 ppm, 96 h; 0.15 ppm, 480 h) but were significantly

less for cod larvae at 2°C (0.18 ppm 96 h; 0.07 ppm, 480 h).

mortality due to the oil was not significant, but developing

Egg

embryos were significantly damaged as shown by post-hatching

mortality (mackerel).

Effective concentrations were less when calculated from

behavioral data, but eventually converged with those calculated

from LT50' s (mortality data). The relationship could be

demonstrated by calculating the ratio ECET50/ECLT50: at 96, 480, and

1000 h these ratios were 0.33, 0.53, and 0.64 for No. 2 fuel oil

(cod larvae).

Although increased measurement sensitivity might account for

some of the observed sensitivity, this study suggests mackerel

and cod larvae are more sensitive to oil than previous reports

have indicated.

An interpretative model describing the relationship between

the rates of loss and rates of effect of individual oil

components was formulated.

8 Carls, M. G., and S. Korn. 1985. Sensitivity of Arctic

marine amphipods and fish to petroleum hydrocarbons. Can.

Tech. Rep. Fish. Aquat. Sci. 1368:11-26.

We determined the sensitivities of six circumpolar benthic

species (amphipods Anonyx nugax, Boeckosimus nanseni, and

Gammaracanthus loricatus; a mysid, Mysis relicta; Arctic cod

Boreogadus saida; and a sculpin, Oncocottus hexacornis) to

water-soluble fractions (WSF) of Cook Inlet crude oil and

naphthalene in separate tests. Exposures were flow-through and

lasted 40 days. Median lethal concentrations (LC50's) of WSF

ranged from 1.6 to 3.8 ppm total aromatics. Naphthalene assays

were conducted at several temperatures (1.5 to 9.6°C) to study

temperature effects on sensitivity to hydrocarbons. LC50's

ranged from 1.35 to 3.35 ppm. General relationships between

exposure temperatures and LC50's were not found. In the absence

of toxicants, upper lethal temperatures for the crustaceans were

surprisingly high: 17-24°C, suggesting the assay temperatures in

themselves were not particularly stressful.
25

We compared the sensitivities of these Arctic marine species

to the sensitivities of temperate species previously tested at

the Auke Bay Laboratory using the same flow-through procedures

and toxicants. We concluded that Arctic species are about equal

in sensitivity to temperate species. However, their habitat is

more vulnerable to the effects of petroleum hydrocarbon pollution

than temperate habitats because colder temperatures lead to

slower losses of hydrocarbons from volatilization and

biodegradation, and oil entrapment under sea ice can result in

very lengthy exposures.

9 Carls, M. G., and S. D. Rice. 1990. Abnormal development

and growth reductions of pollock Theragra chalcogramma

embryos exposed to water-soluble fractions of oil. Fish.

Bull., U.S. 88:29-37.

Exposure of developing pollock Theragra chalcogramma embryos to

static water-soluble fractions (WSF) of Cook Inlet crude oil in

seawater slowed initial development, produced shorter larvae, and

caused morphological abnormalities including membranous vesicles;

body curvatures; deformations of yolk, eye, brain, jaw,

intestine, and pericardial sac; absence of lower jaw; fin

erosion; yolksac bloating; and light pigmentation. These

abnormalities were retained after hatch, and in many cases became

more pronounced as developing structures failed to form properly.

The median concentration of WSF that caused abnormalities (AB50)

was 2.1+0.1 ppm. Exposure during embryonic development reduced

prehatch survival by a maximum of 26%, and caused high mortality

after hatch. The median lethal concentration (1.8+0.6 ppm) was

not significantly different than the AB50. Although exposed

pollock embryos generally survived to hatching, larvae were

malformed, smaller, and had poorer survival potential than

controls.

10 Carls, M. G., and S. D. Rice. 1988. Sensitivity

differences between eggs and larvae of walleye pollock

(Theragra chalcogramma) to hydrocarbons. Mar. Environ. Res.

26:285-297.

Exposure of embryonic and larval walleye pollock (Theragra

chalcogramma) to the water-soluble fractions (WSF) of Cook Inlet

crude oil during embryonic development caused mortality and a

variety of morphological abnormalities. Median WSF

concentrations (1.8 ppm) which caused mortality after hatch did

not differ significantly from those causing morphological

abnormalities (1.6 ppm).

develop abnormalities,

Pollock larvae exposed to WSF did not

but swimming and survival were

significantly affected within 4 h of exposure. The LC50 for

larvae dropped from 1.9 ppm on day 4 to 0.9 ppm on day 10.

The tissue of developing pollock larvae bioaccumulated

significantly more dissolved toluene and naphthalene than those

of embryos. Consequently, when exposed to equal concentrations

of WSF, larval tissues are exposed to much higher hydrocarbon
26

concentrations than embryonic tissues. On the basis of internal

tissue concentrations, larvae were less sensitive than eggs, but

the opposite conclusion is reached if external (WSF)

concentrations are used for estimation of sensitivity.

11 Carls, M. G., and S. D. Rice. 1984. Toxic contributions of

specific drilling mud components to larval shrimp and crabs.

Mar. Environ. Res. 12:45-62.

Toxicities of six drilling muds, mud fractions (supernatants and

suspensions) and common mud components (barite, bentonite, and

ferrochrome lignosulfonate) were tested with stage I larvae of

six species of shrimp and crab. The drilling muds we tested were

not very toxic to these larvae: LC50's for supernatants ranged

from 0.6 to 82% (vol/vol). Shrimp larvae were slightly more

sensitive than crab larvae.

Drilling muds were not rapidly toxic, in contrast to

toxicants such as the water-soluble fractions of oil.

Suspensions were more toxic than supernatants and toxicity was

greatest when particulates remained suspended: for example, the

toxicity of used Cook Inlet mud was primarily due to suspended

solids (88%) rather than chemical toxicity. Ferrochrome

lignosulfonate was relatively toxic alone, but accounted for only

about 6% of the toxicity of used Cook Inlet mud suspensions.

Contributions of particulates to mud toxicities varied

considerably. Barite and bentonite were not very toxic when

tested alone. The toxicity of one mud was caused by its high

alkalinity.

12 Carls, M. G., and S. D. Rice. 1981. Toxicity of oil-well

drilling muds to Alaskan larval shrimp and crabs. In

Environmental Assessment of the Alaskan Continental Shelf.

Final Reports of Principal Investigators, Vol. 11

Biological Studies, p. 1-35. U.S. Dep. Commer., NOAA.

Available Arctic Environmental Assessment Center, 222 W. 8th

Ave., No. 56, Anchorage, AK 99513.

The toxic nature of drilling muds is very different from the

toxic nature of crude oil. Drilling muds affect larval

crustacean swimming ability and survival slowly, whereas the

effects of oil are very rapid. The water-soluble fractions of,

drilling muds are stable in solution, but oil WSF's are not. The

slow nature of the drilling-mud toxicity suggests the toxic

components are not very active as chemical poisons, but cause

extra larval energy expenditure to maintain homeostasis.

Crustacean larvae are more sensitive to drilling muds than adult

crustaceans and fish by approximately an order of magnitude.

However, drilling muds are not particularly toxic: toxic

concentrations ranged from 4 to 40 ppt. Water-column

concentrations of drilling muds capable of causing toxicity are

probably brief and limited to distances <3 m from the point of

platform discharge.
27

13 Collodi, P. 1983. Effect of petroleum hydrocarbons on

hepatic aryl hydrocarbon hydroxylase activity in coho salmon

(Oncorhynchus kisutch). M.S. Thesis, Univ. Alaska, Juneau,

120 p.

Coho salmon were exposed to the water soluble fraction (WSF) of

Cook Inlet crude oil for a maximum of 30 days. This exposure, in

both freshwater and sea water, resulted in a greater than

three-fold increase in hepatic aryl hydrocarbon hydroxylase

enzyme activity between two and five days of exposure.

Persistence of the induced enzyme activity was dependent on the

length and the concentration of WSF exposure. Handling stress

had no effect on AHH activity, but starvation caused a decrease

in activity.

14 Collodi, P., M. S. Stekoll, and S. D. Rice. 1984. Hepatic

aryl hydrocarbon hydroxylase activities in coho salmon

(Oncorhynchus kisutch) exposed to petroleum hydrocarbons.

Comp. Biochem. Physiol. C Comp. Pharmacol. 79:337-341.

Coho salmon exposed to the water soluble fraction (WSF) of Cook

Inlet crude oil for a maximum of 30 days showed a greater than

three-fold increase in hepatic aryl hydrocarbon hydroxylase

activity. The initial increase in enzyme activity appeared

between 2 and 5 days of exposure and increased as a function of

increased exposure time. Persistence of the induced enzyme

activity was dependent on the length and the concentration of WSF

exposure.

15 Evans, D. R., and S. D. Rice. 1974. Effects of oil on

marine ecosystems: A review for administrators and policy

makers. Fish. Bull., U.S. 72:625-638.

A broad selection of literature on the effects of oil on marine

ecosystems is reviewed. The focus is on studies on crude oil,

and should be a useful reference for administrators and policy

makers involved in decisions concerning petroleum developments

and related activities. The characteristics of crude oil and

factors modifying its impact on the marine environment are

discussed. Most research on the toxicity of oil has dealt with

acute effects and data on long-term impacts at the community

level are inconclusive. It is concluded that chronic low-level

pollution is potentially more damaging to ecosystems than

isolated catastrophic spills.

16 Gharrett, J. A. 1986. Effects of a hydrocarboncontaminated diet on survival, feeding, growth, and molting

of juvenile red king crab, Paralithodes camtschatica

(Tilesius). M.S. Thesis, Univ. Alaska, Juneau, 178 p.

Juvenile red king crab (Paralithodes camtschatica Tilesius) were

fed diets containing mono25 ug oil/g wet Mytilus,

and di-aromatic hydrocarbons of up to

or unoiled food for 6 months.
28

Hydrocarbons were rapidly accumulated in tissues, in a

non-selective manner with respect to individual aromatic

compounds. HP tissue always had higher levels-of aromatics than

muscle, but tissue levels never exceeded those in food.

Aromatics were rapidly eliminated. A combination of low oil

ingestion rate, and active metabolism and/or excretion prevented

crabs from accumulating higher hydrocarbon concentrations than

the food. Because crabs are able to maintain low hydrocarbon

burdens, treatment regimes had no significant affect on survival,

or any parameter of feeding, growth, molting, or morphometrics.

17 Gharrett, J. A., and S. D. Rice. 1987. Influence of

simulated tidal cycles on aromatic hydrocarbon uptake and

elimination by the shore crab Hemigrapsus nudus. Mar. Biol.

95:365-370.

Intertidal crab Hemigrapsus nudus were used in laboratory

experiments to determine the influence of periodic exposure to

air on the toxicity, uptake, and elimination of two aromatic

hydrocarbons, toluene and naphthalene. Three tidal cycles were

simulated: i.e., crabs spent 0, 33, and 66% of the time in air

and the remainder of the time in toxicant or seawater solutions.

Naphthalene was 10 to 20 times more toxic than toluene and was

accumulated faster and to higher concentrations than was toluene.

Exposure to air 66% of the time reduced mortality as well as the

rate at which toluene and naphthalene were accumulated and lost.

Once crabs had accumulated the toxicants, however, elimination

was hindered by exposure to air, even for the somewhat volatile

compound toluene.

18 Gharrett, J. A., S. D. Rice, and M. S. Stekoll. 1985.

Influence of an oil-contaminated diet on feeding rates,

growth, and molting success of juvenile red king crabs,

Paralithodes camtschatica. In Proceedings of the

International King Crab Symposium, January 1985, Anchorage,

Alaska, p. 371-375. Univ. Alaska, Fairbanks. Alaska Sea

Grant Coll. Program, Alaska Sea Grant Rep. 85-12.

Feeding rates and molting success are sensitive indicators of the

general condition of crabs. We concluded that daily consumption

of oil-contaminated food was not detrimental to juvenile red king

crabs over a 6-mo period, during which rapid growth and molting

occurred. Aromatic hydrocarbon uptake was detected in the

samples of the digestive gland taken early in the exposure.

Accumulation of high concentrations in crab tissues was not

observed. However, the long-term effects of exposure to oil

contaminated food or sediment in the environment are unknown, and

the small amounts of uptake and the trend of decreased feeding

rates are cause for concern.

19 Karinen, J. F. 1988. Sublethal effects of petroleum on

biota. In D. G. Shaw and M. J. Hameedi (editors),
29

Environmental Studies in Port Valdez, Alaska: A Basis for

Management, p. 293-328. Springer-Verlag, Heidelberg.

Aromatic hydrocarbons in sediments, water, and biota of Port

Valdez near the ballast diffuser are similar, in composition and

concentration, to other oil impacted environments, and sublethal

effects of biota of Port Valdez are probably detectable at this

time.

20 Karinen, J. F. 1985. Occurrence of juvenile king crabs,

Paralithodes camtschatica (Tilesius), in Auke Bay, Alaska,

on sediments with relatively high concentrations of aromatic

hydrocarbons. In Proceedings of the International King

Crab Symposium, January 1985, Anchorage, Alaska, p. 377-387.

Univ. Alaska, Fairbanks. Alaska Sea Grant Coll. Program,

Alaska Sea Grant Rep. 85-12.

In the northeastern corner of Auke Bay, Alaska, juvenile king

crab gather in pods and under slate rocks; it is also the

location of two marinas and two docks. Marina activity may

increase concentrations of hydrocarbons, metals, and synthetic

organic compounds in sediments and organisms, which may affect

the occurrence, health, and reproduction of these crabs; The

purpose of this study was to examine the relationship of marina

sediments and king crab populations in northeastern Auke Bay.

Observations on the occurrence and abundance of crabs in

Auke Bay had been made by divers over a 12-year period.

Determination of hydrocarbons in sediments and in the pink

scallop Chlamys spp., a prey item for king crabs, from the

northeastern corner of Auke Bay had been carried out in 1982 and

1983 as part of the preparation for an environmental-impact

statement on marina expansion. Growth of juvenile red king crabs

from the study area and from adjacent areas of Auke Bay were

estimated and compared.

Year-to-year abundance of juvenile king crabs in the study

area appeared to be constant, though abundance of adults had

declined with time. The marina area showed a relatively high

concentration of aromatic hydrocarbons in sediments, probably

primarily from the marinas. Growth information from other

studies indicated reduced growth rates for juvenile king crabs in

the study area when compared with crabs in other areas.

Juvenile king crabs may continue to appear in the study area

because circulation patterns carry larvae to the northeastern

corner. Lower growth rates in the study area may be due to oil

exposure.

1

21 Karinen, J. F. 1980. Petroleum in the deep sea

environment: Potential for damage to biota. Environ. Int.

3:135-144.

1This summary was written specifically for this Technical Memorandum by

Nancy Barr, ABL.
30

Information on the fate, persistence and biological impact of

petroleum hydrocarbons in shallow marine environments, coupled

with recent data on hydrocarbons in offshore sediments and the

biology of deep sea organisms, have provided new perspectives on

the potential impact of oil on the deep sea environment. A

review of literature on petroleum hydrocarbons in deep sea

sediments, mechanisms for transport of petroleum to the deep sea

floor, interaction of petroleum hydrocarbons and particulate

matter, and the physiology and metabolism of deep sea fish and

crustaceans has resulted in the following conclusions:

1. Hydrocarbons of apparent anthropogenic origin are

accumulating in bottom sediments of coastal margins and in deeper

offshore waters at unknown rates.

2. Several mechanisms exist for their rapid transport of

petroleum hydrocarbons to the deep sea floor.

3. Petroleum hydrocarbons are intimately associated with

particulate matter in the sea, behaving much the same as natural

biogenic material and can potentially modify or interrupt natural

processes.

4. The unique physiology of deep water life forms increases

potential adverse impacts of petroleum hydrocarbons.

5. There is a need to determine trends of temporal and

spatial deposition of hydrocarbons in deep sea sediments and

evaluate the biological impact of this introduction of xenobiotic

compounds on the largest environment on earth.

22 Karinen, J. F. 1977. Assessing oil impacts with laboratory

data applications, limitations, and needs. In B. Melteff

(editor), Oil and Aquatic Ecosystems, Tanker Safety and Oil

Pollution Liability. Proceedings of the Cordova Fisheries

Institute, April 1-3, 1977, Cordova, Alaska, p. 99-110.

Univ. Alaska, Fairbanks, Alaska Sea Grant Rep. 77-8.

This paper outlines some of the problems associated with the

utilization of laboratory data to assess the impact of oil in the

environment and some of the considerations which we should make

in trying to apply the toxicity and sublethal effects data. In

applying these results-to the environment to determine what the

actual effects of oil exposure will be, there are both biological

as well as chemical considerations which we must make. This

paper deals more with the chemical aspects rather than the

biological aspects and is limited to a consideration of the

behavior of oil in water and some chemical factors influencing

the application of laboratory effects data toward assessing oil

impacts.

23 Karinen, J. F., and S. D. Rice. 1974. Effects of Prudhoe

Bay crude oil on molting Tanner crabs, Chionoecetes bairdi.

Mar. Fish. Rev. 36(7):31-37.

Premolt and postmolt juvenile male Tanner crabs, Chionoecetes

bairdi, from Alaska waters were exposed to Prudhoe Bay crude oil

in static bioassays in the laboratory. Crabs in both stages were
31

similarly susceptible to crude oil; the estimated 48-h Tlm

(median tolerance limits) values were 0.56 ml oil/liter. Molting

success decreased with increasing exposure of crabs to oil, and

newly molted crabs autotomized limbs during exposure to oil.

Relating the results of our study to the known behavior of crabs

and the documented behavior of oil spills in the ocean suggests

that oil spilled in Alaska waters would harm the Tanner crab

resources. The impact on all crab resources of chronic low-level

oil pollution from the ballast water discharged into Prince

William Sound is unknown. This study further illustrates our

present state of ignorance concerning the biological effects of

oil in the marine environment.

24 Karinen, J. F., S. D. Rice, and M. M. Babcock. 1990.

Reproductive success in Dungeness crab (Cancer magister)

during long-term exposures to oil-contaminated sediments.

In Outer Continental Shelf Assessment Program, Final Reports

of Principal Investigators, Vol. 67, p. 435-461. U.S. Dep.

Commer., NOAA. Available Arctic Environmental Assessment

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

Dungeness crab (Cancer magister) habitat in the nearshore waters

of Alaska may be at risk from oil pollution. Transport of crude

oil and expanded exploratory drilling in Alaskan waters increase

the likelihood of accidental contamination of coastal sediments,

and oil absorbed by the sediments is likely to persist for years

in subarctic and Arctic waters. In contrast to other

commercially important Alaskan crabs, Dungeness crabs completely

bury themselves, particularly when spawning and incubating their

eggs. Therefore, their eggs are in direct contact with sediments

for extended periods and are vulnerable to any xenobiotics

contaminating the habitat.

In Phase 1 of this study, we exposed adult female crabs to

three dose levels of oiled sediments for one complete

reproductive cycle (10 months) to determine effects on survival,

uptake of hydrocarbons, hatching success, and viability of

larvae. Crabs were exposed to 0, 1.2, 3.7, or 8.6 μ1 Cook Inlet

Crude oil per gram of sediment. Hatching began in mid-April, and

larvae were captured in a trap attached to the outlet of each

tank. At three different times during the hatching period, we

tested larvae, to determine viability.

Dosed crabs produced significantly (0.025<P<0.05) lower

numbers of larvae than control crabs. Control crabs produced a

mean of 368,700 larvae/crab, and the low, mid, and high doses

produced 225,500, 303,900, and 268,100 larvae/crab, respectively.

Larvae from the high-dose tanks survived for significantly

(P<0.005) shorter periods (3.1 days) than larvae from the control

tanks and low- and mid-dose tanks (5.3 days). Eggs from crabs in

the high-dose tanks had significantly elevated levels of aromatic

and aliphatic hydrocarbons.

In Phase 2, we studied the effects of oiled-sediment

exposures on mating and molting. The experimental tanks

contained old (held over from Phase 1) control, old mid-dose, old

high-dose, fresh high-dose, and new control sediments. Female
32

crabs from Phase 1 exposures and previously unexposed male crabs

were used in the old sediment tanks. Previously unexposed male

and female crabs were placed on new sediments.

Clasping behavior and successful molting were low in all

old-dose experimental conditions,

in the new high-dose tanks,

and completely absent in crabs

the new control tank.

but occurred in 75% of the pairs in

The low rate of clasping and molting in

all old, 14-month exposures was probably caused by a combination

of confinement, diet, and oil. The briefer, 4-month exposure to

freshly oiled sediment completely inhibited clasping behavior and

molting.

Fifty-five percent of the female crabs that had not molted

subsequently spawned and established fertile clutches. At the

termination of the exposures, examination revealed the presence

of fresh sperm in the spermathecas of over 90% of the females,

indicating these females had copulated in the hard-shelled

condition. We believe that spawning without molting is an

adaptive response for using energy reserves that may have been

depleted by a combination of stressors.

Exposure to oiled sediments results in lower reproductive

activity in Dungeness crabs,

robust.

and the larvae produced are not as

Therefore, over a period of time, the presence of oil in

the habitat substrate may lower population densities.

25 Korn, S., N. Hirsch, and J. W. Struhsaker. 1977. The

uptake, distribution, and depuration of 14C benzene and 14C

toluene in Pacific herring, Clupea harengus pallasi. Fish.

Bull., U.S. 75:633-636.

Benzene and toluene are water-soluble and toxic monoaromatic

components of petroleum. In this study, a comparison of the

uptake, distribution, and depuration of 14C benzene and 14C

toluene in Pacific herring, Clupea harengus pallasi, was

undertaken to determine which of these may pose the greatest

problem.

Adult herring in non-spawning condition were exposed to

100 ppb 14C benzene or 100 ppb 14C toluene.

and lasted 48 h. In all tissues,

Exposures were static

toluene accumulated to higher

levels than did benzene, despite the faster loss of toluene from

test solutions. Highest accumulations were found in the gall

bladder (34 nl/g toluene and 3.1 nl/g benzene). This was

340 times (toluene) and 31 times (benzene) initial concentrations

in the test solutions. Other tissues exhibited lower levels with

immature gonads showing the lowest. Accumulation was rapid with

toluene peaking before 24 h and benzene within 1-2 d.

of tissues except gallbladder,

Depuration

also rapid--within

intestine and pyloric caeca, was

Toluene

1-2 d after termination of exposures.

is more toxic and probably has greater sublethal

effects than benzene.

26 Korn, S., N. Hirsch, and J. W. Struhsaker. 1976.

distribution, and depuration of

Uptake, 14C-benzene in northern
33

anchovy, Engraulis mordax, and striped bass, Morone

saxatilis. Fish.,Bull., U.S. 74:545-551.

The uptake, distribution, and depuration of water-soluble,

monocyclic hydrocarbon contained in petroleum and refined

products was studied in two species of marine fish. Mature

northern anchovy, Engraulis mordax, and juvenile striped bass,

Morone saxatilis, were exposed to sublethal concentrations of 14C-benzene for 48 h. Residues in tissues exhibiting a high

lipid content or representing apparent major metabolic sites were

measured during the exposure and afterwards when the fish were

transferred to clean seawater. Fish exhibited a rapid uptake

over a wide range of benzene concentrations in the water column.

Accumulation in anchovy was considerably greater than in striped

bass. Results indicate that the pathway of hydrocarbons through

the liver, gallbladder, intestines, and colon is a major

depuration route. Residues were depurated rapidly after

cessation of exposure; in striped bass tissues most residues were

undetectable by 7 days.

27 Korn, S., D. A. Moles, and S. D. Rice. 1979. Effects of

temperature on the median tolerance limit of pink salmon and

shrimp exposed to toluene, naphthalene, and Cook Inlet crude

oil. Bull. Environ. Contam. Toxicol. 21:521-525.

We measured the effects of temperature on the TLm's of pink

salmon (Oncorhynchus gorbuscha) fry and shrimp (Eualus spp. and

Pandalus goniurus) exposed to toluene, naphthalene, and the

water-soluble fraction (WSF) of Cook Inlet crude oil. Static

exposures were used, in which the initial concentration of the

toxicant declines with time, a situation that would also occur

with an oil spill. Both the persistence of oil and the

physiology of animals are affected by temperature, and the

measured TLm's are the net result of both variables operating

simultaneously.

28 Korn, S., and S. Rice. 1981. Sensitivity to, and

accumulation and depuration of, aromatic petroleum

components by early life stages of coho salmon (Oncorhynchus

kisutch). Rapp. P.-V. Reun. Cons. Int. Explor. Mer

178:87-22.

Coho salmon eggs, alevins, and fry were exposed to toluene,

naphthalene, and 2-methyl-naphthalene in a series of short-term

toxicity and hydrocarbon-uptake studies to determine whether

acute toxicity is related to uptake-depuration patterns. Uptake

studies used radiolabeled compounds and radiometric analyses.

The time to reach maximum tissue concentrations of these

hydrocarbons was determined from long-term exposures.

Sensitivity to the aromatics increased from egg to fry with

the greatest increase in sensitivity between the egg and early

alevin. The rates of uptake and depuration of the aromatic
34

hydrocarbons also increased during the development from egg to

fry. Eggs had the slowest rates of uptake and depuration.

Eggs required 10 days to accumulate stable tissue

concentrations of toluene and naphthalene; alevins required 36 h

(both toxicants); fry, 3 h (toluene) and 10 h (naphthalene). The

rate of uptake and toxicity was higher with increased ring size

and increased substitution (2-methyl-naphthalene > naphthalene >

toluene).

Eggs were more tolerant than alevins and fry to short-term

exposures of aromatic hydrocarbons probably because the chorion

prevented rapid uptake. The amount of yolk also influenced

sensitivity because aromatic hydrocarbons were selectively

partitioned into the yolk, thus reducing availability of the

hydrocarbons to the embryo and resulting in lower toxicity.

Although eggs take up hydrocarbons at a slow rate, they may

accumulate lethal levels of hydrocarbons during long-term

exposures and, therefore, would be more sensitive to hydrocarbons

than indicated by short-term experiments.

29 Korn, S., S. D. Rice, D. L. Cheatham, and D. W. Brown.

1985. Contribution of phenol and p-cresol to the toxicity

of crude oil to pink salmon (Oncorhynchus gorbuscha) fry and

kelp shrimp (Eualus suckleyi). In F. J. Vernberg, F. P.

Thurberg, A. Calabrese, and W. B. Vernberg (editors), Marine

Pollution and Physiology: Recent Advances, p. 447-458.

Univ. South Carolina Press, Columbia, SC.

Concentrations of phenolic compounds in the WSFs of Cook Inlet

crude oil, Prudhoe Bay crude oil, and No. 2 fuel oil ranged from

0.040 to 0.088 ppm. Concentrations of phenol that killed 50% of

the animals in 96 h (the 96-h LC50) was 3.7 ppm for pink salmon

and 10.3 ppm for kelp shrimp, LC50s for p-cresol were 3.4 ppm for

pink salmon and 7.4 ppm for kelp shrimp. In 24 h, salmon and

shrimp accumulated 11-22 times the initial exposure concentration

of phenol and p-cresol. Both compounds were completely

eliminated from pink salmon by day 7 or less. Shrimp depurated

the compounds slower than pink salmon.

Phenol and p-cresol contributed less to the toxicity of the

WSF than toluene and naphthalene. Although toluene is one-half

as toxic as phenol, its concentration in the WSF is 50 times

higher than the concentration of phenol. Naphthalene, which is

3-10 times more toxic than phenol, had concentrations in the WSF

that were 2-7 times higher than the concentration of phenol.

Furthermore, the phenols accumulated slowly and are eliminated

rapidly. We, therefore, conclude phenols do not contribute

substantially to the toxicity of the WSF of crude oil.

30 Korn, S., J. W. Struhsaker, and P. Benville. 1976. Effects

of benzene on growth, fat content, and caloric content of

striped bass, Morone saxatilis. Fish. Bull., U.S.

74:694-698.
35

Juvenile striped bass were exposed to constant concentrations of

benzene to determine whether sublethal levels would inhibit

efficient energy utilization by the fish as measured by growth

(wet weight, dry weight), fat content, and caloric content.

Thirty-five fish were maintained up to 4 weeks in each of nine

aquariums; three aquariums at 3.5 μl/liter benzene, three at

6 μl/liter benzene, and three at 0 μl/liter benzene as controls.

Fish were monitored daily and seven fish were sampled from each

aquarium at 0, 7, 14, 21, and 28 days, and weighed (wet and dry);

three fish from each group were subjected to caloric analysis,

and four to fat analysis.

Initial exposure to benzene caused hyperactivity among fish

at the high benzene level and moderately elevated activity levels

at the lower level of exposure. Fish exposed at the high

concentration attempted to feed, but could not locate food

pellets; fish at lower concentrations consumed 50% of the food

offered. By the end of the study, high-dose-level fish consumed

50% of their ration and low-level fish fed normally. Wet

weights, dry weights, and fat content decreased with increased

benzene exposure.1

31 Lauren, D. J., and S. Rice. 1985. Significance of active

and passive depuration in the clearance of naphthalene from

the tissues of Hemigrapsus nudus (Crustacea: Decapoda).

Mar. Biol. 88:135-142.

Adult male crabs Hemigrapsus nudus were exposed statically in

seawater to 14C-naphthalene for 12 h followed by up to 156 h of

depuration. Uptake was rapid, and by 12 h the digestive gland

had accumulated 105 times the water 14C, while other tissues had

accumulated less than 15 times. Depuration was rapid at first,

but slowed by 12 h post-exposure. After 156 h of depuration, the

parent naphthalene that remained was found mainly in the

digestive gland and muscle. However, the highest percentages of

naphthalene metabolites were found in the gills, muscle, and

hemolymph. Metabolites were retained to a greater degree than

the parent compound. Because no significant difference in

depuration rate was found between control individuals injected

with 14C-naphthalene and those with their nephropores and anus

blocked, it was concluded that the gills are the major route of

naphthalene elimination. Thin layer chromatography (TLC) of

extracts of depurated 14C-naphthalene indicated that <10% of the

total naphthalene depurated was composed of metabolites.

In-vitro mixed function oxygenase (MFO) activity was assayed on

15,000 g tissue homogenates, using diphenyloxazole as the

terminal electron acceptor. Specific activity of the gills was

2 to 4 times that of the antennal glands, and no activity was

detected in either muscle, digestive glands, or cardiac and

pyloric stomachs. MFO activities were very low compared to fish

or mammals. It was concluded that metabolism of naphthalene

plays a minor role in the reduction of naphthalene levels, with

the major role being played by simple diffusion of unmetabolized

napthalene across a concentration gradient. This occurs across

the tissue with the largest body-to-water surface area--the gill.
36

32 Mecklenburg, T. A., S. D. Rice, and J. F. Karinen. 1977.

Molting and survival of king crab (Paralithodes

camtschatica) and coonstripe shrimp (Pandalus hypsinotus)

larvae exposed to Cook Inlet crude oil water-soluble

fraction. In D. A. Wolfe (editor), Fate and Effects of

Petroleum Hydrocarbons in Marine Ecosystems and Organisms,

p. 221-228. Pergamon Press, New York.

Larvae of coonstripe shrimp and king crab were exposed to

solutions of the water-soluble fraction (WSF) of Cook Inlet crude

oil in a series of bioassays on intermolt stages I and II and the

molt period from stage I to stage II. Molting larvae were more

sensitive than intermolt larvae to the WSF, and molting

coonstripe shrimp larvae were more sensitive than molting king

crab larvae. When molting larvae were exposed to high

concentrations of the WSF (1.15-1.87 ppm total hydrocarbons) for

as little as 6 h, molting success was reduced by 10-30% and some

deaths occurred. When larvae were exposed to these high

concentrations for 24 h or longer, molting declined 90-100% and

the larvae usually died. The lowest concentrations tested

(0.15-0.55 ppm total hydrocarbons) did not inhibit molting at any

length of exposure, but many larvae died after molting.

lethal concentrations (LC50' s) based on 144 h of observation for

molting coonstripe shrimp and 120 h for molting king crab were

much lower than the 96-h LC50's, showing that the standard 96-h

LC50 is not always sufficient for determining acute oil toxicity.

Although our LC50 's for intermolt larvae are higher than levels

of petroleum hydrocarbons reported for chronic and spill

situations, some of our LC50 's for molting larvae exposed 24 h

and longer are similar to or below these environmental levels.

Comparisons of sensitivity to oil between different crustacean

species or life stages should be based on animals tested in the

same stage of the cycle, such as intermolt.

33 Merrell, T. R. 1988. Fisheries resources. In D.G. Shaw

and M. J. Hameedi (editors), Environmental Studies in Port

Valdez, Alaska: A Basis for Management, p. 203-224.

Springer-Verlag, New York.

Pink, chum, and coho salmon and Pacific herring are the most

important fisheries resources in Port Valdez from both commercial

and recreational standpoints. Early life stages of these species

are the most vulnerable to oil pollution: pink and chum salmon

eggs in intertidal spawning streambeds, and fry during early

marine life; coho salmon smolts during their seaward migration

through Port Valdez; and Pacific herring eggs and larvae during

their first few weeks of life before they have dispersed away

from spawning sites.

The new Solomon Gulch Hatchery is expected to reach full

production of salmon by 1989, when more than 6 million pink,

324,000 chum, and 60,000 coho salmon will return each year as

adults. Most of the pink and chum salmon will be caught by

common property commercial net fisheries, and most of the coho
37

salmon by sport fishermen. Unless a catastrophic oil spill

occurs in Port Valdez, or concentrations of toxic components from

the treated tanker ballast effluent become substantially higher

in Port Valdez than at present, it is unlikely that oil pollution

will have significant adverse impacts on hatchery or natural

stocks of salmon, herring, or other fishery resources.

The possibility of eventual subtle indirect effects of

hydrocarbons on salmon and herring, and their habitat, cannot be

ruled out, however. Hydrocarbon concentrations from the ballast

effluent are gradually increasing in sediment and water and could

eventually have harmful effects, such as altering migratory

behavior of juvenile and adult salmon, or diminishing the

abundance of prey species upon which salmon and herring are

dependent during early sea life.

34 Moles, A. 1980. Sensitivity of parasitized coho salmon fry

to crude oil, toluene, and naphthalene. Trans. Am. Fish.

soc. 109:293-297.

The effect of parasitism by glochidia of Anodonta oregonensis (a

freshwater mussel) on the sensitivity of coho salmon fry,

Oncorhynchus kisutch, to oil was determined by exposing fry with

different levels of parasitism to several concentrations of

either the water-soluble fraction of Prudhoe Bay Crude oil or the

aromatic hydrocarbons toluene and naphthalene. Fry infested with

20-35 glochidia were significantly (P < 0.05) more sensitive to

each of the toxicants than uninfested fish. Sensitivity

increased linearly with increased parasite numbers.

Interpretation and application of results of toxicity tests

should take into account the kinds and intensities of parasitism

found both in the test animals and in the wild populations of

fish.

35 Moles, D. A. 1977. Effect of parasitism by glochidia of

Anodonta oregonensis Lea on the sensitivity of coho salmon

fry, Oncorhynchus kisutch Walbaum, to the water-soluble

fraction of Prudhoe Bay crude oil and the effect of crude

oil on the development of the glochidium. M.S. Thesis,

Univ. Alaska, Juneau, 50 p.

Larvae of the freshwater mussel Anodonta oregonensis Lea are

incubated amongst the branchial lamellae of the mother and when

released, glochidia become protelean parasites on fins of fish

such as coho salmon.

Ninety-six hour static bioassays using the water soluble

fractions (WSF) of Prudhoe Bay crude oil and benzene were

conducted on the adult mussels to determine the sensitivity of

the mussels to the toxicants. Crude oil water-soluble fraction

had no lethal effect and the median tolerance limit (TLm) for

mussels in benzene was 500 ppm.

Exposing egg bearing mussels to the water-soluble fraction

(WSF) of Prudhoe Bay crude oil for 96-h and subsequently holding

the mussels in the laboratory for two months resulted in an
38

inhibition of the development of the glochidia. The eggs of

mussels exposed to oil concentrations of 2.4 ppm total aromatics

had nearly completed development while mussels exposed to

10.94 ppm still carried fertilized eggs.

Coho salmon fry were inoculated with glochidia at three

levels of infestation (10-45, 46-65, and 66-100 parasites per

fish) and exposed to concentrations of Prudhoe Bay WSF ranging

from 0.95 ppm to 11.73 ppm total aromatics for 96 h. TLm's

ranged from 10.38 ppm total aromatics for non-parasitized fish to

2.26 ppm for heavily (66-100) parasitized fish.

The conclusions of the study are: 1) adult freshwater

mussels are highly tolerant to oil pollution, 2) the development

of glochidia incubating in the adult mussel can be inhibited by

exposure to crude oil WSF with the degree of inhibition

increasing with increasing oil concentration and, 3) the greater

the number of glochidia on coho salmon fry, the greater the

sensitivity of the salmon to water-soluble fraction of Prudhoe

Bay crude oil.

36 Moles, A., M. M. Babcock, and S. D. Rice. 1987. Effects of

oil exposure on pink salmon, Oncorhynchus gorbuscha, alevins

in a simulated intertidal environment. Mar. Environ. Res.

21:49-58.

Pink salmon, Oncorhynchus gorbuscha, alevins (5 and 60 days after

hatching) were continuously or intermittently exposed for 30 days

to the water-soluble fraction (WSF) of Cook Inlet crude oil in

fresh water or in a simulated freshwater-seawater cycle. Alevins

exposed to 0.7-2.4 mg/liter WSF in the simulated tidal cycle were

more sensitive to oil, had reduced yolk reserves, and accumulated

more hydrocarbons than did alevins exposed to the same

concentrations in fresh water. Alevins in fresh water were more

sensitive to continuous than to intermittent exposures. In all

exposures, 60-day alevins were more severely affected than were

5-day alevins.

37 Moles, A., S. Bates, S. D. Rice, and S. Korn. 1981.

Reduced growth of coho salmon fry exposed to two petroleum

components, toluene and naphthalene, in fresh water. Trans.

Am. Fish. Soc. 110:430-436.

Coho salmon, Oncorhynchus kisutch, fry were exposed for 40 days

to stable, sublethal concentrations of toluene or naphthalene in

fresh water. All fry were fed equal daily rations of Oregon2

Moist Pellet Formula II. Dry weights, wet weights, and lengths

of fry exposed to the two highest concentrations of each toxicant

for 40 days were significantly less than controls (P < 0.01).

Growth per day, determined from weights and lengths, decreased

linearly with increased concentrations. Fry exposed to

2

Reference to trade names does not imply endorsement by National Marine

Fisheries Service, NOAA:
39

naphthalene had a slower growth rate than fry exposed to

equivalent concentrations (percentage of the 96-h median lethal

concentration of LC50) of toluene. Concentrations 18% of the LC50

of naphthalene and 26% of the LC5o of toluene had no effect on

dry weight, wet weight, or length of exposed fry.

38 Moles, A., and S. D. Rice. 1983. Effects of crude oil and

naphthalene on growth, caloric content, and fat content of

pink salmon juveniles in seawater. Trans. Am. Fish. Soc.

112:205-211.

Juvenile pink salmon Oncorhynchus gorbuscha were exposed for

40 days to stable, sublethal concentrations of naphthalene and

the water-soluble fraction of Cook Inlet crude oil. All fish

were fed equal daily rations of Oregon Moist Pellet Formula II.

Change in length of the fish was not a sensitive measure of

toxicity. Fish exposed for 40 days to concentrations of

toxicants as low as 33% of the 96-h LC50 weighed significantly

less than control fish (P < 0.05). Juveniles exposed to the

water soluble fraction of crude oil had slower growth rates than

those exposed to the same concentration (percentage of the LC50)

of naphthalene. Fish exposed to either naphthalene or the

water-soluble fraction of crude oil had decreased caloric

content; however, fat content of the fish was not affected.

Chronic marine oil pollution at a concentration as low as

0.40 mg/liter total aromatic hydrocarbons could reduce growth of

juvenile pink salmon.

39 Moles, A., S. D. Rice, and S. Andrews. 1985.

Continuous-flow devices for exposing marine organisms to the

water-soluble fraction of crude oil and its components.

Can. Tech. Rep. Fish. Aquat. Sci. 1368:53-61.

The devices produced stable concentrations of aromatic

hydrocarbons that can be used in continuous-flow toxicity tests.

The crude-oil mixing device produces a stable water-soluble

fraction of 2.5 mg/1 total aromatic hydrocarbons for 30-40 days.

The device uses a gentle flow of water to dissolve aromatic

components in a layer of crude oil floating on a 2-m column of

seawater. Because the water does not pick up oil droplets as it

passes through the column,

rather than a dispersion.

a water-soluble fraction is produced

The other device, a syringe pump,

introduces compounds directly into a water stream and produces a

stable solution of monoaromatic hydrocarbons of any desired

concentration or mixture up to the maximum solubility of the

compounds.

40 Moles, A., S. D. Rice, and S. Korn. 1979. Sensitivity of

Alaskan freshwater and anadromous fishes to Prudhoe Bay

crude oil and benzene. Trans. Am. Fish. Soc. 108:408-414.
40

The sensitivity of various species and life stages of Alaskan

freshwater and anadromous fishes to benzene and the water-soluble

fraction of Prudhoe Bay crude oil was determined with 96-h

toxicity tests. Freshwater juveniles of the six salmonid species

tested had similar sensitivities: Median tolerance limits

(TLm's) for crude oil ranged from 2.7 to 4.4 mg/liter; TLm's of

benzene ranged from 11.7 to 14.7 μl/liter. Threespine

sticklebacks and, to a lesser extent, slimy sculpins were more

tolerant than salmonids. Eggs of pink salmon and coho salmon

were quite tolerant to crude oil (TLm > 12 mg/liter) and benzene

(TLm = 339-542 μl/liter). Emergent fry were the most sensitive

freshwater stage (crude oil TLm = 8.0 mg/liter; benzene Tlm =

12.3-17.1 μl/liter). Out-migrant salmonids tested in seawater

were twice as sensitive as out-migrant salmonids tested in fresh

water, apparently because of the additional stress of entering

seawater and the physiological changes associated with this

transition. Seawater sensitivities were 1.1-3.6 mg/liter for

crude oil and 5.5-8.5 μl/liter for benzene.

41 Myren, R. T., and J. J. Pella. 1977. Natural variability

in distribution of an intertidal population of Macoma

balthica subject to potential oil pollution at Port Valdez,

Alaska. Mar. Biol. 41:371-382.

Natural variability in the abundance of an intertidal population

of the lamellibranch Macoma balthica (Linnaeus, 1758) was

measured during 1971 and 1972 in a study area near the proposed

oil storage and tankship loading facility at the southern

terminus of the Trans-Alaska pipeline in Port Valdez, Alaska.

M. balthica were divided for analysis into a large and a small

size category. Small temporal changes in population densities

throughout the entire study area were detected for both size

categories over several of the 7 sampling times of the 2-year

period. Large and persistent differences in density were found

among elevation contour intervals for either size category.

Densities of both size categories were more stable at the higher

elevations of the study site. Large M. balthica were more

homogeneously distributed along a given elevation contour

interval than the small category. Mobility and time available to

redistribute at a horizontal location would explain the more

homogeneous distribution of large M. balthica if competition for

food resources exists.

42 O'Clair, C. E., J. L. Hanson, J. S. MacKinnon, J. A.

Gharrett, N. I. Calvin, T. R. Merrell, Jr. 1978. Baseline/

reconnaissance characterization littoral biota, Gulf of

Alaska and Bering Sea. In Environmental Assessment of the

Alaskan Continental Shelf. Annual Reports of Principal

Investigators for the Year Ending March 1978. Volume 4,

Receptors--Fish, Littoral, Benthos, p. 256-415. U.S. Dep.

Commer., NOAA. Available Arctic Environmental Assessment

Center, 222 W. 8th Ave., No. 56, Anchorage, AK 99513.
41

Distribution and relative abundance patterns of littoral plants

and invertebrates at representative sites in the eastern Gulf of

Alaska are described. Eighteen sites were chosen for study. In

addition we examined, insofar as the data permit, those factors

which are likely to play important roles in structuring

intertidal communities.

Understanding how communities are organized is important for

predicting the effects of oil and gas development on community

composition.

Within the limitations of our data we examine the role of an

important interaction in intertidal communities, competition for

space, especially competition among dominant competitors and

accompanying effects on subdominant. Our data indicates that

total species richness tends to be greater in patches of

intertidal area dominated by Mytilus edulis than in patches

dominated by Fucus distichus, and that the difference is

accounted for by increased species richness of small subdominant

in Mytilus dominated areas. Mytilus does not appear to have a

greater adverse effect on competitively inferior large

subdominant than does Fucus.

43 O'Clair, C. E., and S. D. Rice. 1985. Depression of

feeding and growth rates of the seastar Evasterias

troschelii during long-term exposure to the water-soluble

fraction of crude oil. Mar. Biol. 84:331-340.

To test the effect of petroleum hydrocarbons on predation by the

seastar Evasterias troschelii (Stimpson, 1862) on the mussel

Mytilus edulis (L.), we exposed the predator with the prey to six

concentrations of the water-soluble fraction (WSF) of Cook Inlet

crude oil. During a 28 d exposure in a flow-through system,

seastars were more sensitive to the WSF than mussels: the LC50

for the seastars was 0.82 ppm at Day 19 and, although no mussels

were exposed to WSF for more than 12 d, none died. Daily feeding

rates and growth of seastars were significantly reduced at all

concentrations above 0.12 ppm. At 0.20, 0.28 and 0.72 ppm WSF,

daily feeding rates were, respectively, 53, 37, and 5% of the

control rate; at higher concentrations, the seastars did not

feed. These laboratory results show that E. troschelii is more

sensitive to chronic low levels of the WSF of crude oil than the

prey. Oil pollution could reduce predation on mussels and

increase their dominance in the intertidal zones.

44 Pelto, M. J., R. W. Williamson, and J. Knull. 1978.

Dispersion of drogues:

in Port Valdez, Alaska.

an estimation of pollutant dispersal

Northwest and Alaska Fisheries

Center Processed Report, 10 p. Northwest and Alaska Fish.

Cent., Natl. Mar. Fish. Serv., NOAA, 2725 Montlake Blvd. E.,

Seattle, WA 98112.

Assemblages of drogues in Port Valdez dispersed at rates which

increase with time. The rates of dispersal were found by

computing the variance (D2) of the distances transited by the
42

drogues and fitting the results to D2 - atb. A mean value of 1.8

was determined for b with a range between 0.1 and 2.5.

Rates of change of the variance of drogue distances found

imply that effective diffusivity in the direction of flow was

between 3.2 x 103 and 14.2 x 103 cm

2

/sec. Diffusivity in the

transverse direction was between 0.6 x 103

 and 6.8 x 103 cm

2

/sec.

These values are an order of magnitude less than found by another

type of drogue method in Lake Michigan and establish lower limits

to rates of diffusion in Port Valdez.

45 Rice, S. D. 1985. Effects of oil on fish. In F. R.

Engelhardt (editor), Petroleum Effects in the Arctic

Environment, p. 157-182. Elsevier Applied Science

Publishers, New York.

Literature related to the effects of oil on fish is reviewed with

particular emphasis on implications for spilled oil in the

Arctic. Although toxicity of oil and its components to

individual fish is well documented in laboratory studies, the

effects of oil spills on fish are less understood. In the

laboratory, concentrations of <1 ppm aromatic hydrocarbons have

been found to affect growth in fish. Outside the laboratory,

fish are more likely to be affected by chronic sources of oil

pollution than by a one-time spill with short-term, high

concentrations. Eggs and larvae are the most sensitive life

stages of fish: concentrations of <1 ppm aromatic hydrocarbons

probably affect their development in many species. Although fish

at all life stages are apparently vulnerable to damage from oil

spills, devastation to fishery stocks as a result of spills is

not documented. Local populations have, however, been eliminated

from industrialized bays and estuaries where oil may be a

contributing pollutant.

Little is known about spilled oil in the Arctic, but Arctic

populations may be relatively vulnerable because aromatic

hydrocarbons may persist longer in shallow, cold waters like

those found on the northern continental shelf.1

46 Rice, S. D. 1985. Fisheries and oil: facts and figures.

Alaska Fish and Game 18[17](6):35-37.

For ten years, biologists at the Auke Bay Laboratory, National

Marine Fisheries Service, have studied the effects of oil on

selected marine animals. They have found that the lethal

concentration of Alaskan crude oil is about 1-2 ppm aromatic

compounds for salmon, 1-3 ppm for king crab and shrimp, and over

10 ppm for intertidal animals such as prickleback eels and

mussels after 96-h exposures. For shrimp and crab larvae

concentrations as low as 0.2 ppm are lethal, and salmon smolts in

seawater were found to be twice as sensitive to oil as smolts in

freshwater.

Researchers noted that aquatic animals, especially fish, can

rid themselves of petroleum hydrocarbons when placed in fresh

water. Marine eggs and larvae, and sedentary animals such as
43

clams, may not be able to avoid contaminated areas. Laboratory

studies indicate that crabs living in subtidal areas with

fine sediment substrates may accumulate hydrocarbons in tissues

and egg-bearing success may be affected. Animals like crabs and

salmon may detect oil through their sense of smell and, in the

laboratory, avoid oil after detection.

Researchers concluded that wetlands are probably the habitat

most vulnerable to adverse impacts from oil pollution because

they are shallow, highly productive nursery areas for many

oil-sensitive species and because decontaminating bottom

sediments would be difficult. Rocky shores are less productive

and could be cleansed of oil by wave action, but seasonal

concentrations of birds and marine mammals along these shores

could be vulnerable to oil-spill effects.1

47 Rice, S. D. 1977. A review of oil toxicity studies

conducted at the Auke Bay Laboratory. In B. Melteff

(editor), Oil and Aquatic Ecosystems, Tanker Safety and Oil

Pollution Liability. Proceedings of the Cordova Fisheries

Institute, April 1-3, 1977, Cordova, Alaska, p. 111-113.

Univ. Alaska, Fairbanks, Alaska Sea Grant Rep. 77-8.

Although each of our publications has specific conclusions, the

two most significant general conclusions are:

1. We have generally found crustacean larvae to be the most

sensitive life stage, especially when molting.

2. Alaskan species may be more vulnerable to oil than

species from warmer waters, since lower temperatures cause toxic

aromatic hydrocarbons to persist longer. Temperature effects on

oil toxicity and animal sensitivity are complex and warrant

further study.

48 Rice, S. D. 1974. Toxicity and avoidance tests with

Prudhoe Bay oil and pink salmon fry. In 1973 Proceedings of

the Joint Conference on Prevention and Control of Oil

Spills, March 13-15, 1973, Washington, DC, p. 667-670.

American Petroleum Institute, Washington, DC.

With the potential of oil pollution harming Alaska's marine

resources, experiments were conducted at the National Marine

Fisheries Service, Auke Bay Fisheries Laboratory to determine the

concentrations of Prudhoe Bay crude oil that are acutely toxic to

pink salmon fry in fresh water and seawater and also the

concentrations of this oil that fish avoid. Observed 96-h TLm

values were 88 mg of oil/liter of water in fresh water and

213 mg/liter in seawater in June and 100 mg/liter in seawater in

August. Among fry held in seawater, older fry were more

susceptible to oil toxicity than younger fry and older fry were

also more sensitive in their detection and avoidance of oil;

older fry in seawater avoided oil concentrations as low as 1.6 mg

of oil/liter of water. The avoidance of oil by salmon fry was

quite apparent and suggests that there is potential for oil

pollution to change their migration behavior.
44

49 Rice, S. D., and M. M. Babcock. 1989. Effects of habitat

and environmental variables on red king crabs, and settling

of glaucothoe. In L. E. Jarvela and L. K. Thorsteinson

(editors), Proceedings of the Gulf of Alaska, Cook Inlet,

and North Aleutian Basin Information Update Meeting,

February 7-8, 1989, Anchorage, Alaska, p. 49-53. Available

National Marine Fisheries Service, Auke Bay Laboratory.

Bering Sea red king crab (Paralithodes camtschatica) populations

crashed in the early 1980's. There is considerable speculation

on the causes of the population decline. Understanding of the

population dynamics of king crabs is limited by the lack of

biological knowledge, particularly for the larval and juvenile

life stages. The impacts of environmental and habitat variables

on the early life stages remain largely unknown.

The objectives of this research were to determine the role

of environmental and habitat variables on red king crab eggs,

larvae, and juveniles. Many of these variables influence

year-class strength. Adults were not the focus of the study,

except for ovigerous females that were incubating eggs. Both

laboratory and field studies were conducted at Auke Bay,

Southeast Alaska.

A comprehensive report detailing the results of our research

on red king crabs is scheduled to be finished in spring 1990.

50 Rice, S. D., M. M. Babcock, C. C. Brodersen, M. G. Carls,

J. A. Gharrett, S. Korn, A. Moles, and J. W. Short. 1987.

Lethal and sublethal effects of the water-soluble fraction

of Cook Inlet crude oil on Pacific herring (Clupea harengus

pallasi) reproduction. U.S. Dep. Commer., NOAA Tech. Memo.

NMFS F/NWC-111, 63 p.

Pacific herring (Clupea harengus pallasi) are a valuable

commercial fishery in the northwestern Pacific Ocean, where they

are harvested primarily just before spawning when the edible

ovaries are at maximum ripeness. Our studies examined the

effects of oil on important stages of the herring reproductive

process.

crude oil,

Using the water-soluble fraction (WSF) of Cook Inlet

we studied effects of lethal and sublethal exposures

on prespawn adult Pacific herring, eggs, yolk sac larvae, and

feeding larvae. We also studied the effects of feeding

oil-contaminated prey to herring larvae. The results of our

study are summarized:

1. Prespawn adult herring exposed to WSF had a 2- and

12-day LC50 (the median concentration that killed 50% of the

herring) of 2.3 parts per million (ppm) aromatic hydrocarbons.

2. Adults exposed 12 days to 1.6 ppm produced eggs that had

normal hatching success.

3. Eggs exposed 2 days to 5.3 ppm had normal hatching

success; eggs exposed 12, days had LC50 of 1.5 ppm.

4. Yolk-sac larvae exposed ? ? ?h to 6.1 ppm survived;

yolk-sac larvae exposed from 16 h to 6 days had LC50's of 2.8 to

2.3 ppm.
45

5. Feeding larvae exposed 7 days had a LC50 of 1.8 ppm;

exposed 21 days, 0.36 ppm.

6. Uptake of hydrocarbons in adult herring was rapid in

muscle, liver, testes, and mature and immature ovaries, but

equilibrium was not reached in 10 days of exposure.

7. Muscle tissue generally accumulated the highest levels

of hydrocarbons; immature ovarian tissue accumulated almost two

times the levels found in mature ovarian tissue.

8. In adults, initial depuration was rapid but slowed after

24 h, and 10% of the hydrocarbons were still present after 7 days

in clean water. After 14 days, hydrocarbon levels were not

significantly higher in the control levels.

9. Uptake in larvae was more rapid than in adults and

reached equilibrium within 4 h. Retention was lower in larvae

than adults, and after 24 h, only 2% of the 14C-labeled

naphthalene remained in larval tissues.

10. Larval growth was significantly reduced after 7 days of

exposure to 0.3 ppm, and growth reductions were greater after

longer exposures and higher concentrations.

11. Growth of larvae was not significantly reduced by a diet

of oil contaminated prey.

Feeding larvae of Pacific herring are killed by shorter

exposures and lower concentrations than are the eggs or the

adults. This is therefore the life stage most likely to be

impaired by oil. Even if oil is present at levels too low to

threaten the survival of herring, fisheries could be impacted if

the rapid bioaccumulation and persistent retention of oil

hydrocarbons in the edible ovarian tissues made the herring

unmarketable.

51 Rice, S. D., M. M. Babcock, C. C. Brodersen, J. A. Gharrett,

and S. Korn. 1987. Uptake and depuration of aromatic

hydrocarbons by reproductively ripe Pacific herring and the

subsequent effect of residues on egg hatching and survival.

In W. B. Vernberg, A. Calabrese, F. P. Thurberg, and F. J.

Vernberg (editors), Pollution Physiology of Estuarine

Organisms, p. 139-154. Univ. South Carolina Press,

Columbia, SC.

Adult Pacific herring (Clupea harengus pallasi) were exposed to

the water-soluble fraction (WSF) of Cook Inlet crude oil to

determine uptake and depuration of aromatic hydrocarbons in

tissues (including ovaries) and survival of ova and spawn.

Adults accumulated high concentrations of aromatic hydrocarbons

in tissues (including ovaries), but tissue concentrations did not

reach equilibrium during the 10-d sampling.

rapidly during the first 24 h,

Depuration occurred

but then took 14 d before tissue

concentrations were not significantly different from controls.

Exposure to WSF did not affect the survival of ova and spawn. If

the adults survived, their spawn hatched. In a companion test,

artificially spawned and fertilized eggs from unexposed adults

were exposed to several concentrations of WSF.

tolerant to short-term (2-d),

Herring eggs were

direct exposures to high

concentrations of WSF but had a lower (1.5 ppm) LC50 (median
46

lethal concentration) than did adults (2.2 ppm) for 12-d

exposure. The reproductive process of herring just prior to

spawning did not appear to be as sensitive to oil as did eggs.

52 Rice, S. D. and J. F. Karinen. 1976. Acute and chronic

toxicity, uptake and depuration, and sublethal metabolic

response of Alaska marine organisms to petroleum

hydrocarbons. In Environmental Assessment of the Alaskan

Continental Shelf. Annual Reports of Principal

Investigators for the Year Ending March 1976, Vol. 8,

Effects of contaminants, p. 25-118. U.S. Dep. Commer.,

NOAA. Available Arctic Environmental Assessment Center,

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

This study was designed to determine the acute and chronic

toxicity of crude oil and its component fractions on

physiological and behavioral mechanisms of selected Arctic and

subarctic organisms and to determine recovery rates of selected

organisms in laboratory and field studies.

Conclusions to date: Bioassays with invertebrates require

modification of standard bioassay procedures because they do not

complete their response in 96-h. Temperature has little effect

on toxicity, so that data generated at 12°C can be extrapolated

to colder climates. Oil exposures stimulate metabolism in fish,

rather than depress metabolism, as in crabs.

53 Rice, S. D., J. F. Karinen, and C. C. Brodersen. 1985.

Effects of oiled sediment on juvenile king crab. In Outer

Continental Shelf Assessment Program, Final Reports of

Principal Investigators, Vol. 29, p. 287-310. U.S. Dep.

Commer., NOAA. Available Arctic Environmental Assessment

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

This l-year project to determine effects of oiled sediment on

juvenile king crab began in April 1982. Juvenile king crab

(Paralithodes camtschatica) were exposed to the water-soluble

fraction of Cook Inlet crude oil (flow-through, stable

concentrations for 30 days) or to oiled sediments for 3 months.

The higher exposure concentrations of the water-soluble fraction

were toxic and affected survival, growth, feeding rate, and scope

for growth. Adverse effects were visible in just a few days. In

contrast, the oiled sediments did not cause any measurable

adverse effects on survival, feeding rate, growth, molting

success, or scope for growth during the 3-month exposure,

including those crabs exposed to the highest concentrations--2%.

Aromatic hydrocarbons were detected in some tissues of the crabs,

including the crabs exposed to oiled sediment. Most experimental

evidence suggests that exposures to oiled sediment will have

minimal impact directly on survival and growth of juvenile king

crab. However, the fact that hydrocarbons were detected in the

tissues means that there is some contact with the hydrocarbons,

and adverse effects are possible,

3 months would be required.

although exposures longer than
47

54 Rice, S. D., J. F. Karinen, and S. Korn. 1978. Acute and

chronic toxicity, uptake and depuration, and sublethal

response of Alaska marine organisms to petroleum

hydrocarbons. In D. A. Wolfe (editor), Marine Biological

Effects of OCS Petroleum Development, p. 11-24. U.S. Dep.

Commer., NOAA Tech. Memo. ERL-OCSEAP-1.

Oil effects studies at the Auke Bay Laboratory are reviewed. The

studies can be broken down into two basic themes: (1) Toxicity

challenge experiments, where an attempt is made to identify

sensitive species, life stages, factors that affect toxicity, or

components that are most responsible for toxicity; and

(2) sublethal physiological response, where an attempt is made to

identify, measure, and characterize physiological responses that

are indicative of oil stress. Results from the series of

sublethal effects studies may have application to monitoring

effects of oil and defining the mode of action of toxicants.

55 Rice, S. D., S. Korn, C. C. Brodersen, S. A. Lindsay, and

S. A. Andrews. 1981. Toxicity of ballast-water treatment

effluent to marine organisms at Port Valdez, Alaska. In

Proceedings 1981 Oil Spill Conference, Atlanta, Georgia,

March 2-5, p. 55-61. American Petroleum Institute,

Washington, DC.

Approximately 12 million gallons of oily ballast water is taken

ashore and treated daily at the Alyeska treatment plant, where

tankers take on crude oil at the terminus of the Trans Alaska

pipeline near Valdez, Alaska. Most oil is removed, but light

aromatic hydrocarbons (1 to 16 parts per million) remain in the

large volume of discharged effluent. Between May and July, the

concentration of aromatic hydrocarbons in the treated effluent

(measured by gas chromatography) generally declined as seasonal

temperatures increased. We measured the toxicity of the effluent

on site at Valdez. For the larvae of crustaceans the median

lethal concentration LC50 was between 10 and 20 percent of

treated effluent in 96-h static tests. For salmon fry and shrimp

in repeated acute flow-through assays, the (LC50) was quite

consistent, between 20 and 40 percent of treated effluent.

Because the concentrations of aromatic hydrocarbons was much

lower in the later tests but the toxicity of the effluent was not

lower, toxicants other than aromatic hydrocarbons must contribute

significantly to the toxicity of the effluent from the

ballast-water treatment plant.

56 Rice, S. D., S. Korn, and J. F. Karinen. 1981. Lethal and

sublethal effects on selected Alaskan marine species after

acute and long-term exposure to oil and oil components. In

Environmental Assessment of the Alaskan Continental Shelf.

Annual Reports of Principal Investigators for the Year

Ending March 1981, Vol. 4, Effects of Contaminants,

p. 61-78. U.S. Dep. Commer., NOAA. Available Arctic
48

Environmental Assessment Center, 222 W. 8th Ave., No. 56,

Anchorage, AK 99513.

This report consists of an Annual Report that summarizes status

and programs for FY 80 studies, followed by Quarterly Report

summarizing status of FY 81 studies. All of our studies are

applied laboratory studies, where we expose a variety of marine

species and life stages, to determine survival and effects of

short and long-term exposures to water-soluble fractions of oil.

We had two major objectives in FY 80: (1) to determine the

vulnerability of pink salmon alevins exposed to oil in a

simulated intertidal spawning environment. (2) to synthesize and

publish results to previous OCSEAP research.

57 Rice S. D., S. Korn, and J. F. Karinen. 1980. Lethal and

sublethal effects on selected Alaskan marine species after

acute and long-term exposure to oil and oil components. In

Environmental Assessment of the Alaskan Continental Shelf.

Annual Reports of Principal Investigators for the Year

Ending March 1980, Vol. 3, Effects, Contaminant Baselines,

p. 1-12. U.S. Dep. Commer., NOAA. Available Arctic

Environmental Assessment Center, 222 W. 8th Ave., No. 56,

Anchorage, AK 99513.

Fish were more sensitive than the Arctic invertebrates tested and

were comparable to pink salmon (subarctic species). Arctic

animals were highly adaptable to temperature changes, although

there were no effects of temperature on sensitivity of Arctic

animals to oil. Results indicate little difference in

sensitivity of Arctic and subarctic animals to oil. Lower

environmental temperatures in the Arctic could result in oil

persisting longer after a spill due to lower volatility and

biodegradation of oil components and because oil would become

trapped or immobilized in ice. There are fewer species in the

Arctic and food chains are very short. If a species is affected,

there would be few replacement species. The Arctic habitat is

more vulnerable, and once changed, less able to adjust, even

though individual species are generally very hearty and tolerant

of natural environmental extremes and limited amounts of

pollution.

58 Rice, S. D., S. Korn, and J. F. Karinen. 1978. Lethal and

sublethal effects on selected Alaskan marine species after

acute and long-term exposure to oil and oil components. In

Environmental Assessment of the Alaskan Continental Shelf.

Final Reports of Principal Investigators, Vol. 1, Biological

Studies, p. 1-32. U.S. Dep. Commer., NOAA. Available

Arctic Environmental Assessment Center, 222 W. 8th Ave.,

No. 56, Anchorage, AK 99513.

Annual Report: The research involves physiological and bioassay

tests of applied research on species indigenous to the Gulf of

Alaska, Bering Sea, and Beaufort Sea. The major emphasis has
49

shifted from strictly descriptive acute toxicity determinations

to mechanistic studies and sublethal tests that will eventually

allow prediction of oil impact on the biota. The studies have

two basic themes. Toxicity challenge experiments, where

sensitive species, life stages, factors that affect toxicity, or

components that are most responsible for toxicity are identified;

and sublethal physiological response, where physiological

responses that are indicative of oil stress are identified,

measured, and characterized.

59 Rice, S. D., S. Korn, and J. F. Karinen. 1977. Lethal and

sublethal effects on selected Alaskan marine species after

acute and long-term exposure to oil and oil components. In

Environmental Assessment of the Alaskan Continental Shelf.

Annual Reports of Principal Investigators for the Year

Ending March 1977, Vol. 12, Effects, p. 23-124. U.S. Dep.

Commer., NOAA. Available Arctic Environmental Assessment

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

Laboratory and field studies were conducted to determine recovery

rates of selected organisms and ecosystems from perturbations

resulting from either contamination or other disturbances

associated with petroleum development. The research involved

physiological and bioassay tests of applied research on species

indigenous to the Gulf of Alaska, Bering Sea, and Beaufort Sea.

The major emphasis of research has shifted from strictly

descriptive acute toxicity determinations to mechanistic studies

and sublethal tests that will eventually allow prediction of oil

impact on the biota. Studies to determine the acute toxicity of

the water-soluble fraction (WSF) of crude oil continue with

experiments with species not tested previously. Experiments

also continue with larvae of species not tested previously.

Emphasis is on intertidal species, such as mussels, barnacles,

snails, and sea urchins.

60 Rice, S. D., and D. A. Moles. 1991. Humpies and

hydrocarbons: Relating laboratory effects studies to

potential effects from the Exxon Valdez oil spill.

Unpublished manuscript. National Marine Fisheries Service,

Auke Bay Laboratory.

The potential effects from the Exxon Valdez oil spill on the pink

salmon, Oncorhynchus gorbuscha, are examined in light of

controlled laboratory studies. Eggs and alevins are tolerant of

short term water-borne oil exposures, but are especially

vulnerable in Prince William Sound where whole oil could

contaminate the intertidal redds. Out migrant fry in seawater

are about twice as sensitive as fry in freshwater, but exposure

to high levels of hydrocarbons in seawater from a spill is

unlikely. If spilled oil affects out migrant fry, laboratory

studies suggest that decreased growth will be the most likely

effect. Returning adults have declining abilities to metabolize

hydrocarbons, but their exposures are likely to be minimal.
50

61 Rice, S. D., D. A. Moles, J. F. Karinen, S. Korn, M. G.

Carls, C. C. Brodersen, J. A. Gharrett, and M. M. Babcock.

1984. Effects of petroleum hydrocarbons on Alaskan aquatic

organisms: A comprehensive review of all oil-effects

research on Alaskan fish and invertebrates conducted by the

Auke Bay Laboratory, 1970-81. U.S. Dep. Commer., NOAA Tech.

Memo. NMFS F/NWC-67, 128 p.

This report reviews and summarizes all oil-effects research by

the Auke Bay Laboratory from the beginning of these studies in

1970 through 1981. Both published and unpublished results from

62 studies are included, regardless of funding source. Research

is reviewed according to subject (e.g. toxicity, sublethal

effects, studies at Port Valdez). A bibliography and abstracts

are also included.

62 Rice, S. D., A. Moles, and J. W. Short. 1975. The effect

of Prudhoe Bay crude oil on survival and growth of eggs,

alevins, and fry of pink salmon, Oncorhynchus gorbuscha. In

Proceedings of 1975 Conference on Prevention and Control of

Oil Pollution, p. 503-507. American Petroleum Institute,

Washington, DC.

Standard 96-h bioassays with "total" oil solutions in fresh water

and seawater determined differences in sensitivity of the

developing life stages of pink salmon (Oncorhynchus gorbuscha).

Eggs were the most resistant and emergent fry (yolk sac absorbed)

the most sensitive to acute 4-day exposures. In fresh water, the

96-h median tolerance limit (TLm) of fry was 12 ppm as measured

in subsurface water by infrared spectrophotometry. In seawater,

it was 6 ppm.

Three life stages of alevins were exposed to IO-day

sublethal exposures of the water-soluble fraction to determine

what doses might affect growth. Growth was affected most

severely in alevins exposed during later developmental stages.

Decreased growth was observed in fry after 10-day exposures at

about 10% of the 96-h TLm limit for that life stage.

63 Rice, S. D., D. A. Moles, T. T. Taylor, and J. F. Karinen.

1979. Sensitivity of 39 Alaskan marine species to Cook

Inlet crude oil and No. 2 fuel oil. In Proceedings of 1979

Oil Spill Conference (Prevention, Behavior, Control,

Cleanup), p. 549-554. American Petroleum Institute,

Washington, DC.

The sensitivities of 39 subarctic Alaska species of marine fish

and invertebrates to water-soluble fractions of Cook Inlet crude

oil and No. 2 fuel oil were determined.

Sensitivity was correlated to habitat. Pelagic fish and

shrimp were the most sensitive animals to Cook Inlet crude oil

with 96-h median tolerance limits (TLm's) from 1-3 mg/l total

aromatic hydrocarbons. Benthic animals, including fish, crabs,

and scallops were moderately tolerant (TLm's to Cook Inlet crude
51

oil of 3-8 mg/l total aromatic hydrocarbons). Intertidal

animals, including fish, crabs, starfish, and many molluscs, were

the most tolerant forms to water-soluble fraction of petroleum

(TLm's greater than 8-12 mg/l of total aromatic hydrocarbons).

Most of the intertidal animals were not killed by static oil

exposures. No. 2 fuel oil was more toxic to most species than

Cook Inlet crude oil.

Sensitive pelagic animals are not necessarily more

vulnerable to oil spills than tolerant intertidal forms--oil may

damage intertidal environments more easily and adverse effects

may persist longer than in damaged pelagic environments.

64 Rice, S. D., J. W. Short, C. C. Brodersen, T. A.

Mecklenburg, D. A. Moles, C. J. Misch, D. L. Cheatham, and

J. F. Karinen. 1976. Acute toxicity and uptake-depuration

studies with Cook Inlet crude oil, Prudhoe Bay crude oil,

No. 2 fuel oil and several subarctic marine organisms.

Northwest Fisheries Center Processed Report, 90 p.

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

NOAA, 2725 Montlake Blvd. E., Seattle, WA 98112.

Several life stages of Alaskan fish, shrimp, scallops, and crab

were subjected to laboratory experiments to determine levels of

acute toxicity of Cook Inlet and Prudhoe Bay crude oils and fuel

oil and to determine the rate and degree of depuration of

hydrocarbons from sublethal concentrations of water-soluble

fractions of Cook Inlet crude oil.

Acute toxicity of crude oil was determined by standard

bioassays using both oil-water dispersions and water-soluble

fractions.

Results of the study are reported as median tolerance limits

in ppm oil and are given by species at specified temperature

ranges. Both larval and adult forms were tested. Larvae were

most sensitive, and median tolerance limits of all non-larval

stages of the eight species tested were similar.

Rate and amount of hydrocarbon uptake and depuration were

determined by exposing test organisms to seawater containing less

than 20% of the WSF concentration that would be expected to kill

50% of the animals in 96 h. Exposure and depuration periods each

lasted for 96 h. Gas chromatography was used to determine

hydrocarbon concentrations in samples of tissues taken

periodically during the test periods. All species tested

accumulated significant amounts of aromatic hydrocarbons;

depuration rates varied with species.

Behavioral observations of larvae near surface oil slicks

and physiological studies of the effect of dissolved oil

fractions on crab respiration were also made.1

65 Rice, S. D., J. W. Short, and J. F. Karinen. 1977 .

Comparative

sensitivity.

oil toxicity and comparative animal

In D. A. Wolfe (editor), Fate and Effects of

Petroleum Hydrocarbons in Marine Ecosystems and Organisms,

p. 78-94. Pergamon Press, New York.
52

A review is presented of studies concerning: (1) the behavior of

oil in water, (2) the methodology problems associated with

bioassays, (3) the comparative toxicity of oils, oil-water mixes,

and oil components, (4) the comparative sensitivity of different

life stages and species. Conclusions relating to these subjects

are presented in list form, incorporating recommendations and

guidelines for future work. Improvements in methodology are

advocated, in order that sensitivity tests may be applied to new

species and to previously investigated species with greater

accuracy.

66 Rice, S. D., J. W. Short, and J. F. Karinen. 1976.

Toxicity of Cook Inlet crude oil and No. 2 fuel oil to

several Alaskan marine fishes and invertebrates. In

Sources, Effects, and Sinks of Hydrocarbons in the Aquatic

Environment, p. 395-406.

Sciences, Washington, DC.

American Institute of Biological

We used 96-h static bioassay to determine the TLm's (median

tolerance levels) of 27 different invertebrate and vertebrate

Alaskan marine species exposed to WSF's (water-soluble fractions)

of Cook Inlet crude oil and No. 2 fuel oil. Concentrations of

oil in the exposure doses of the WSF's were determined by

infrared spectrophotometry.

No. 2 fuel oil was somewhat more toxic than the Cook Inlet

crude oil to some of the species. Fish were consistently among

the more sensitive species with 96-h TLm's from 0.81 to 2.94 ppm.

Some invertebrates were as sensitive as fish, while others were

quite resistant. Intertidal invertebrates were consistently

among the most resistant species.

It appears that Alaskan marine species may be slightly more

sensitive than similar species residing in more temperate

regions. However, the differences in observed sensitivity may be

due to the greater toxicity of oil at lower temperatures (because

of greater persistence of hydrocarbons) rather than to actual

increases in the sensitivity of the animals.

67 Rice, S. D., and R. E. Thomas. 1989. Effect o f

pre-treatment exposures of toluene or naphthalene on the

tolerance of pink salmon (Oncorhynchus gorbuscha) and kelp

shrimp (Eualis [sic] suckleyi). Comp. Biochem. Physiol. C

Comp. Pharmacol. 94:289-293.

1. Toluene pre-treatment exposures did not affect survival

of kelp shrimp or pink salmon fry in subsequent bioassays.

2. Naphthalene pre-treatment exposures to kelp shrimp

caused increased sensitivity in subsequent bioassays.

3. Naphthalene pre-treatment exposures to pink salmon

caused increase in tolerance in subsequent bioassays. Tolerance

increase was proportional to concentration and duration of

pre-treatment exposures,

depuration.

and decreased with periods of
53

68 Rice, S. D., R. E. Thomas, and J. W. Short. 1977. Effect

of petroleum hydrocarbons on breathing and coughing rates

and hydrocarbon uptake-depuration in pink salmon fry. In F.

J. Vernberg, A. Calabrese, F. P. Thurberg, and W. B.

Vernberg (editors), Physiological Responses of Marine Biota

to Pollutants, p. 259-277. Academic Press, New York.

We report the changes in breathing rates during the extended

exposure of pink salmon, Oncorhynchus gorbuscha, fry to oil, and

examine the return to near normal rates during extended exposure.

Cook Inlet and Prudhoe Bay crude oil and No. 2 fuel oil were used

in the experiments. The 3 oils caused similar increases in

breathing and coughing rates in the fry, and these rates

increased in proportion to the oil concentrations, as measured by

UV but not by IR (suggesting that naphthalenes rather than

paraffins are responsible for this effect). Breathing and

coughing rates of the fry remained above normal during exposure

to a constant dose of oil for 72 h.

the first 24 h of exposure,

High breathing rates during

elimination of most aromatics by

20 h, and the continued high breathing rates during the constant

dose exposure for 72 h indicate that salmon fry can cope with a

sublethal exposure to hydrocarbons, but at the cost of an

increased metabolic rate. Increased metabolic rates may be

detrimental to survival if the stress persists for long periods

of time.

69 Schwartz, J. P. 1985. Effect of oil-contaminated prey on

the feeding and growth rate of pink salmon fry (Oncorhynchus

gorbuscha). In F. J. Vernberg, A. Calabrese, and W.

Vernberg (editors), Marine Pollution and Physiology: Recent

Advances, p. 459-476. Univ. South Carolina Press, Columbia,

SC.

Pink salmon fry fed 0.6, 3.2 and 6.5 ppm oil-contaminated prey

(OCP) for 10 d ate less than fry fed uncontaminated prey. Fry

fed 0.6 ppm OCP ate more than fry fed 6.5 ppm OCP. Fry feeding

rates at each OCP dose level did not improve when the fish were

fed uncontaminated prey for 5 d after exposure.

The growth rate of pink salmon fry exposed to OCP for 50 d

decreased with increased dose levels of OCP. The wet weight of

fry fed 0.6 ppm was significantly greater than fry fed 3.2 and

6.5 ppm OCP after 50 d. Decreases in fry feeding and growth

rates were not proportional to OCP hydrocarbon concentrations,

but reductions in fry feeding rate correlated with reduction in

growth rate. Reductions in the growth of fry exposed to the 96-h

LC50 (0.7 ppm) of the water soluble fraction (WSF) of Cook Inlet

crude oil were greater than the reduction in growth of fry fed

OCP.

Pink salmon fry accumulated bicyclic aromatic hydrocarbons

from OCP during the first 12 h of exposure, but concentrations

returned to levels below detection after 10-d exposure to OCP.

Fry exposed to WSF rapidly accumulated both mono- and bicyclic

hydrocarbons within 24 h.
54

OCP were not acutely toxic to pink salmon fry. Fry were

able to feed and grow at OCP hydrocarbon concentrations that were

5-10 times the 96-h LC50 for the WSF of Cook Inlet crude oil in

sea water. However, reductions in the feeding rate of pink

salmon fry both during and after exposure to OCP, and reductions

in fry growth rate from exposure to OCP, have the potential for

reducing fry survival during early sea life.

70 Schwartz, J. P. 1984. The effects of oil-contaminated prey

on the feeding, growth, and related energetics of pink

salmon, Oncorhynchus gorbuscha Walbaum, fry. Ph.D. Thesis,

Univ. Alaska, Fairbanks, 99 p.

Pink salmon, Oncorhynchus gorbuscha Walbaum, fry were exposed to

oil-contaminated prey (OCP) in a series of experiments to

determine the effects of oil exposure via the diet on the ability

of pink fry to survive. Brine shrimp, Artemia salina, nauplii

were contaminated with petroleum hydrocarbons by exposure to the

water-soluble fraction (WSF) of Cook Inlet crude oil and fed to

the fish. Feeding rates were measured for 10 days using OCP and

for 5 days using uncontaminated prey (post-exposure period). In

a separate experiment, fry growth was measured over a 50 day

period. In another experiment, fry oxygen consumption, food

absorption and utilization, and ammonia excretion was measured to

determine the effects of OCP on fry metabolic activity.

Fry feeding rates were reduced by exposure to OCP, and

remained suppressed during the post-exposure period. Chronic

exposure to OCP for 50 days reduced fry growth. OCP were not

lethal to the fry. There was no change in fry oxygen consumption

or ammonia excretion from exposure to OCP, but the fish exposed

to OCP absorbed less food than controls and continued to absorb

less food for 7 days after exposure. Results indicate that

exposure to OCP can reduce fry growth primarily by reducing food

intake, but additional nutrition is lost from the non-absorption

of ingested food. Reductions in growth could decrease fry

survival, and thereby reduce the number of returning adult pink

salmon.

71 Short, J. W., S. D. Rice, and D. L. Cheatham. 1976.

Comparison of two methods for oil and grease determination.

In D. W. Hood and D. C. Burrell (editors), Assessment of the

Arctic Marine Environment: Selected Topics, p. 451-465.

Univ. Alaska, Fairbanks.

A gravimetric method is used by government regulatory agencies

for determining levels of oil pollutants in discharge waters.

This method involves extraction with an organic solvent,

evaporation at elevated temperatures, and gravimetric

determination of the residue. The authors compared oil content

determined by the gravimetric method with oil content determined

by infrared spectrophotometry for toxic water-soluble fractions

of Prudhoe Bay and Cook Inlet crude oils and a No. 2 fuel oil.
55

The gravimetric method is adequate for grease but not for

the oils. Recovery of a synthetic grease standard was

98 percent, whereas the recovery of the three pure oils ranged

from 52 to 65 percent by the gravimetric method. Recovery of all

the oils and the grease standard was essentially 100 percent by

the infrared method. The differences between the two methods are

ascribed to significant losses of volatile compounds from the

oils during the evaporation step of the gravimetric method.

Gravimetric estimates of oil in toxic concentrations of

water-soluble fractions ranged from 0 to 36 percent of those

determined by the infrared method. Oil content of the No. 2 fuel

oil water-soluble fractions was below detectable limits of the

gravimetric method (1.5 mg/liter). Four-day median tolerance

limits of shrimp (Eualus fabricii) and scallops (Chlamys rubida),

as evaluated by the infrared water-soluble fractions of the three

oils, were between 0.25 mg/liter (No. 2 fuel oil) and

3.82 mg/liter (crude oils).

It is concluded that the gravimetric method is sensitive to

heavier compounds, but these have only a casual relationship to

acute toxicity. Concentrations of oil in water known to have

adverse effects are much lower than can be detected by the

standard gravimetric method. When oil concentrations in water

are to be measured and correlated with chemical toxicity, the

gravimetric procedure should be supplemented with a method

specific for the more soluble and volatile components.

72 Stickle, W. B., M. A. Kapper, T. C. Shirley, M. G. Carls,

and S. D. Rice. 1987. Bioenergetics and tolerance of the

pink shrimp (Pandalus borealis) during long-term exposure to

the water-soluble fraction and oiled sediment from Cook

Inlet crude oil. In W. B. Vernberg, A. Calabrese, F. P.

Thurberg, and F. J. Vernberg (editors), Pollution Physiology

of Estuarine Organisms, p. 87-106. Univ. South Carolina

Press, Columbia, SC.

Adult Pandalus borealis were exposed to the water-soluble

fraction (WSF) of and sediments oiled with Cook Inlet crude oil.

Fifteen times more total aromatic hydrocarbons were

bioaccumulated into the soft tissues of the shrimp cephalothorax

during 10-d exposure to 550 ppb WSF aromatic hydrocarbons than to

84-d exposure to 2% oiled sediment. Little bioaccumulation of

aromatic hydrocarbons into abdominal muscle of either group

occurred. The tolerance of shrimp to the WSF fraction declined

linearly with duration of exposure, to a 28-d LC50 value of

28 ppb total aromatic hydrocarbons. Scope for growth, or the

energy budget, of shrimp remained positive at all sublethal WSF

concentrations but declined significantly below control values,

as a result of a decreased rate of food consumption, in shrimp

exposed to concentration which were eventually lethal.

Insufficient mortality occurred in shrimp exposed to oiled

sediment to calculate an LC50 value; in fact, shrimp exposed to

the highest concentrations exhibited increased scope for growth

compared with control shrimp as a result of increase food

consumption. Shrimp exposed to the WSF were far more vulnerable
56

to oil effects on their survival and bioenergetics than shrimp

exposed to oiled sediment.

73 Stickle, W. B., S. D. Rice, and A. Moles. 1984.

Bioenergetics and survival of the marine snail Thais lima

during long-term oil exposure. Mar. Biol. 80:281-289.

The carnivorous snail Thais lima was fed Mytilus edulis during a

28-d exposure to the water soluble fraction (WSF) of Cook Inlet

crude oil. The LC50 of T. lima declined from >3,000 ppb aromatic

hydrocarbons on Day 7 to 818 + 118 ppb on Day 28. The LC50 of

M. edulis declined from > 3,000 ppb aromatic hydrocarbons on Day

7 to 1,686 + 42 ppb on Day 28. Predation rate declined linearly

with increasing aromatic hydrocarbon concentration up to 302 ppb;

little predation occurred at 538 ppb and none at 1,160 or

1,761 ppb. Snail absorption efficiency averaged 93.5% and did

not vary as a function of WSF dose. Total energy expenditure

(R+U) increased at 44 ppb aromatics and declined at lethal WSF

exposures. At sublethal WSF exposures, percentages of total

energy expenditure were: respiration (87%), ammonia excretion

(9%) and primary amine loss (4%). These percentages did not vary

as a function of WSF dose or time. 0xygen:nitrogen ratios were

not affected by WSF concentration or time and indicated that

T. lima derived most of its energy from protein catabolism. The

uptake of aromatic hydrocarbons into the soft tissues of snails

and mussels was directly related to the WSF concentration.

Naphthalenes accounted for 67 to 78% of the aromatic hydrocarbons

in T. lima and 56 to 71% in M. edulis. The scope for growth was

negative above 150 ppb WSF aromatic hydrocarbons and above

1,204 ppb soft-body aromatic hydrocarbons. These snails were

physiologically stressed at an aromatic hydrocarbon concentration

which was 19% of the 28-d WSF LC50 (818 + 118 ppb) and/or 48% of

the 28-d LC50 of soft tissue aromatics (2,502 ppb).

74 Stickle, W. B., Jr., S. D. Rice, C. Villars, and W. Metcalf.

1985. Bioenergetics and survival of the marine mussel,

Mytilus edulis L., during long-term exposure to the

water-soluble fraction of Cook Inlet crude oil. In F. J.

Vernberg, F. P. Thurberg, A. Calabrese, and W. B. Vernberg

(editors), Marine Pollution and Physiology: Recent

Advances, p. 427-446. Univ. South Carolina Press, Columbia,

SC.

The sensitivity of Mytilus edulis on exposure to the WSF of Cook

Inlet crude oil increased with duration of exposure. Scope for

growth declined in an inverse manner with WSF concentration to

zero at 1163 ppb aromatic hydrocarbons, this value represents

82% of the 28-d LC50. Therefore, M. edulis is not a particularly

sensitive species to utilize for the study of sublethal stress

during WSF exposures over long-term (one month) exposure periods.

The primary determinant of a decline in scope with increased

concentration of WSF was a decline in algal consumption. Scope

for growth regressed in an inverse manner-against total tissue
57

aromatic hydrocarbons at WSF concentrations up to 1083 ppb over

28-d exposure, but no relationship existed between scope for

growth and total tissue concentration of aliphatic hydrocarbons.

At a higher WSF aromatic hydrocarbon concentration (1083 ppb),

uptake of aromatic hydrocarbons was probably reduced because of a

reduced algal clearance (= feeding) rate. 0xygen:nitrogen ratios

were low indicating protein catabolism, and did not decline with

increased WSF exposure; however, mussels were in the postspawning

phase of their reproductive cycle.

75 Stickle, W. B., T. D. Sabourin, and S. D. Rice. 1982.

Bensitivity and osmoregulation of coho salmon, Oncorhynchus

kisutch, exposed to toluene and naphthalene at different

salinities. In W. B. Vernberg, A. Calabrese, F. P.

Thurberg, and F. J. Vernberg (editors), Physiological

Mechanisms of Marine Pollutant Toxicity, p. 331-348.

Academic Press,. New York.

Coho salmon smolts were more sensitive to toluene and naphthalene

in seawater than in freshwater. Tolerance dropped linearly from

0 through 10, 20, and 30%. Smolt tolerance at 30% were 54% and

63% of the 48-h TLm in freshwater for toluene and naphthalene,

respectively. Smolt tolerances to toluene and naphthalene were

the same after 12, 22, and 42 d of acclimation to seawater as

they were after only 1 d of acclimation. The increase in

sensitivity was not transient nor did it appear related to

acclimation stress because the smolts gained 30% in weight in

42 d.

Toluene and naphthalene affected serum osmolality and ions

but only at lethal concentrations of 100 and 130% of the 48-h

TLm. At those exposure concentrations, osmolality, Na+, and Clmoved down the diffusion gradients between the serum of the

smolts and ambient water: decreased in freshwater smolts and

increased in seawater smolts. At the same concentration, K+

concentrations in the serum increased, even in freshwater smolts,

indicating cellular damage. Exposures of 70% of the 48-h TLm had

no effect on serum osmolality or ions. Consequently, we conclude

that the increase in sensitivity of smolts in seawater is not

related to a failure in ion-regulation ability, but rather the

loss of ion-regulating ability at lethal exposures is symptomatic

of other toxic actions. Increased sensitivity of smolts in

seawater is not transient and the cause remains unknown.

76 Taylor, T. L., and J. F. Karinen. 1977. Response of the

clam, Macoma balthica (Linnaeus), exposed to Prudhoe Bay

crude oil as unmixed oil, water-soluble fraction, and

oil-contaminated sediment in the laboratory. In D. A.

Wolfe (editor), Fate and Effects of Petroleum Hydrocarbons

in the Marine Ecosystems and Organisms, p. 229-237.

Pergamon Press, New York.

The small clam,

subjected to oil

Macoma balthica (Linnaeus 1758), will likely be

slicks layered on the mud and to water-soluble
58

fractions (WSF) of crude oil or oil-contaminated sediment.

Groups of adult clams in or on their natural sediment were

exposed in flow-through aquaria at 7°-12°C to various

concentrations of Prudhoe Bay crude oil layered on the mud

surface, the water-soluble fraction of the crude oil, and

oil-treated sediment (OTS).

Gentle settling of crude oil over clam beds had negligible

effects on clams observed for 2 months. Water-soluble and

oil-treated sediment fractions of Prudhoe Bay crude oil inhibited

burrowing and caused clams to move to the sediment surface.

Responses were directly proportional to concentrations of the WSF

or amount of OTS. The l-h and 72-h effective median

concentrations of the WSF for the responses of burrowing by

unburied clams and surfacing by buried clams were 0.234 and

0.367 ppm naphthalene equivalents respectively. The interpolated

amount of OTS needed for a 50% surfacing response within 24 h was

0.67 g OTS cm-2.

Although short-term exposures of clams to the WSF of crude

oil and OTS caused few deaths, behavioral responses of clams to

oil may be of great importance to their survival in the natural

environment. In these laboratory tests, many of the clams

recovered, but in nature clams that come to the sediment surface

may be eaten by predators or die from exposure.

77 Thomas, R. E., and S. D. Rice. 1987. Effect of

water-soluble fraction of Cook Inlet crude oil on swimming

performance and plasma cortisol in juvenile coho salmon

(Oncorhynchus kisutch). Comp. Biochem. Physiol. C Comp.

Pharmacol. Toxicol. 87:177-180.

1. Swimming performance of juvenile coho salmon decreased

and plasma cortisol increased following 48-h exposure to the

water-soluble fraction (WSF) of Cook Inlet crude oil at 75% of

the LC50.

2. Exposure of 25 and 50% of the LC50 did not significantly

reduce swimming performance.

3. Plasma cortisol concentrations were highest in fish

exposed to both the combined stress of WSF exposure and of forced

swimming in a stamina tunnel.

78 Thomas, R. E., and S. D. Rice. 1986. The effects of

salinity on uptake and metabolism of toluene and naphthalene

by Dolly Varden, Salvelinus malma. Mar. Environ. Res.

18:203-214.

Dolly Varden (Salvelinus malma) were force-fed 14C-toluene or 14C-naphthalene and held for 12, 24 and 48 h in freshwater or

seawater (30%) in order to determine the effect of salinity on

uptake and metabolism of aromatic hydrocarbons. Fish held in

seawater had slightly more

held in freshwater.

14C removed from the gut than did fish

Whether held in freshwater or seawater,

toluene was more readily metabolized by fish than was

naphthalene; after 48 h, 2-3 times more 14C from toluene was in
59

the metabolite fraction of liver and central nervous system

tissues. Fish held in seawater metabolized significantly less

toluene and naphthalene than those in freshwater. The increased

toxicity of toluene and naphthalene to Dolly Varden held in

seawater appears to be the result of the decreased metabolism of

these compounds by the fish.

79 Thomas, R. E., and S. D. Rice. 1986. Effect of temperature

on uptake and metabolism of toluene and naphthalene by Dolly

Varden char, Salvelinus malma. Comp. Biochem. Physiol. C

Comp. Pharmacol. Toxicol. 84:83-86.

14C 

1. Tissue concentrations and metabolism of 14C toluene and

naphthalene were measured in Dolly Varden char at exposure

temperatures of 4 and 12°C after oral administration of the

hydrocarbons.

2. Tissue concentrations of toluene and toluene metabolites

were slightly less at 12 than 4°C.

3. Metabolism of toluene was significantly greater at

12 than 4°C.

4. Tissue concentrations of naphthalene and naphthalene

metabolites were less at 12 than 4°C after 48 h exposure.

5. Naphthalene metabolism was not consistently different

between fish exposed at 4 and 12°C.

6. Failure to adequately metabolize toluene at reduced

temperatures appears to contribute to the increased sensitivity

of fish to toluene when exposed at low temperatures.

7. Lack of temperature effect on naphthalene metabolism

correlates with lack of temperature-induced changes in the

toxicity of naphthalene to fish.

80 Thomas, R. E., and S. D. Rice. 1985. Effect of

pretreatment exposure to toluene and naphthalene on the

subsequent metabolism of dietary toluene and naphthalene by

Dolly Varden, Salvelinus malma. In F. J. Vernberg, F. J.,

F. P. Thurberg, A. Calabrese and W. A. Vernberg (editors),

Marine Pollution and Physiology: Recent Advances,

p. 505-520. Univ. South Carolina Press, Columbia, SC.

Pretreatment exposure of Dolly Varden to naphthalene in sea water

resulted in increased metabolism of orally-administered 14C-naphthalene, but pretreatment exposure to toluene did not

alter the rate of metabolism of orally-administered 14C-toluene.

Naphthalene is less polar than toluene and thus requires

significant bioconversion before it is excreted by fish. Toluene

is more polar and is readily excreted by fish, therefore

inducement of increased bioconversion of toluene in the fish is

not needed.

Several pretreatment factors influenced the rate of

naphthalene bioconversion, including; 1) length of pretreatment

exposure, 2) concentration of pretreatment exposure, and

3) duration of depuration after pretreatment exposure.
60

Pretreatment exposure of fish to naphthalene (75% of the

96-h LC50) for 48 h doubled or tripled the percentage of

naphthalene metabolites in brain, liver, and muscle tissue, and

decreased the amount of carbon-14 recovered in those tissues.

Pretreatment exposure for only 24 h caused some increases in

tissue metabolites, but no significant differences in percent

metabolites of tissue burden were measured between the

pretreatment fish and controls which were not pretreated.

The concentration of the pretreatment is directly related to

the increase in bioconversion rate of naphthalene. Exposure to

25 and 50% of the 96-h LC50 did not result in increased

metabolism of naphthalene or in lower tissue burdens of the

hydrocarbons. Exposure at 75% of the LC50 resulted in

significant increases in tissue metabolites and decreases in

total tissue hydrocarbon burden.

Periods of depuration following pretreatment exposure

resulted in a decrease in the induced rate of metabolism of

naphthalene. Depuration for as little as 24 h resulted in the

return of the naphthalene bioconversion rate to that of control

fish.

Pretreatment exposure of fish can induce increased

metabolism of aromatic hydrocarbons, such as naphthalene, and may

increase the survival of fish in subsequent exposures. Dinuclear

and polynuclear aromatic hydrocarbons are quite lipophilic and

must be metabolized before they can be excreted. Inducement of

increased metabolism of these compounds, resulting from a

previous exposure, increases excretion and therefore reduces

hydrocarbon accumulation in the tissues of fish.

81 Thomas, R. E., and S. D. Rice. 1982. Metabolism and

clearance of phenolic and mono-, di-, and polynuclear

aromatic hydrocarbons by Dolly Varden Char. In W. B.

Vernberg, A. Calabrese, F. P. Thurberg, and F. J. Vernberg

(editors), Physiological Mechanisms of Marine Pollutant

Toxicity, p. 161-176. Academic Press, New York.

Once hydrocarbons were absorbed from capsules placed in the gut

of Dolly Varden char, size and polarity of the hydrocarbons

influenced the elimination, metabolism, and tissue distribution.

Size of the hydrocarbon appeared to be the most critical factor

in excretion of hydrocarbons by the gills. The gills of fish can

easily excrete phenolic and mononuclear aromatic compounds. Some

naphthalene was excreted by the gills, but virtually none of the

polynuclear aromatic hydrocarbons were excreted through the

gills. Even though the partition coefficients (log of

octanol/water partition) of phenol and cresol are about 1/10 of

toluene, these similar-sized hydrocarbons were excreted from the

gills in approximately equal amounts.

Polar phenolic compounds were excreted into the cloacal

chamber but not toluene or the larger polynuclear aromatic

hydrocarbons. Partition coefficient is apparently a more

important factor than size in excretion of hydrocarbons into the

cloacal chamber because the excretion of phenol and cresol into
61

the cloacal chamber was more than 10 times that of similar-sized

toluene.

The excretion of the largest hydrocarbons tested, anthracene

and benzo[a]pyrene, was minimal in 24 h (3.2% and l.2%,

respectively). These compounds were slowly absorbed from the gut

probably because they are relatively nonpolar and have a high

partition coefficient (octanol/water), therefore are more

difficult to remove from a lipid matrix (membrane). The mobility

of these compounds between tissues is limited, and they probably

have to be metabolized before excretion. Consequently,

metabolism in the liver and secretion into the bile is probably

the most important pathway for excretion of large molecular

weight hydrocarbons; however, this is a relatively slow proces

that takes much longer than 24 h.

82 Thomas, R. E., and S. D. Rice. 1981. Excretion of aromatic

hydrocarbons and their metabolites by freshwater and

seawater Dolly Varden char. In F. J. Vernberg,

A. Calabrese, F. P. Thurberg, and W. B. Vernberg (editors),

Biological Monitoring of Marine Pollutants, p. 425-448.

Academic Press, New York.

The gills were the most important pathway for excretion of

carbon-14 from 14C naphthalene. Most of the carbon-14 excreted

by the gills was still attached to the parent compound. About

10% of the excreted carbon-14 appeared in the cloacal chamber,

mostly as metabolites. Less than 1% of the total carbon-14 was

excreted in the urine, predominantly as metabolites.

Tissues retained a significant amount of carbon-14 at 24 h.

Although muscle contained large amounts of carbon-14 because of

its mass, the gall bladder had the highest specific activity.

The brain also retained significant quantities of carbon-14.

Although more 14C toluene was excreted and metabolized than 14C

naphthalene, more 14C naphthalene was retained in the tissue. A

lower percentage of the carbon-14 was recovered in 14C

naphthalene metabolites than in 14C toluene metabolites.

Seawater and freshwater Dolly Varden char excreted similar

amounts of carbon-14; however, the percentage of metabolites in

the excretions and tissues of seawater fish was lower than the

percentage of metabolites in excretions and tissues of freshwater

fish. For example, we recovered greater amounts of carbon-14

with a lower percentage of metabolites from the brain-spinal cord

of seawater fish than from the brain-spinal cord of freshwater

fish--possibly explaining why seawater Dolly Varden are more

sensitive to aromatic hydrocarbons and the water-soluble fraction

of oil than freshwater Dolly Varden.

83 Thomas, R. E., and S. D. Rice. 1979. The effect of

exposure temperature on oxygen consumption and opercular

breathing rates of pink salmon fry exposed to toluene,

naphthalene, and water-soluble fractions of Cook Inlet crude

oil and No. 2 fuel oil. In W. B. Vernberg, A. Calabrese,

F. P. Thurberg, and F. J. Vernberg (editors), Marine
62

Pollution: Functional Responses, p. 39-52. Academic Press,

New York.

Breathing rates were measured in pink salmon fry exposed to

equivalent concentrations of water soluble concentrations of Cook

Inlet crude oil (containing mononuclear and dinuclear aromatics)

and No. 2 fuel oil (containing dinuclear aromatics) at 4° and

l2°C to determine whether responses differ with temperature.

Opercular breathing and oxygen consumption in fry exposed to

toluene (mononuclear) and naphthalene (dinuclear) were measured

and indicated that increased breathing rates reflect increased

energy demands as toxicants are metabolized. Exposure of the fry

to 107%, 70%, and 45% of the 24-h median tolerance limits of

naphthalene resulted in increased breathing rates; exposure to

toluene concentrations of 94% and 69% produced increased

breathing rates, but breathing rates at exposures of 45% and 30%

remained unchanged. For both toluene and naphthalene, increases

in breathing rates with increased exposure were linear.

Results of exposure to the four toxicants at 4° and 12°C

were not uniform. At 4°C, 17 of 24 fry exposed to toluene had

died 15 h after initial exposure; no deaths occurred among fry

exposed to toluene at 12°C or fry exposed to other substances at

either temperature. Greatest increase in breathing rates

occurred with exposures to naphthalene and No. 2 fuel oil at 4°C,

with breathing rates among fry nearly doubled. Results of the

study indicate that lower temperatures may influence the effect

of hydrocarbons on coldwater fish and that oils with relatively

high concentrations of mononuclear aromatic hydrocarbons may be

particularly damaging.1

84 Thomas, R. E., and S. D. Rice. 1975. Increased opercular

rates of pink salmon (Oncorhynchus gorbuscha) fry after

exposure to the water-soluble fraction of Prudhoe Bay crude

oil. J. Fish. Res. Board Can. 32:2221-2224.

The opercular rates of pink salmon (Oncorhynchus gorbuscha) fry

were measured during 24-h exposure to sublethal concentrations of

the water-soluble fraction of Prudhoe Bay crude oil. Opercular

rates increased significantly for as long as 9 and 12 h after

exposure to water-soluble fractions prepared from oil-water

solution of 2.83 and 3.46 ppm. The increases in rates were

proportional to increases in dose. Recording changes in

opercular rates appears to be a suitable method for detecting

sublethal physiological effects of stress, because the observed

changes occurred at approximately 20% of the 96-h LC50.

85 Thomas, R. E., S. D. Rice, M. M. Babcock, and A. Moles.

1989. Differences in hydrocarbon uptake and mixed function

oxidase activity between juvenile and spawning adult coho

salmon (Oncorhynchus kisutch) exposed to Cook Inlet crude

oil. Comp. Biochem. Physiol. C Comp. Pharmacol. 93:155-159.
63

1. Coho salmon smolts accumulated up to 30 times more

hydrocarbon in selected tissues than did coho salmon jacks when

exposed to water-soluble fraction of Cook Inlet crude oil.

2. Aryl hydrocarbon hydroxylase (AHH) activity continued to

increase during 12 days of exposure in coho smolts, in contrast

to jacks in which AHH activity peaked at 2 days and remained

elevated during 20 days of exposure.

3. Cytochrome P-450 concentrations peaked in 48 h and

returned to control levels in exposed smolts. In exposed and

control jacks, cytochrome P-450 decreased during the 20 day

experimental period. Concentrations of P-450 were significantly

greater, however, in exposed jacks than in control jacks.

4. Aryl hydrocarbon hydroxylase (AHH) activity and

cytochrome P-450 concentrations were greater in non-exposed coho

smolts and jacks than in non-exposed pre-spawning adults.

5. Coho jacks were not as responsive as coho smolts in terms

of P-450 induction; however, they were more responsive than

anticipated for a fish in a terminal life stage.

86 Thomas, R. E., S. D. Rice, M. M. Babcock, and A. Moles.

1988. Differences in hydrocarbon uptake and mixed function

oxidase activity between juvenile and spawning adult coho

salmon exposed to Cook Inlet crude oil. Am. Zool.

28(4):104A. (Abstract only).

Coho salmon smolts accumulated up to 30 times more hydrocarbon in

selected tissues than did coho salmon jacks when exposed to

water-soluble fraction of Cook Inlet crude oil. Aryl hydrocarbon

hydroxylase (AHH) activity continued to increase during 12 days

of exposure in coho smolts in contrast to jacks in which AHH

activity peaked at 2 days and remained elevated during 20 days of

exposure.

Cytochrome P-450 concentrations peaked in 48 h and returned

to control levels in exposed smolts.

jacks,

In exposed and control

cytochrome P-450 decreased during the 20 day experimental

period; however, concentrations were significantly greater in

exposed jacks than in control jacks. Aryl hydrocarbon

hydroxylase (AHH) activity and cytochrome P-450 concentrations

were greater in non-exposed coho smolts and jacks than in

non-exposed pre-spawning adults. Coho jacks were not as

responsive as coho smolts in terms of P-450 induction; however,

they were more responsive than anticipated for a fish in a

terminal life stage.

87 Thomas, R. E., S. D. Rice, and S. Korn. 1987. Reduced

swimming performance of juvenile coho salmon (Oncorphynchus

[sic] kisutch) exposed to the water-soluble fraction of Cook

Inlet crude oil. In W. B. Vernberg, A. Calabrese, F. P.

Thurberg, and F. J. Vernberg (editors), Pollution Physiology

of Estuarine Organisms, p. 127-137. Univ. South Carolina

Press, Columbia, SC.
64

The magnitude of the effect exposure to the water-soluble

fraction (WSF) of Cook Inlet crude oil on swimming performance of

juvenile coho salmon (Oncorhynchus kisutch) was found to be

dependent on both the concentration of the WSF and the length of

the time that fish were exposed. Swimming activity was reduced

to a greater extent at higher concentrations than at lower

concentrations. Some of the lower concentrations, which did not

significantly reduce swimming performance when fish were exposed

for only 48 h, had a greater effect when fish were exposed for

5 or 13 d. When fish were transferred to clean water for a

period of depuration prior to determining swimming performance,

control levels of swimming activity were restored within 8 h of

transfer.

It is suggested that exposure of salmon to the WSF of crude

oil induces an increased metabolism which reduces the energy

available for swimming. When fish are no longer exposed to the

WSF, this energy is no longer required and swimming performance

is restored to normal.
65

Prince William Sound and Related Papers/Reports

88 Barr, L. 1970. Alaska's fishery resources: The shrimps.

U.S. Fish Wildl. Serv. Fish. Leafl. 631, 10 p.

Shrimp fishing began in Alaska over 50 years ago. Recently the

annual domestic catch has been as high as 40 million pounds.

Japanese and Soviet Union fishermen operating in Alaska waters

have caught as much as 70 million pounds annually in recent

years.

The five commercially important shrimp of Alaska belong to

the family Pandalidae; the most important is the pink shrimp,

Pandalus borealis.

The complicated life histories of these shrimp are all

similar. The shrimp develop first as males and after several

years transform to females, which they remain for the rest of

their lives.

United States fishermen use otter trawls, beam trawls, and

pots, and deliver their catch to ports in Alaska; foreign

fishermen use larger otter trawls and process the catch at sea.

The Alaska Department of Fish and Game and the Bureau of

Commercial Fisheries are studying shrimp. They are sampling the

commercial catch, trying to improve the product, and conducting

exploratory fishing and biological research.

89 Barr, L. 1970. Diel vertical migration of Pandalus borealis

in Kachemak Bay, Alaska. J. Fish. Res. Board Can. 27:669-676.

Catches of Pandalus borealis by surface-to-bottom pot fishing in

Kachemak Bay, Alaska, showed that this species was distributed in

large numbers throughout the water column at night and suggested

a diel vertical migration. The shrimp apparently left the

vicinity of the bottom about dusk and returned about dawn; the

apparent length of time they were off bottom was directly related

to the length of the night. Bottom trawling clearly showed lower

availability of P. borealis on bottom at night than during the

day. Catches of shrimp by pot fishing and trawling indicated

that small P. borealis have a greater tendency to migrate

vertically than the larger individuals.

90 Calvin, N. I., and R. J. Ellis. 1978. Quantitative and

qualitative observations of Laminaria dentigera and other

subtidal kelps of southern Kodiak Island, Alaska. Mar.

Biol. 47:331-336.

Observations were made and quantitative samples of non-canopy

kelps were collected, using SCUBA, at 9 subtidal sites off

southern Kodiak Island in May, 1976.

observations were made.

At a 10th site, only

The most abundant species in the

quadrants were

gardneri,

Laminaria dentigera, L. yezoensis, Pleurophycus

Agarum cribrosum, and Alaria marginata. We found

fertile plants of all these species. L. dentigera dominated at
66

all sampled sites except one within a bed of Nereocystis

luetkeana, where P. gardneri was dominant. L. yezoensis was

dominant at the site which was visited but not sampled. The

average wet weight of non-canopy kelps in the 55 samples was

12 kg m-2, and the quadrant range was 2 to 35 kg m-2

. The

reported ranges of P. gardneri and Alaria marginata are extended

westward from Montague Island (147°22'W) to Bumble Bay, Kodiak

Island (154°43'W), where A. marginata was found at a depth of

8.5 m.

91 Calvin, N. I., and S. C. Lindstrom. 1980. Intertidal algae

of Port Valdez, Alaska: species and distribution with

annotations. Bot. Mar. 23:791-797.

We found 18 species of Chlorophyta, 20 species of Phaeophyta, and

42 species of Rhodophyta at 7 intertidal sites in Port Valdez,

Alaska. Of these species, Kornmannia zostericola,

Antithamnionella pacifica, Cryptonemia borealis, Halosaccion

firmum, Neodilsea integra, and Ceramium gardneri were found

beyond their previously reported range. Dumontia simplex, known

from Japan and previously reported in Alaska only at Amchitka

Island in the Aleutian Islands, was found at one site. Fucus

distichus was the most abundant and ubiquitous algal species in

the intertidal zone, and species of Alaria were notably absent

from the shore.

92 Dahlberg, M. L. 1979. History of fishery and summary

statistics of the sockeye salmon, Oncorhynchus nerka, runs

to the Chignik Lakes, Alaska, 1888-1966. NOAA (Natl.

Oceanic Atmos. Adm.) Tech. Rep. NMFS (Natl. Mar. Fish.

Serv.) SSRF (Spec. Sci. Rep. Fish.) 735, 16 p.

Annual runs of sockeye salmon to the Chignik Lakes, Alaska,

decreased from an average of 1.9 million during the period

1922-39 to an average of 0.9 million during the period 1949-66.

In order to study the dynamics of the runs' historic catch,

escapement and age structure data were compiled by spawning stock

and brood year. This history of fishing and management of the

runs from inception of the fishery until 1966 is described. The

high seas and coastal distributions of Chignik sockeye salmon

indicated significant interception by the fishery in only one

area other than the Chignik Bay and Chignik Lagoon; the fishery

at Cape Igvak started in the mid-1960's. Results of the study

were used to construct parent-progeny relationships that formed

the basis for a management strategy to restore the runs to former

level of abundance.

93 Dahlberg, M. L. 1973. Stock-and-recruitment relationships

and optimum escapements of sockeye salmon stocks of the

Chignik Lakes, Alaska. Rapp. P.-V. Reun. Cons. Penn. Int.

Explor. Mer 164:98-105.

The Chignik River system of western Alaska offers opportunity to

formulate a model management strategy for enhancement of sockeye

salmon runs based on precise harvest regulation. This is
67

possible because the system is segregated from other similar

systems and accurate statistics on catch and escapement are

available for a 38 year period.

By statistically studying spawner-return relationships, a

model of spawner-returns was derived for Chignik Lake and Black

Lake stocks, the two contributors to the Chignik River system,

and for the combined stocks. Alternative models were examined.

Optimum escapements were estimated for each stock and for

combined stocks. Results indicate that higher yields can be

realized by fishing each stock at its optimum rate of

exploitation rather than harvesting a mixture of stocks at a

common rate of exploitation.1

94 Dahlberg, M. L. 1968. Analysis of the dynamics of sockeye

salmon returns to the Chignik Lakes, Alaska. Ph.D. Thesis,

Univ. Washington, Seattle, 337 p.

Records of the abundance and age structure of sockeye salmon

(Oncorhynchus nerka (Walbaum)) returns to the Chignik Lakes,

Alaska, were studied to determine factors responsible for the

marked decrease of the populations over the past 50 years. The

average abundance of the stocks has dwindled in recent years to

less than one-half the average abundance during the period from

1922 to 1939. Based on the tacit assumption that a potential for

increase still exists in the populations, a plan for improved

management of the stocks is proposed; prerequisites for the plan

are adopted as specific objectives of the study.

The prerequisites of the plan for rebuilding the stocks

required (1) compilation of the historic records of catch,

escapement and age composition of the runs since 1888,

(2) determination of the relationship between the abundance of

mature progeny and size of parental spawning stocks and

(3) determination of the most accurate methods of forecasting the

magnitude and time of return of the yearly runs of sockeye to

Chignik.

Density-dependent and density-independent factors possibly

related to the decrease in abundance are presented and discussed.

A differential decrease in the abundance of the two stocks is

attributed tentatively to increase density dependent mortality in

one stock, probably a result of ecological changes in the nursery

lake brought about by consistent underescapement. The proposed

management strategy for increased production of Chignik sockeye

requires increased escapement levels, precisely regulated, to the

two nursery lakes. This should result in ecological changes

which should enhance the production of sockeye salmon.

95 Frame, G. W. 1976. Alaska's fishing black bear. Pac.

Discovery 29(3):19-25.

The author spent one summer observing black bear behavior at

Olsen Creek which flows into Olsen Bay in Prince William Sound,

Alaska. Specifically, he observed behavior of 18 individually

identified black bears fishing and eating chum and pink salmon.

Although both species were available in roughly equal numbers,

the preferred target was chum salmon (possibly because they are
68

slower and larger). Three types of fishing and eating behavior

were identified (in order of frequency of occurrence)--(l) active

fishing for chums and preference for females with eggs;

(2) feeding on dead salmon caught by other bears; and,

(3) feeding on salmon that had spawned out and died from natural

causes. The eggs were the preferred part of the fish, with the

head and nape areas of the fish following in preference. The

author estimated that black bears killed 8% of the salmon before

they had an opportunity to spawn; therefore, they did not pose a

serious threat to population maintenance.1

96 Frame, G. W. 1974. Black bear predation on salmon at Olsen

Creek, Alaska. Z. Tierpsychol. 35:23-38.

Activities of black bears (Ursus americanus) were observed on a

stream used for spawning by chum salmon (Oncorhynchus keta) and

pink salmon (0. gorbuscha) at Olsen Bay, Prince William Sound,

Alaska, during the summer of 1967. Black bears were most active

for 2 hours beginning at dawn and for several hours before and

after dusk. The most common activity was eating salmon killed

and abandoned by other black bears. An average of seven attempts

was made for each salmon captured and retained. Black bears

preferred eggs over the rest of the carcass and were able to

recognize live unspawned female salmon. Although equal numbers

of male and female salmon were captured, twice as many females

were retained;

when captured.

65% of these females were relatively unspawned

Black bears captured about 8% of the female

salmon population before they spawned. There was no evidence of

fishing territories.

97 Harry, G. Y. 1964. The shrimp fishery of Alaska. Proc.

Gulf Caribb. Fish. Inst. 16:64-71.

Until 1958, the shrimp fishery of Alaska was confined to

southeastern Alaska where from 1 to 2 million pounds of

hand-picked shrimp were packed annually for more than 30 years.

In 1958 the shrimp fishery expanded westward, with new plants in

Seldovia, Kodiak, and Seward. Alaska has the potential for a

much larger shrimp fishery than exists at present. Four species

of shrimp are of commercial importance; Pandalus borealis,

P. platyceros, P. hypsinotus, and Pandalopsis dispar, with pink

shrimp (P. borealis) being of greatest importance.

Shrimp fishing by foreign powers in water off Alaska has

been primarily in the Bering Sea. The initial Japanese effort in

1961 took about 19 million pounds of shrimp, and in 1962,

40 million pounds were taken. In 1963, the U.S. Bureau of

Commercial Fisheries received reliable reports of Soviet shrimp

fishing in the Bering Sea.

98 Haynes, E. 1983. Distribution and abundance of larvae of

king crab, Paralithodes camtschatica, and Pandalid shrimp in

the Kachemak Bay area, Alaska, 1972 and 1976. NOAA (Natl.

Oceanic Atmos. Adm.) Tech. Rep. NMFS (Natl. Mar. Fish.

Serv.) SSRF (Spec. Sci. Rep. Fish.) 765, 64,p.
69

Distribution and abundance of larvae of king crab, Paralithodes

camtschatica, northern shrimp,

P. goniurus,

Pandalus borealis, humpy shrimp,

coonstripe shrimp, P. hypsinotus, and sidestripe

shrimp, Pandalopsis dispar, were studied in the Kachemak Bay

area, Alaska, in 1972 and 1976. In both 1972 and 1976, larvae of

king crab, northern shrimp, and humpy shrimp first appeared in

outer Kachemak Bay; their abundance was greatest in the central

portion of the outer bay. Two additional species were studied in

1972, coonstripe shrimp and sidestripe shrimp. In 1972, the

center of abundance of sidestripe shrimp larvae was similar to

that of larvae of king crab, northern shrimp, and humpy shrimp.

Coonstripe shrimp larvae were most abundant in the inner bay and

along the northern shore of the outer bay.

The direction in which larvae were transported out of outer

Kachemak Bay was only in partial agreement with suspected

water-current patterns and may have been influenced by behavior

of the larvae. Continued abundance of larvae in outer Kachemak

Bay may be caused by entrainment of the larvae in gyres.

Depending on species and area, pandalid shrimp larvae are

released at different times and over different periods. For

example, larvae of northern shrimp appeared in plankton catches

earlier than larvae of humpy shrimp. Coonstripe shrimp had the

longest release period of all the shrimp studied.

From the percentage of glaucothoe in the samples, king crab

larvae probably settled in the Bluff Point area in outer Kachemak

Bay. Larvae of pandalid shrimp probably settle in outer Kachemak

Bay and possibly lower Cook Inlet, but exact locations cannot be

determined only by observing changes in morphology of the larvae.

Vertical depth distributions of larvae of king crab and

pandalid shrimp were generally similar.

king crab,

Early-stage larvae of

northern shrimp, and humpy shrimp migrated vertically

in a diel cycle. A thermocline did not prevent migration to

surface waters.

99 Haynes, E. 1981. Description of stage II zoeae of snow

crab, Chionoecetes bairdi, (Oxyrhyncha, Majidae) from

plankton of lower Cook Inlet, Alaska. Fish. Bull., U.S.

79:177-182.

Chionoecetes bairdi Rathbun 1924 (subfamily Oregoniinae) is the

only species of snow crab (genus Chionoecetes) that occurs in

Cook Inlet, Alaska, and contributes about 20% of the total value

of the commercial fisheries harvest of the area. The larval

stages of C. bairdi consist of one prezoeal, two zoeal, and one

megalopal stage. The zoeae are readily distinguished from known

zoeae of other genera of the subfamily Oregoniinae (Hyas and

Oregonia) by size (the zoeae of C. bairdi are nearly twice as

large as zoeae of Hyas and Oregonia) and by slight differences in

morphology, especially setation of the antennule and length of

the posterior lateral spines. The prezoeae, Stage I zoeae, and

megalopa of C.

In this report,

bairdi have been described from known parentage.

I describe the Stage II zoeae from plankton and

compare them with other known Oregoniinae zoeae from the North

Pacific Ocean.
70

100 Haynes, E. 1980. Stage I zoeae of a crangonid shrimp,

Crangon franciscorum angustimana, hatched from ovigerous

females collected in Kachemak Bay, Alaska. Fish. Bull.,

U.S. 77:991-995.

Information on the larval stages of crangonid shrimp of the North

Pacific Ocean is meager. Needler (1941) described the first

zoeal stage of Crangon septemspinosa (as Crago septemspinosus

Say) hatched in the laboratory from ovigerous females and the

remaining four zoeal stages from plankton collected near Prince

Edward Island, Canada. Kurata (1964) described the larval stages

of c. affinis de Haan and various larval stages of six,

unidentified Crangon spp. from Japanese waters. He obtained the

first zoeal stage of C. affinis from known parentage, but the

remaining stages were collected from plankton. Markarov (1967)

briefly described larvae of C. dalli Rathbun and C. septemspinosa

(Say) which were collected from plankton along the western

Kamchatka shelf. He suggested that C. dalli was an analog of

C. allmani Kinahan and C. septemspinosa was an analog of

C. crangon (Linnaeus). Crangon allmani and C. crangon are

eastern Atlantic species. He assumed that the C. affinis larvae

described by Kurata (1964) was actually larvae of

C. septemspinosa. Loveland (1968) described larvae of

C. alaskensis Rathbun reared in the laboratory from females

collected near Anacortes, Washington.

Morphology of Stage I larvae is closely related to Caridean

development and can be used to estimate the number of larval

stages, classify species, categorize larvae for identification

purposes, and identify subsequent larval stages (Needler 1938;

Pike and Williamson 1961, 1964; Kurata 1964; Ivanov 1971; and

others). In this report I describe and illustrate the first

zoeal stage of C. franciscorum angustimana Rathbun from ovigerous

females and compare these zoeae with Stage I zoeae of crangonids

described by other authors. Also, I show that the criterion of

the absence of exopodites on the second pair of pereopods for

distinguishing larvae of Crangon from other genera of the

Crangonidae is invalid for Crangonidae of the North Pacific

Ocean.

101 Haynes, E. 1979. Description of larvae of the northern

shrimp, Pandalus borealis, reared in situ in Kachemak Bay,

Alaska. Fish. Bull., U.S. 77:157-173.

Northern shrimp, Pandalus borealis, were reared in situ in

Kachemak Bay, Alaska, from Stage I (first zoeal) through Stage

VIII (second juvenile). Each of the six larval stages and first

juvenile stage is described and illustrated, and a brief

description is given for the second juvenile stage. Apparently

larvae of P. borealis in Alaska waters have at least one less

stage than larvae of P. borealis in either British Columbia,

Greenland, or Japan waters. Of the known larvae of the North

Pacific Ocean, larvae of P. borealis are most similar

morphologically to larvae of P. goniurus but are separable from

them by being slightly larger in size and, in zoeal Stages I-III,

by bearing more setae on certain appendages, particularly the

antennal scale and certain mouth parts. From Stage IV to
71

megalopa, the rostrum of P. borealis has more dorsal teeth, the

second pereopods are more developed, and the pleopods are fringed

with more setae than for larvae of P. goniurus. The criterion of

the lack of an outer seta on the maxillule for distinguishing

zoeae of Pandalus from certain other Caridea is shown to be

invalid.

102 Haynes, E. 1970. Age and growth of giant Pacific sea

scallops in the Gulf of Alaska. Proc. Natl. Shellfish.

Assoc. 60:14. (Abstract only).

Giant Pacific sea scallops, Patinopecten caurinus, in the Gulf of

Alaska have a typically sigmoid-shaped growth curve with an

inflection point at about 3 years of age or 55 mm shell height.

Increase in shell height decreases thereafter, tending toward an

upper asymptote that varies with geographical area. Sea scallops

grow fastest off Kodiak Island and slowest off southeastern

Alaska; between these areas their growth rate is intermediate.

Within-age variability in shell height is greatest in scallops

2-4 years old and is much reduced and nearly constant thereafter,

suggesting the presence of a compensating growth mechanism which

limits variability in growth as scallops get older. Causes for

the differences in scallop growth among the different areas have

not been established but may be related to certain environmental

conditions.

103 Haynes, E. B., and J. C. McMullen. 1970. Relation between

meat weight and shell height of the giant Pacific sea

scallop, Patinopcten caurinus, from the Gulf of Alaska.

Proc. Natl. Shellfish. Assoc. 60:50-53.

Meat weight-shell height relations were obtained for giant

Pacific sea scallops, Patinopecten caurinus, collected from two

areas in the Gulf of Alaska.

shell height in mm.

W is meat weight in g and L is

Over the range of shell height; examined, the mean weight of

scallop meats differed significantly (1% level) between the

areas.

104 Heard, W. R., R. L. Wallace, and W. L. Hartman. 1969.

Distributions of fishes in fresh water of Katmai National

Monument, Alaska, and their zoogeographical implications.

U. S. Fish Wildl. Serv. Spec. Sci. Rep. Fish. 590, 20 p.

Katmai National Monument covers 10,916 km on the base of the

Alaska Peninsula and is divided by the Aleutian Mountain Range

into two principal drainage areas. Streams north of the Aleutian

Range flow into Bristol Bay of the Bering Sea, and those south of

the mountains flow into Shelikof Strait of the North Pacific

Ocean. The large multilake Naknek River system is the dominant

drainage area on the Bristol Bay side of the monument, whereas

small single lakes and short streams and rivers constitute many
72

separate drainages on the Shelikof Strait side. Twenty-four

species of fish occur in the Bristol Bay drainages of the

monument, but only eight species were collected in streams and

lakes draining into Shelikof Strait. Evidently the Aleutian

Range has been a barrier to the southward movement of freshwater

fishes in the monument. All eight species in Shelikof Strait

drainages are capable of dispersal through salt water, whereas

several forms in Bristol Bay drainages require fresh water for

dispersal. Variable numbers of species occur in the

interconnecting lakes of the Naknek River system. Naknek Lake,

the downstream terminus of the lake system, contains 24 known

species and each upstream lake contains fewer species than the

one into which it drains. The present distribution of fishes in

this system is discussed in terms of the sequential timing of

species invasion and the postglacial development of barriers.

105 Helle, J. H. 1989. Relation between size-at-maturity and

survival of progeny in chum salmon, Oncorhynchus keta

(Walbaum). J. Fish Biol. 35(A):99-107.

Data on age- and size-at-maturity, growth, and abundance of chum

salmon were collected from 1959 to 1977 at Olsen Creek in Prince

William Sound, Alaska. Age composition of spawners (3 to

6-year-olds) varied from year to year: 4-year-old fish were the

dominant age group in most (16 out of 19) years and 6-year-old

fish usually represented less than 1% of the returns. Mean size

of older spawners was significantly larger than that of younger

spawners. Size-at-maturity was similar among fish from different

broods maturing at different ages in the same year.

Size-at-maturity and survival of progeny were significantly

related. The larger the mean size of spawners, the higher the

survival rate to adulthood of their progeny. Possible reasons

for this relationship are discussed.

106 Helle, J. H. 1984. Age and size at maturity of some

populations of chum salmon in North America. In P. A.

Moiseev (editor), Proceedings of the Pacific Salmon Biology

Conference, Yuzhno-Sakhalinsk, USSR, 3-13 October 1978,

p. 126-143. Pacific Scientific Institute of Fisheries and

Oceanography (TINRO),

Vladivostok, USSR.

4 Shevchenko Alley 690600,

In North America, chum salmon, Oncorhynchus keta, spawn in

watersheds from the Mackenzie River in northern Canada southward

to northern California, U.S.A. (Bakkala 1970). However, only

small spawning populations exist north of Kotzebue Sound, Alaska,

U.S.A., or south of the central Oregon coast in the United

States.

Published records of size and age of chum salmon at maturity

in North America have been sporadic (see Marr 1943; Henry 1954;

Thorsteinson, Noerenberg, and Smith 1963; Bakkala 1970).

Further, most of the reported observations described size and age

at maturity for commercial catches of mixed stocks.

Comprehensive records of size and age of individual spawning

populations of chum salmon are very limited.
73

In 1959, I began sampling the spawning population of chum

salmon at Olsen Creek, a small coastal stream in Port Gravina,

Prince William Sound, Alaska, for size and age throughout the

spawning season. The objectives of this study were to

(1) describe the variation in size and age composition of chum

salmon during a spawning season (intraseasonal), and (2) describe

the variation in size and age of spawning chum salmon in

different years (interseasonal).

In 1971-78, I sampled several other spawning populations of

chum salmon in different geographical areas--Noatak River, near

Kotzebue, Alaska; Chilkat River, in northern southeastern Alaska

near Haines; Disappearance Creek, in Cholmondeley Sound on Prince

of Wales Island near Ketchikan, Alaska; Fish Creek, at the head

of Portland Canal near Hyder, Alaska; Quilcene Hatchery, at

Walcott Slough on Hood Canal near Brinnon, Washington; and

Whiskey Creek, in Netarts Bay, Oregon. The chum salmon run at

Walcott Slough is maintained by artificial propagation by the

U.S. Fish and Wildlife Service, Quilcene National Fish Hatchery.

The run at Whiskey Creek is maintained by natural spawning and an

experimental hatchery operated by Oregon State University. The

objectives of these additional observations were to compare size

and age at time of spawning of chum salmon that were

geographically and ecologically diverse.

107 Helle, J. H. 1982. Some effects of the marine environment

on age at maturity and growth of chum salmon in Prince

William Sound, Alaska. In B. R. Melteff and R. A. Neve

(editors), Proceedings of the North Pacific Aquaculture

Symposium, p. 91. (Abstract only.) Univ. Alaska,

Fairbanks, Sea Grant Rep. 82-2.

Influence of the marine environment on age and size at maturity,

early marine growth, and abundance of chum salmon, Oncorhynchus

keta, from Olsen Creek in Prince William Sound, Alaska, was

studied during the years 1959 to 1978 (Helle, 1979).

Age composition of the spawners returning to Olsen Creek

varied from year to year, but they were predominately 3-, 4-, and

5-year fish. Some 6-year fish returned between 1968 and 1975,

but this age group usually did not represent more than 1 percent

of the returns. Mean age composition for the brood years 1956 to

1972 for males was 15,

fish, respectively.

66, and 19 percent for 3-, 4-, and 5-year

Mean age composition for females of the same

broods showed higher percentages of older fish: 9, 67, and

23 percent for 3-, 4-, and 5-year fish, respectively.

Interseasonally, mean age at maturity increased as number of

fish in a brood increased. Intraseasonally, age of new chum

salmon spawners at Olsen Creek decreased as the season

progressed. Mean size of older spawners was significantly larger

than the mean size of younger spawners, but the ranges in size of

the three age groups overlap each other so size is not a good

criterion for estimating age of chum salmon.

Numbers of circuli and distances-between annuli on adult

scales were used to estimate growth of chum salmon during their

first two years at sea. Growth during the first year at sea was

related to sea-surface temperatures and marine weather parameters

in Prince William Sound and in the northern Gulf of Alaska. 
74

Growth during the first year at sea was not significantly related

to age at maturity; however, growth during the second year at sea

was negatively related to age at maturity.

Size at maturity was related to sea-surface temperatures and

marine weather parameters in the northern Gulf of Alaska and

Prince William Sound during the year of return. Fluctuations in

size at maturity were more similar between fish from different

broods returning during the same year than between fish from the

same broods maturing at different ages. A highly significant

relationship was found between survival of progeny and mean size

of parents.

108 Helle, J. H. 1981. Significance of the stock concept in

artificial propagation of salmonids in Alaska. Can. J.

Fish. Aquat. Sci. 38:1665-1671.

Hatcheries could make a significant contribution to the

enhancement of salmonid fisheries if application of this

technology is based on principles of the stock concept. Evidence

for the stock concept is reviewed and discussed in relation to

transplantation of stocks, mating procedures, maintenance of

genetic diversity, different methods of artificial propagation,

and location of artificial propagation facilities. A strategy is

developed for management of both wild and artificially propagated

stocks.

109 Helle, J. H. 1979. Influence of marine environment on age

and size at maturity, growth, and abundance of chum salmon,

Oncorhynchus keta (Walbaum), from Olsen Creek, Prince

William Sound, Alaska. Ph.D. Thesis, Oregon State Univ.,

Corvallis, 118 p.

Effects of the marine environment on age and size at maturity,

early marine growth, and abundance of chum salmon, Oncorhynchus

keta, were studied at Olsen Creek during 1959-77.

Chum salmon return to Olsen Creek as predominately 3-, 4-,

and 5-year fish; however, age composition varied from year to

year. The mean age composition for the brood years 1956-72 for

males was 15%, 66%, and 19% for 3-, 4-, and 5-year fish,

respectively. Mean age composition for females of the same

broods showed slightly higher percentages of older fish: 9%,

67%, and 23% for 3-, 4-, and 5-year fish, respectively. Some

6-year chum salmon returned to Olsen Creek between 1968 and 1975;

but, only in 1973 did the number of 6-year fish (3%) represent

more than 1% of the returns. Population sizes tended to be

larger during these years, and mean age increased as the number

of fish in a brood increased. Intraseasonally, age of new chum

salmon spawners at Olsen Creek decreased as the season

progressed. Mean size of older spawners was greater than the

mean size of younger spawners; but, the ranges in size of the

three age groups overlap each other so size is not a good

criterion for estimating age of chum salmon.

Measurement of circuli and distances on adult scales were

used to estimate growth of chum salmon during their first two

years of marine life. Both number of circuli and distances on
75

scales of juvenile chum salmon after their first summer in Prince

William Sound were shown to be related to length of the fish.

Growth during the first season at sea was not related to age at

maturity; however, amount of growth acquired during the second

marine season was negatively related to age at maturity. Growth

during the first summer at sea was related to sea surface

temperatures and marine weather parameters in Prince William

Sound in the northern Gulf of Alaska. Location of chum salmon

from Olsen Creek during their second year at sea is unknown.

Fluctuations in size (length) at maturity were more similar

between fish from different broods returning during the same year

than they were for fish that matured at different ages from the

same broods. Length at maturity was related to marine weather

factors during their last summer at sea in the northern Gulf of

Alaska and Prince William Sound and also mean summer sea surface

temperature in Prince William Sound during the year of return.

Total survival of each brood was estimated from the ratio of

number of progeny (returns) to number of parents (spawners). No

direct relationships were found between survival and growth

during the first or second season in the sea, sea surface

temperatures, or upwelling indices along the coast. However, a

highly significant relationship was found between the survival of

progeny and mean length of the parents.

110 Helle, J. H. 1970. Biological characteristics of

intertidal and fresh-water spawning pink salmon at Olsen

Creek, Prince William Sound, Alaska, 1962-63. U.S. Fish

Wildl. Serv. Spec. Sci. Rep. Fish. 602, 19 p.

Prince William Sound is unique among major pink salmon-producing

areas in that a significant portion of the spawning takes place

in the intertidal zones of streams. Olsen Creek is one of the

major spawning streams in the Sound.

The percentage of fines (solids passing through an 0.833 mm

sieve) in spawning-bed materials increased progressively from

higher to lower intertidal levels, i.e., higher intertidal levels

contained coarser spawning gravel than lower levels.

Although less than one-third of the spawning area available

in the Olsen Creek drainage is subject to tidal influence,

70 percent of the total pink salmon spawners occupied this area

in 1962 and 30 percent in 1963. Late-run fish of the even-year

line spawned only in the intertidal area; fish of both the early

and late runs of the odd-year line spawned in both the intertidal

and fresh-water areas. The size of the spawning populations was

estimated by a repetitive stream survey technique, which is

described and compared with the three methods used in 1960 and

1961.

The length of pink salmon was compared between sexes,

between spawning areas, and between times of spawning. Fish in

the even-year line that spawned in the small intertidal creeks

tended to be smaller than those that used the main stream, but in

the odd-year line this difference was confined to females. The

mean lengths of females were about the same in 1962 and 1963, but

females from the odd-year line were more fecund. In both years a
76

significant positive correlation was shown between lengths of

females and numbers of eggs.

111 Helle, J. H. 1966. Behavior of displaced adult pink

salmon. Trans. Am. Fish. Soc. 95:188-195.

Studies to show whether pink salmon Oncorhynchus gorbuscha

(Walbaum) will repeat their original choice of spawning area when

captured and transported to another location were done in Olsen

Creek, Prince William Sound, Alaska in 1961, 1962, and 1963.

Fish to be displaced were anesthetized with MS-222 (except in

1961 when no anesthesia was used) and marked with Petersen disk

tags. Equal numbers of control fish were treated in an identical

manner and left at the donor site.

In 1961, 100 pink salmon were displaced from Olsen Creek

beyond the entrance of the bay, a distance of more than 3 miles

from the creek mouth. Of the 54 fish accounted for, 91% returned

to Olsen Creek.

In 1962 and 1963, the displacements were confined to Olsen

Bay. Fish displaced "on-route" achieved a higher rate of return

than fish displaced "off-route". Further, under conditions of

extremely low stream flow, displaced fish did not return to their

original choice of spawning area. When stream flows were normal

or high, they showed a strong tendency to return to areas of

their original choice.

112 Helle, J. H. 1960. Characteristics and structure of early

and late spawning runs of chum salmon, Oncorhpchus keta

(Walbaum), in streams of Prince William Sound, Alaska. M.S.

Thesis, Univ. Idaho, Moscow, ID, 53 p.

Due to a poor escapement of pink salmon in 1957 the commercial

fishery for pink and chum salmon was closed in 1959. This

closure provided an opportunity to observe on the spawning

grounds the total return of chum salmon to Prince William Sound

in 1959. This study describes the abundance, distribution, age,

size, and fecundity of chum salmon that spawned in some of the

major streams along the eastern shore of Prince William Sound

between Valdez Arm and Orca Inlet.

113 Helle, J. H., R. S. Williamson, and J. E. Bailey. 1964.

Intertidal ecology and life history of pink salmon at Olsen

Creek, Prince William Bound, Alaska. U.S. Fish Wildl. Serv.

Spec. Sci. Rep. Fish. 483, 26 p.

Intertidal spawning of pink salmon is of major importance in

Prince William Sound. Studies were initiated at Olsen Bay in

1960 to ascertain how much these intertidal spawners contributed

to the total production of pink salmon.

Olsen Creek is inundated with tidewater about 80 percent of

the time at the 3-foot tide level and about 7 percent of the time

at the 11-foot level. Saline water was shown to penetrate the
77

gravel at redd depth during high tides. The highest

concentration at the 11-foot tide level was 9.3‰ during a

14.5-foot tide. Temperature changes of up to 10o

F, would occur

within 1 hour at evaluations up to the 8-foot level on floodtide.

The occurrence of spawners in 1960 and 1961 was bimodal;

however, in 1960 the late run utilized only the intertidal

spawning area, while in 1961 the late run utilized both the

intertidal and fresh-water areas.

run spawned in both environments.

During the 2 years the early

In 1960, 98,574 pink salmon

spawned in Olsen Creek and 1961, 135,905 spawned. During both

years 74 percent of the total run spawned in the intertidal

portion of the stream.

Temporal and spatial distribution of spawners, size

differences, and seasonal changes in sex ratios provide evidence

for the existence of discrete spawning groups or races.

Live egg densities and survival over winter to the

preemergent fry stage were progressively greater from the lower

to the higher levels in the intertidal area. Overwinter survival

between egg and fry stages below the 4-foot level was 0.

Survival at the 7- to 9-foot level and the 10- to 11-foot level

was 20 and 54 percent respectively.

114 Hubbard, J. D. 1971. Distribution and abundance of

intertidal invertebrates at Olsen Bay in Prince William

Bound, Alaska, one year after the 1964 earthquake. In The

Great Alaska Earthquake of 1964: Biology, p. 137-157.

National Academy of Sciences, Washington, DC.

In the summer of 1965, the Bureau of Commercial Fisheries began a

study of intertidal invertebrates in the Olsen Bay area of Prince

William Sound; this included an evaluation of the effect on

intertidal organisms of uplift caused by the 1964 Alaska

earthquake. Intensive sampling of four areas representative of

the three most common types of habitat in this protected bay

revealed a vertical distribution and abundance of invertebrates

species strongly influenced by substrate composition and tidal

exposure. Mean tide level (+6.2 ft) exhibited particular

importance as the upper limit of numerous organisms. It was

found that the uplift produced measurable changes. in the

distributions of certain intertidal organisms. The amount of

uplift, approximately 3.0 to 3.5 ft in Olsen Bay, could be

estimated by comparing the positions of some of these organisms

before and after the earthquake. The most obvious and reliable

quantitative index of the uplift was provided by pre- and

postearthquake limits of barnacle populations, whereas supportive

evidence was obtained from examination of certain bivalve mollusk

distributions. Differential mortality, noted especially between

different species of mollusks and between different age-classes

of certain species, was apparently attributable to

earthquake-related processes. While reproductive success of some

species appeared unaffected by the earthquake, other species have

apparently experienced little reproduction since the earthquake.
78

115 Kirkwood, J. B. 1962. Inshore-marine and freshwater life

history phases of the pink salmon, Oncorhynchus gorbuscha

(Walbaum) and the chum salmon 0. keta (Walbaum) in Prince

William Bound, Alaska. Ph.D. Thesis, Univ. Louisville,

Louisville, KY, 300 p.

The salmon resources are of great economic importance to the

people of Alaska. Pink and chum salmon are most abundant along

the coast of central and southeastern Alaska. Numbers of pink

and chum salmon returning to Prince William Sound each year has

generally declined since 1943. The current study was designed to

determine the causes of this decline. Since it seemed likely

that the major limiting factors probably occurred while the

salmon were in fresh water, emphasis was placed on studying this

phase of the life histories of both species.

The 194 streams listed as spawning streams were stratified

according to the annual abundance of pink salmon spawners and

time of spawning during the 10 years prior to 1957. Aerial

surveys were used to determine the sizes of spawning populations

so that these populations could be correlated with fry production

and subsequent adult returns. Surveys were conducted at

12 streams during 1957 and 40 streams during 1958 and 1959.

During 1958, 6,273 adult chum salmon were sampled to study

the age, length and sex composition of the spawning populations

in Prince William Sound. Larval development and mortality were

studied during the fall and winter of 1958-1959 to determine the

time and causes of mortality of the salmon while they were in the

gravel. Fry-migration studies consisted in capturing a portion

of the migrating fry in each study stream and intertidal area as

they moved from the gravel to the sea.

Larger populations of pink salmon spawned in study streams

during 1958 than in either 1957 or 1959, and a greater percentage

of the spawners utilized the intertidal zones during 1958.

The survival of eggs and larvae in intertidal and stream

zones is discussed and the survival is correlated with air and

water temperatures in one stream. Other possible mortality

factors are explored.

Population estimates of fry migrations in stream zones of

study streams, categories of streams, and Prince William Sound

are presented. The survival of fry during migration to sea is

greatly affected by predators, particularly birds and other fish.

A discussion is given of the recruitment of fry correlated

with parent spawners. Points included in this discussion include

the success of fry production resulting from even- and odd-year

spawning populations and the success of reproduction in

intertidal and stream zones.

116 Kirkwood, J. B., and R. M. Yancey. 1965. Effects of the

March 27 earthquake upon the shellfish resources of Alaska.

Proc. Alaska Sci. Conf. 15:162.

Between June 8 and June 24, 1964, a general survey was conducted

to learn the effects of the March 27, 1964, earthquake upon

shellfish populations and their habitats. Fishermen, packers,
79

state management personnel, and others were visited in Cordova,

Anchorage, Seldovia, Homer, Kasitsna Bay, Kodiak, Olga Bay, Sand

Point, and Squaw Harbor. Razor clam beaches also were visited at

Cordova, Ninilchik, Poly Creek, and Swickshak. The area affected

by the earthquake has been estimated at more than 30,000 square

miles. More than 22,000 square miles located east of the

tectonic hinge-zone subsided and at least 12,000 square miles

located west of this zone uplifted. The major effect on

shellfishes occurred near Cordova. Razor clam beaches rose about

six feet in the Copper River area; butter clams (Saxidomus

nuttallii) and cockles (Clinocardium nuttallii) were found dead

on exposed beaches in Orca Inlet after the quake; and submarine

landslides may have caused mortality to shellfishes in western

Prince William Sound where several square miles of dead rockfish

(Sebastodes sp. and Sebastolobus sp.) and cod (Gadus sp.) were

observed on March 28, 1964. Other razor clam beaches were

probably not affected since these areas were not uplifted or

lowered greatly by the earthquake. Subsidence of the area which

includes Kasitsna Bay will reduce the availability of butter

clams there by about 80%.

king crab, shrimp,

There was no evidence of mortality of

and Dungeness crab or major changes in their

habitats except possibly in the Prince William Sound area.

117 Leatherwood, S., A. E. Bowles, E. Krygier, J. D. Hall, and

S. Ignell. 1984. Killer whales (Orcinus orca) in southeast

Alaska, Prince William Sound, and Shelikof Strait; A review

of available information. Rep. Int. Whaling Comm.

34:521-530.

Information on killer whales is summarized from a formal

sightings network, 1976-1981 (Southeast Alaska), aerial surveys,

1976-1978 and small boat surveys, 1976-1982 (Prince William

Sound), aerial surveys, 1982-1983 (Shelikof Strait), and

interviews with marine scientists, fishermen and other

knowledgeable mariners (all areas). Although present year-round,

killer whales in each area increase in numbers and concentrate in

specific areas in late summer through early fall, apparently in

response to concentrations of salmon. In Prince William Sound,

one group apparently remains within a limited home-range (less

than 25 km radius) in Knight Island Passage each summer. Minimum

counts from portions of the three areas are 93, 80 and 66 whales,

respectively. Data indicate concurrent presence of killer whales

in other unstudied portions of all three areas; so, 'populations'

in each area certainly number in excess of 100 animals. Herds

include an average of 4-6 animals (n=890) in Southeast Alaska.

Of the three areas surveyed, Prince William Sound and Southeast

Alaska appear most suitable as areas for long-term observational

studies of killer whales of the sort pioneered in inland waters

of Washington and British Columbia.

118 Lindstrom, S. C., and N. I. Calvin. 1975. New records of

marine red algae from the Gulf of Alaska. Syesis 8:405-406.
80

The documented ranges of eight species of red marine algae are

extended into the northeast Gulf of Alaska as a result of

intertidal sampling done in 1974 as part of baseline surveys of

area continental shelf baseline studies conducted in anticipation

of oil development. The extensions are as follows: Erythrocladia

subintegra --found at Anchor Cove, not previously reported in

Alaska; Erythrotrichia carnea--found at Zaikof Bay, not

previously reported from Alaska; Farlowia mollis--Macleod Harbor,

previously reported northern limit as Dixon Entrance; Gigartina

exasperata --found near Ocean Cape, previously reported northern

limit as northern California; Nemalion elminthoides--MacLeod

Harbor, previous northernmost collection from Khaz Bay, Chichagof

Island; Botryoglossum farlowianum, Callophyllis violacea, and

Serraticardia macmillani --found in the drift zone at Middleton

Island; none had previously been reported north of Khaz Bay.1

119 Meehan, W. R. 1966. Growth and survival of sockeye salmon

introduced into Ruth Lake after removal of resident fish

populations. U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish.

532, 18 p.

Sockeye salmon (Oncorhynchus nerka) in three lakes on Afognak

Island, Alaska, were studied. Ruth Lake was treated with

rotenone to remove resident fish. Midarm Lake, which has no

salmon, and Little Kitoi Lake, which has a small run of sockeye,

were used as controls to compare survival and growth of

introduced fry and natural fry with survival and growth of

introduced fry in Ruth Lake. Other factors that might influence

sockeye production, such as plankton and bottom fauna, were also

considered.

In general, growth and survival of fry and biological

production were greater in the treated lake. Growth and survival

decreased as fry densities increased.

120 Meyer, R. M. 1968. The Dungeness crab fishery around

Kodiak, Alaska. Comm. Fish. Rev. 30(8-9):44-47.

Alaska seiners and power barges form the bulk of the fleet that

fishes for Dungeness crabs in Kodiak waters from May to October.

Crews normally number three men who may set, pull, and reset over

ten 30-pot strings a day in shallow, near-shore waters. The pots

are baited with herring, clams, or squid. As each pot is pulled,

the catch is placed in tanks filled with circulating sea water to

insure live delivery to the processing plants. The crabs are

usually butchered, cooked, and frozen at the plant and are

shipped south for further processing.

121 Moyle, Peter. 1966. Feeding behavior of the

Glaucous-winged Gull on an Alaska salmon stream. Wilson

Bull. 78:175-190.
81

The feeding behavior of the Glaucous-winged Gull was studied on

Olsen Creek, a salmon stream flowing into Prince William Sound,

Alaska. It was found that feeding took place in two distinct

situations: (1) on the banks of the stream on salmon carcasses

pulled up by gulls or bears and (2) in the stream itself, on

salmon eggs drifting with the current. The behavior patterns

centering around the defense of salmon carcasses appeared to be

very similar to the territorial displays of the breeding season

with these exceptions; actual territories did not exist, the

 carcasses were defended only by hungry birds, and the attacking

gull won more often than the defender in disputes over a carcass.

Upright Displays, Oblique-cum-Long-Call Displays, and Mew Call

Displays were all observed frequently during carcass defense.

The effectiveness of such behavior, however, tended to break down

in the presence of bears feeding on freshly caught salmon,

particularly when the salmon were females. The gulls also seemed

to exhibit preference for female over male carcasses of spawned

out salmon. Birds in immature plumage usually could not defend a

salmon carcass against adult gulls, although first year juveniles

had a certain immunity to attack by their unresponsiveness to

adult threat displays. Apparently, even adults cannot defend

themselves against other adult gulls while diving for drift eggs

in the stream. At the approach of a low-flying attacker, the

swimming gull must either fly up or be bowled over. If drift egg

feeding occurs in a riffle, however, the riffle can be defended

like a salmon carcass. Paddling, probably to stir up salmon eggs

caught in the surface gravel also occurred in the riffles.

122 O'Clair, C. E., and S. T. Zimmerman. 1987. Biogeography

and ecology of intertidal and shallow subtidal communities.

In D.W. Hood and S. T. Zimmerman, (editors), The Gulf of

Alaska: Physical Environment and Biological Resources,

p. 305-343. NOAA, U.S. Dep. Commer. Available

Superintendent of Documents, U.S. Gov. Print. Office,

Washington, DC 20402.

Recent studies of the natural shore communities of the Gulf of

Alaska provide a descriptive foundation for future work on the

primary factors that determine both geographical and local

distribution and abundance patterns for algal and invertebrates

populations. However, there is still a lack of experimental

evaluation to determine the role that physical disturbances,

gradients in physical regimes,

in determining these patterns.

and biological interactions play

Our analysis of both the biotic composition and the

zoogeographic affinities of those invertebrates of the major

phyla revealed no major biogeographical discontinuities between

Yakutat and the eastern Aleutian Islands. However, we found that

the intertidal flora and fauna of the western Aleutians (Amchitka

and Shermya Islands) differed markedly from the flora and fauna

of the eastern Gulf. The distribution of species among trophic

levels was similar between these two regions, but the western

Aleutians had more Asiatic and fewer North American species, and
82

had a greater proportion of endemic species of Mollusca,

Crustacea, and Echinodermata than were found in the eastern Gulf.

Physical disturbances were only of overriding importance in

controlling community structure at three of the 29 study sites.

Gradients in the regimes for salinity, turbidity, and exposure

altered both the community composition and the relative

abundances of intertidal species, such as Semibalanus balanoides

and Balanus glandula, which have a tolerance for a broad range of

values for these factors. Pisaster ochraceus and Evasterias

troschelli do not appear to play key roles in the organization of

intertidal communities in Alaska because Mytilus californianus is

rare there and M. edulis is vulnerable to the activities of other

predators and perhaps to physical disturbance as well.

When most intertidal species are lifted even slightly above

their upper vertical limits by land-level changes, they either

die or emigrate. This supports the contentions that the upper

limits of most intertidal organisms are physiologically

determined. The exceptions are Balanus glandula, Semibalanus

cariosus, and Chthamalus dalli which can survive uplift of nearly

1 m above their upper limits. It normally takes at least three

years for communities to redevelop to their former condition

after an uplift.

123 Olsen, J. C. 1973. Pandalid shrimp life history research

at Kachemak Bay, Alaska. Mar. Fish. Rev. 35(3-4):62-64.

Kachemak Bay, an arm of lower Cook Inlet, is the site of NMFS

field studies of Pandalus borealis, the most important commercial

shrimp species in Alaska. Research activities focus on shrimp

life histories, behavior, and population dynamics; effects of

fishing on shrimp stocks; annual fluctuations in shrimp stock

abundance; characteristics and ecology of shrimp habitat; and the

role of pandalid shrimp in the organic production system of the

North Pacific. Weekly sampling of the commercial shrimp catches,

begun in 1970, provides information on growth, recruitment, and

mortality of P. borealis and P. goniurus. Larval studies of

shrimp, begun in 1971, have focused on understanding time and

place of hatching, diurnal vertical distribution patterns,

survival estimates, and annual larval production. Identification

and description of larval stages will be accomplished through

laboratory cultures and field collections.1

124 Orsi, J. A., R. K. Gish, and B. L. Wing. 1991. Northern

range extensions of four nearshore marine fishes in Alaska.

Can. Field-Nat. 105:82-86.

Northern range extensions of Oxylebius pictus (Hexagrammidae),

Syngnathus leptorhynchus (Syngnathidae), Synchirus gilli

(Cottidae), and Citharichthys stigmaeus (Bothidae) are reported.

All four species were collected north of 60 N in Prince William

Sound in the Gulf of Alaska from April through June 1989. This

is the first published account of a verified specimen of

O. pictus in Alaska.
83

125 Phinney, D. E. 1970. Spawning ground catalog of the

Chignik River system, Alaska. U.S. Fish Wildl. Serv. Data

Rep. 41, 147 p.

All known information about the sockeye salmon runs and the

spawning grounds of the Chignik River system, Alaska, is

cataloged in this report. The system, which is composed of two

lakes, Chignik and Black, supports the largest run of sockeye

salmon on the south side of the Alaska Peninsula. The catalog

lists for each spawning stream or beach the name, location,

physical description, description of the sockeye salmon runs, and

a chronological listing of the spawning ground surveys.

126 Quast, J. C., and E. L. Hall. 1972. List of fishes of

Alaska and adjacent waters with a guide to some of their

literature. NOAA (Natl. Oceanic Atmos. Admin.) Tech. Rep.

NMFS (Natl. Mar. Fish. Serv.) SSRF (Spec. Sci. Rep. Fish.)

658, 47 p.

The authors list 432 species known to occur in Alaska water,

supplemented by 137 species that have been recorded from

neighboring waters and, in the authors' opinion, should be

considered when new collections are identified. Species entries

are annotated to include common names, recorded range, useful

references, localities represented by specimens in the collection

of the Auke Bay Fisheries Laboratory, and comments on taxonomy.

Recorded geographic ranges are extended for 26 species:

Ophidiidae--Spectrunculus radcliffei; Scorpaenidae--Sebastes

emphaeus, S. nigrocinctus, S. wilsoni; Cottidae--Eurymen gyrinus,

Gymnocanthus detrisus, G. pistilliger, Hemilepidotus zapus,

Icelus spatula, I. uncinalis, Myoxocephalus jaok, Nautichthys

pribilovius, Triglops scepticus; Agonidae--Agonopsis emmelane,

Aspidophoroides bartoni, Ocella verrucosa; Cyclopteridae--

Careproctus melanurus, C. rastrinus, Cyclopteropsis phrynoides,

Liparis bristolense, L. ochotensis, Paraliparis caudatus,

P. deani, Temnocora candida; Scombridae--Thunnus thynnus;

Pleuronectidae--Limanda proboscidea.

127 Reid, G. M. 1971.

of herring,

Age composition, weight, length, and sex

Clupea pallasii, used for reduction in Alaska,

1929-66. NOAA (Natl.

(Natl. Mar. Fish.

Oceanic Atmos. Admin.) Tech. Rep. NMFS

Serv.) SSRF (Spec. Sci. Rep. Fish.) 634,

25 p.

Sampling data from the reduction fisheries for herring, Clupea

pallasii, in southeastern Alaska (1929-66), Prince William Sound

(1937-58), Kodiak (1936-59) are summarized. The data include the

weight of the catches, the weight allowed by quota, and age

composition, average weight, average length, and sex ratios.
84

128 Sears, H. S., and S. T. Zimmerman. 1977. Alaska intertidal

survey atlas, 402 p. National Marine Fisheries Service,

Auke Bay Laboratory.

This atlas describes the intertidal habitat types from Yakutat to

Shumagin Pass to Cape Prince of Wales on the Bering Strait. All

surveys were made from low flying aircraft, with some ground

truthing. This work was an extension of a more detailed study at

specific sites along the Alaska coast. The flights were started

in summer 1975, but were interrupted by the crash and sinking of

the survey aircraft, and were completed in 1976. Data for all

beaches described in the atlas,

River,

except for Yakataga to Copper

were collected during the 1976 period.

129 Skud, B. E., H. M. Sakuda, and G. M. Reid. 1960.

Statistics of the Alaska herring fishery 1878-1956. U.S.

Fish Wildl. Serv. Bur. Comm. Fish. Stat. Dig. 48, 21 p.

Statistics of the herring fishery in Alaska are summarized in

this report for the years 1878 to 1956 insofar as detailed data

were available.

Catches are recorded by statistical areas in Southeastern

Alaska, Prince William Sound, and Kodiak and represent a revision

of previously published figures obtained by converting meal

production to pounds of fish. Products of the fishery and the

statistics of operation are summarized for all Alaska.

130 Tait, H. D., and J. B. Kirkwood. 1962. Estimating

abundance of pink and chum salmon fry in Prince William

Sound, 1957. U.S. Fish Wildl. Serv. Spec. Sci. Rep. Fish.

429, 21 p.

Salmon fry enumeration studies conducted on eight streams in the

Prince William Sound area of Alaska provided estimates of the

numbers of pink and chum salmon fry produced in streams of that

area in 1957. The studies were conducted to provide a basis for

predicting returns of adult salmon. Methods of deriving

estimates of fry production from trapping experiments and

excavations of pre-emergent fry in intertidal gravel were

developed. Timing and duration of migration of chum and pink

salmon fry were recorded, and recommendations were made for

future fry sampling programs.

131 Thorsteinson, F. V. 1965. Effects of the Alaska earthquake

on pink and chum salmon runs in Prince William Sound. Proc.

Alaska Sci. Conf. 15:267-280.

The earthquake that struck Alaska March 27, 1964, centered in

northwestern Prince William Sound and produced drastic

environmental changes in the spawning habitat of pink

(Oncorhynchus gorbuscha) and chum (0. keta) salmon runs homing to

streams of that area. Its total impact cannot be measured until
85

spring, 1965 when the survival of salmon eggs deposited in

uplifted or downwarped stream areas in 1964 can be determined.

In this report, some results of pre-earthquake ecological studies

on a Prince William Sound stream are presented, and their

implications regarding future abundance of pink salmon in view of

the earthquake-produced changes in spawning habitat are

discussed.

132 Thorsteinson, F. V. 1965. Aftermaths of the Alaska earthquake in Prince William Bound. Pac. Fisherman 63(6):10-11.

Prince William Sound pink salmon spawning streams were adversely

affected by the March 27, 1964, earthquake. Elevations of

226 streams used by 75 percent of the total population were

changed. Thirty seven percent of the fish in odd year runs and

56 percent of fish in even year runs were potentially affected.

In streams affected by subsidence, previously used intertidal

spawning areas were drowned, forcing fish to move upstream to

productive tide levels. In streams located in areas of uplift,

low intertidal streambeds were raised to levels that might be

useful for spawning but siltation led to low egg survival. Some

streams in uplifted areas proved unstable as they degraded or

moved laterally to form new flow patterns, and, as a result,

deposited eggs were sometimes lost. Eggs and developing embryos

in all streams may have been affected by compaction of spawning

beds. In general, egg and fry mortalities among pink salmon were

expected to be higher than average until streams stabilized and

digging activity of spawning salmon removed fine materials from

newly exposed spawning beds.1

133 Thorsteinson, F. V., J. H. Helle, and D. G. Birkholz. 1971.

Salmon survival in intertidal zones of Prince William Sound

in uplifted and subsided areas. In The Great Alaska

Earthquake of 1964: Biology, p. 194-219. National Academy

of Sciences, Washington, DC.

Large numbers of pink and chum salmon, spawned each year in

intertidal portions of streams in the Prince William Sound, were

affected by the Alaska earthquake of March 27, 1964. Detailed

studies of the effects of changes in land level were made at

Olsen Creek (uplifted about 4 ft), a stream for which a large

amount of pre-earthquake data existed. General studies were made

at three other streams-- two in areas that were uplifted 10 ft and

one in an area that subsided 6 ft. Despite relocation of

intertidal spawning beds in relation to tidal levels, the areas

occupied and the distribution by the runs of returning pink and

chum salmon were at approximately the same levels and in the same

proportions as before the earthquake. The first spawners

occupied upper intertidal areas, and those that came later

spawned at lower levels;

tide level.

very few fish spawned below the 6 ft

Survival of eggs was high in upper areas and lower

in downstream areas. Stream slope adjustment caused by changes

in elevation created unstable streambed conditions. Scour, fill,
86

and channel shifts caused mortalities to eggs and alevins. We

estimate that these secondary effects of the earthquake caused

the disappearance of 7.27 million pink salmon eggs and

1.1 million chum salmon eggs from Olsen Creek intertidal

streambeds during the summer and fall of 1965 and 0.54 million

pink salmon alevins and 0.43 million chum salmon alevins during

the 1965-1966 winter.

134 Thorsteinson, F. V., W. H. Noerenberg, and H. D. Smith.

1963. The length, age, and sex ratio of chum salmon in the

Alaska Peninsula, Kodiak Island, and Prince William Sound

area of Alaska.

Fish. 430, 84 p.

U.S. Fish Wildl. Serv. Spec. Sci. Rep.

Data on length, age, and sex ratio of chum salmon from the Alaska

Peninsula area from 1951 through 1957, the Kodiak Island area

from 1948 through 1951 and 1955 through 1957, and the Prince

William Sound area from 1952 through 1958 show that age and

length composition in these areas varied in a similar manner.

Lengths of fish in the 3-, 4-, and 5-year age classes overlapped

to such an extent that length was not a useful guide to age. The

average age composition for the combined samples was about

10 percent 3-year-olds, 75 percent 4-year-olds, and 15 percent

5-year-olds. Mean age decreased as the season advanced. The

percentage of males decreased slightly as the runs progressed.

135 Zimmerman, S. T., J. L. Hanson, J. T. Fujioka, N. I. Calvin,

J. A. Gharrett, and J. S. MacKinnon. 1978. Intertidal

biota and subtidal kelp communities of the Kodiak Island

area. In Environmental Assessment of the Alaskan

Continental Shelf.

Investigators,

Final Reports of Principal

Vol. 4, Biological Studies, p. 316-508. U.S.

Dep. Commer., NOAA. Available Arctic Environmental

Assessment Center, 222 W. 8th Ave., No. 56, Anchorage, AK

99513.

The distributions and abundances of principal intertidal plants

and invertebrates are described at about 40 sites in the Kodiak

Island region.

Comparison of species composition and biomass on adjacent

transects at two sites indicated that spatial variability within

sites was high, obscuring seasonal patterns and confounding

between-site comparisons. Nevertheless, when sites were classed

according to exposure, topography, and substrate, exposed flat

bedrock beaches appeared to have lower species number and biomass

than other stable bedrock beaches. Mytilus edulis appeared to

reach highest densities in protected or partly protected areas

and were uncommon at exposed locations.

Densities of biota were lower at sites affected by frequent

physical disturbance such as scouring by sand or gravel and

unstable substrates (cobble and boulders) at exposed sites.

Benthic kelps were widely distributed and high in biomass in the

region of Kodiak Island. Very large individuals of Nereocystis
87

luetkeana and Laminaria dentigera were recorded. The total

biomass of Fucus spp. on rocky beaches in the Kodiak Island

region was estimated to be 3x107 to 1x108 kg. Diversity indices

did not reveal any clear patterns in species diversity of plants

or animals within or between sites.
88

1992 Addendum

These two manuscripts represent research conducted before the

Exxon Valdez oil spill; both manuscripts are currently in draft

form and nearing completion.

136 Karinen, J. F., M. M. Babcock, D. W. Brown, W. D. MacLeod,

Jr., L. S. Ramos, and J. W. Short. In Preparation.

Hydrocarbons in intertidal sediments and mussels from Prince

William Bound, Alaska, 1977-1980: Characterization and

probable sources. National Marine Fisheries Service, Auke

Bay Laboratory.

We collected and analyzed samples of sediments and mussels

(Mytilus trossulus) for alkane and aromatic hydrocarbons from

eight sampling stations adjacent to the oil tanker vessel

transportation corridor through Prince William Sound, Alaska,

during the period 1977 to 1980, to determine baseline

concentrations of these analytes prior to any pollution that

might result from oil tanker traffic through the sound.

Results indicate chronic, low-level hydrocarbon

contamination that probably originates from small fuel spills,

ballast water discharges, and fuel-combustion exhaust emissions

of occasional vessel activity adjacent to three stations:

Constantine Harbor, Rocky Bay, and Mineral Flats. The other five

stations show no indication of petroleum hydrocarbon

contamination; detected aromatic hydrocarbons are present only

sporadically and at concentrations that are generally near

detection limits. Exceptions are perylene, which is found at

concentrations well above detection limits at all sampling

station outside Port Valdez, and which probably has natural

sources; and phenanthrene, which is found sporadically at all

sampling stations and which also may come from natural sources.

Concentrations of individual n-alkanes vary substantially in

sediments and in mussels. The most abundant n-alkanes in

sediments include odd carbon-numbered alkanes of molecular weight

greater than tetradecane (C-14). Concentrations of these

n-alkanes are generally in the range of 10 to 100 ng/g dry

sediment weight, but exceed 1000 ng/g at Constantine Harbor. The

most abundant n-alkanes in mussels include decane (C-10) through

heptadecane (C-17), and pristane, at concentrations generally

ranging from 10 to several hundred ng/g dry tissue weight.

Sources of alkanes in sediments include terrigenous plant

waxes, marine plankton, and possibly marine macrophytic algae at

all the sampling stations, and petroleum-derived alkanes in

addition at Constantine Harbor.

Except in areas affected by local vessel traffic, intertidal

sediments and mussels in Prince William Sound are remarkably free

of petroleum-contaminant hydrocarbons during the period of this

study. The hydrocarbons found in sediments and mussels

unaffected by vessel traffic can be adequately explained by

known, natural sources. As a result, sediments and mussels

contaminated by crude oil from the Exxon Valdez oil spill should
89

be particularly apparent, due to the general absence

confounding sources of petroleum hydrocarbons.

of

137 Myren, R. T., G. Perkins, and T. R. Merrell. In

Preparation. Reduced abundance of Macoma balthica near an

oil tanker terminal in Port Valdez, Alaska, 1971-1984.

National Marine Fisheries Service, Auke Bay Laboratory.

Populations of Macoma balthica, a small intertidal clam in

mudflats, were monitored for 14 years (1971-1984) at Port Valdez,

to determine whether treated ballast water effluent would cause a

decrease in the populations. The Alyeska terminal began treating

oily ballast water from tankers in August 1977, averaging

2 million gallons of low-level oil in water per day. Populations

of large Macoma clams were stable for the 7 years before treated

ballast water discharge began, but declined to levels of less

than 20% by 1982-84. The population decline coincides with the

massive continuous discharge of effluent contaminated with low

levels of hydrocarbons and other unmeasured toxic compounds, but

cause and effect is not proven.
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91

INDEXES

Keyword Index

Note: All numbers correspond to reference numbers in the

bibliography and abstracts.

abundance 98, 109, 114, 130

accumulation (see hydrocarbon uptake)

acute toxicity (see toxicity)

Alaska Peninsula 134

alevins (see salmon, life stages)

algae 91, 118

amphipods 8

anchovy 26

Anodonta oregonensis 34, 35

Arctic 8

aromatic hydrocarbons 1, 3, 25, 26, 27, 29, 31, 34, 37, 38, 40,

67, 75, 78, 79, 80, 81, 83

artificial propagation 108

aryl hydrocarbons (see mixed function oxidase activity)

Atlantic cod 7

Atlantic mackerel 7

atlas, intertidal 128

avoidance 1, 48

ballast-water 55, 137

baseline 42, 136

bay pipefish 124

bay shrimp (see shrimp, species)

bear 95, 96

behavior 7

benzene 1, 3, 25, 26, 29, 40

benzopyrene 81, 82

biodegradation, hydrocarbon metabolism 78, 79, 81, 82

bioenergetics 43, 70, 72, 73, 74

biogeography 122

breathing rates 68, 83

caloric content 38

Chignik Lakes, River 92, 93, 94, 125

chinook salmon (see salmon, species)

chronic toxicity (see toxicity)

chum salmon (see salmon, species)

clam 41, 76, 137

cod (see Atlantic cod)

coho salmon (see salmon, species)

contaminated diet (see oil exposure type)

continuous-flow devices 39

Cook Inlet 98

Coonstripe shrimp

cortisol 77

(see shrimp, species)
92

coughing rates 67

crab, species

Dungeness 2, 24, 120

king 4, 5, 16, 18, 20, 32, 49, 53, 98

shore 17, 31

Tanner/snow 2, 23, 24, 99

crab, life stages

eggs 2, 24, 53

larvae, zoeae, glaucothoe 4, 5, 11, 32, 49, 98, 99

juveniles 20, 53

Crangonid shrimp (see shrimp, species)

creosol 29

crude oil (see oil, type)

depuration, hydrocarbon 17, 25, 26, 31, 51, 52, 54, 64, 68

development 9

devices 39

diet (see feeding)

Dolly Varden char 78, 79, 80, 81, 82

drilling muds 11, 12

Dungeness crab (see crab, species)

earthquake, 1964 114, 116, 131, 132, 133

effluent (see ballast water)

eggs (see life stages: crab, fish, salmon, shrimp)

epithelium 1

escapements 93

excretion, hydrocarbon 81, 82

fat content 29, 38

feeding 16, 43, 69, 70, 121

fish populations 104

fishery

crab 120

herring 127, 129

resources 33

salmon 92

shrimp 88, 97

fishes 8, 126

glaucothoe (see life stages: crab, shrimp)

glochidia, glochidium 34, 35

grease 71

greenling 124

growth 9, 16, 18, 29, 37, 38, 43, 62, 69, 70, 102, 105, 107,

109, 119

Gulf of Alaska 102, 103, 118, 122

gull 121

Hemigrapsus nudus 17, 31

herring (see Pacific herring)

hydrocarbon metabolism (see biodegradation)

hydrocarbon uptake 17, 25, 26, 28, 50, 51, 52, 54, 64, 68, 78,

79, 85

hydroxylase activity (see mixed function oxidase activity)
93

intertidal 17, 36, 41, 42, 91, 110, 113, 122, 128, 133, 135

invertebrates 114

Kachemak Bay 89, 98, 100, 101, 123

Katmai National Monument 104

kelp 90, 135

kelp shrimp (see shrimp, species)

king crab (see crabs, species)

Kodiak Island 89, 120, 134, 135

LC-50 (see toxicity)

larvae (see life stages)

life stages

fish (Atlantic cod, Atlantic mackerel, herring, pollock)

eggs, alevins, larvae 6, 7, 9, 10, 51

littoral 42

long-term exposure (see toxicity)

low temperature (see temperature effects)

mackerel (see Atlantic mackerel)

Macoma balthica 41, 76, 137

manacled sculpin 124

metabolism, hydrocarbon

metabolism,

(see biodegradation)

metabolic rate 52, 66

migration (see avoidance)

mixed function oxidase activity 13, 14, 85

molting 16, 18, 23, 32

morphology 1

mussels 74

naphthalene 27, 31, 34, 37, 38, 67, 75, 78, 80, 81, 83

narcosis 4

northern shrimp (see shrimp, species)

oil effects (see bioenergetics, feeding, growth, heart rates,

metabolism, molting, morphology, osmoregulation, oxygen

consumption, reproduction,

stamina)

scope for growth, swimming

oil exposure medium

diet 6, 16, 18, 69, 70, 80

sediment 2, 20, 24, 53, 72, 76

water-borne 6

water-soluble fraction 4, 5, 9, 10, 13, 23, 27, 32, 34, 39,

43, 50, 63, 66, 72, 74, 76, 77, 83, 84, 86, 87

oil, type

Cook Inlet 5, 27, 32, 50, 63, 66, 74, 77, 83, 85, 86, 87

No. 2 fuel oil 7, 63, 66

Prudhoe Bay 23, 34, 35, 40, 48, 62, 76, 84

olfactory epithelium 1

Olsen Bay 95, 114

Olsen Creek 95, 96, 105, 106, 107, 109, 110, 111, 112, 113, 121

osmoregulation 75

oxygen consumption 83

Pacific herring 6, 25, 50, 51, 127, 129

painted greenling 124
94

Pandalid shrimp (see shrimp, species)

parasitism 34

petroleum (see oil, type)

phenol 29, 81

pink salmon (see salmon, species)

pink shrimp (see shrimp, species)

pipefish 124

plankton 99

pollock (see Walleye pollock)

Port Valdez 19, 33, 41, 44, 55, 91, 137

predation 95, 96

pretreatment effects 67, 80

Prince William Sound 95, 96, 105, 106, 107, 109, 110, 111, 112,

113, 115, 117, 121, 124, 130, 131, 132, 133, 134, 136

reproduction 2, 24, 50, 51, 85, 86

reviews 15, 19, 21, 22, 33, 45, 46, 47, 60, 61, 65, 66, 117

Ruth Lake 119

salinity effects 40, 75, 78, 82

salmon, species 40, 108, 121, 133

chum 95, 96, 105, 106, 107, 109, 112, 115, 130, 131

coho 13, 14, 28, 34, 35, 37, 75, 77, 85, 86, 87

pink 1, 27, 29, 36, 38, 40, 48, 60, 62, 67, 68, 69, 70, 83,

84, 110, 111, 113, 115, 130, 131, 132

sockeye 40, 92, 93, 94, 119, 125

salmon, life stages

alevins 36, 60, 62, 115

eggs 28, 60, 62, 115

fry 34, 35, 37, 40, 48, 60, 62, 68, 69, 70, 75, 83, 84,

115, 130

sanddab 124

scallops 102, 103

scanning electron microscopy 1

scope for growth (see bioenergetics)

sculpin 124

sea scallops 102, 103

seastar 43

sediments (see oil exposure medium)

Shelikof Strait 117

shell height 103

shellfish 116

shore crab (see crab, species)

shrimp, species 27

Bay shrimp 3

coonstripe 5, 32

crangonid 100

kelp 4, 29, 67

northern 101, 123

Pandalid 72, 98, 123

pink 72, 88, 89, 97

shrimp, life stages 123

larval 5, 11, 32, 101

zoeae 100

snail 73
95

snow crab (see crab, species; Tanner crabs)

sockeye salmon (see salmon, species)

Southeast Alaska 117

spawning 85, 86, 110, 111, 112, 121, 125, 132

speckled sanddab 124

stock-and-recruitment 93

stock concept 108

streams 95, 96, 105, 106, 107, 109, 110, 111, 112, 121, 132

Striped bass 3, 26, 30

sublethal effects (see oil effects)

subtidal kelp 90

survival 105, 119, 133

swimming stamina 77, 87

Tanner crabs (see crab, species)

temperature effects 27, 79, 83

tidal cycles 17, 36, 41, 42

tolerance (see toxicity)

toluene 25, 27, 34, 37, 67, 75, 78, 79, 80, 83

toxic contributions 29

toxicity 47

acute 3, 4, 5, 8, 10, 27, 29, 34, 35, 40, 48, 50, 52, 55,

62, 63, 64, 65, 66, 67, 75

long-term 24, 28, 43, 52, 56, 57, 58, 59, 72, 73, 74

uptake (see hydrocarbon uptake)

Valdez (see Port Valdez)

Walleye pollock 9, 10

water-borne oil (see oil exposure medium, water-borne)

water-soluble fraction (see oil exposure medium, water soluble

fraction)

whales

killer 117

zoeae (see crab, life stages)
96

Author Index

Note: All numbers correspond to reference numbers in the

bibliography and abstracts.

Andrews, S. 39, 55

Babcock, M. M. 1, 2, 24, 36,49, 50, 51, 61, 85, 86

Bailey, J. E. 113

Barr, L. 88, 89

Bates, S. 37

Benville, P. E. 3, 30

Birkholz, D. G. 133

Bowles, A. E. 117

Brodersen, C. 4, 5, 50, 51, 53, 55, 61, 64

Brown, D. W. 29

Calvin, N. I. 42, 90, 91, 118, 135

Carls, M. G. 6, 7, 8, 9, 10, 11, 12, 50, 61, 72

Cheatham, D. L. 29, 64, 71

Collodi, P. 13, 14

Dahlberg, M. L. 92, 93, 94

Ellis, R. J. 90

Evans, D. R. 15

Frame, G. W. 95, 96

Fujioka, J. T. 135

Gharrett, J. A. 16, 17, 18, 42, 50, 51, 61, 135

Gish, R. K. 124

Hall, E. L. 126

Hall, J. D. 117

Hanson, J. L. 42, 135

Harry, G. Y. 97

Hartman, W. L. 104

Haynes, E. 98, 99, 100, 101, 102, 103

Heard, W. R. 104

Helle, J. H. 105, 106, 107, 108, 109, 110, 133

Hirsch,

111, 112, 113,

N. 25, 26

Hubbard, J. D. 114

Ignell, S. 117

Kapper, M. A. 72

Karinen, J. F. 2, 5, 19, 20, 21, 22, 23, 24, 32, 52, 53, 54, 56,

57, 58, 59, 61, 63, 64, 65, 66, 76, 136

Kirkwood, J. B. 115, 116, 130
97

Knull, J. 44

Korn, S. 3, 8, 25, 26, 27, 28, 29, 30, 37, 40, 50, 51, 54, 55,

56, 57, 58, 59, 61, 87

Krygier, E. 117

Lauren, D. J. 31

Leatherwood, S. 117

Lindsay, S. A. 55

Lindstrom, S. C. 91, 118

MacKinnon, J. S. 42, 135

MacLeod, W. D. 136

McMullen, J. C. 103

Mecklenburg, T. A. 5, 32, 64

Meehan, W. R. 119

Merrell, T. R. 33, 42, 137

Metcalf, W. 74

Meyer, R. M. 120

Misch, C. J. 64

Moles, D. A. 27, 34, 35, 36, 37, 38, 39, 40, 50, 60, 61, 62, 63,

64, 73, 85, 86

Moyle, P. 121

Myren, R. T. 41, 137

Noerenberg, W. H. 134

O'Clair, C. E. 42, 43, 122

Olsen, J. C. 123

Orsi, J. A. 124

Pella, J. J. 41

Pelto, M. J. 44

Perkins, G. 137

Phinney, D. E. 120

Prohaska, P. G. 136

Quast, J. C. 126

Ramos, L. S. 136

Reid, G. M. 127, 129

Rice, S. D. 5, 9, 10, 11, 12, 14, 15, 17, 18, 23, 24, 27, 28,

29, 31, 32, 36, 37, 38, 39, 40, 43, 45, 46, 47, 48, 49, 50;

51, 52, 53, 54, 55, 56, 57, 58, 59, 60, 61, 62, 63, 64, 65,

66, 67, 68, 71, 72, 73, 74, 75, 77, 78, 79, 80, 81, 82, 83,

84, 85, 86, 87

Sabourin, T. D. 75

Sakuda, H. M. 129

Schwartz, J. P. 69, 70

Sears, H. S. 128

Shirley, T. C. 72

Short, J. W. 5, 50, 62, 64, 65, 66, 68, 71
98

Skud, B. E. 129

Smith, H. D. 134

Stekoll, M. S. 14, 18

Stickle, W. B. 72, 73, 74, 75

Struhsaker, J. W. 25, 26, 30

Tait, H. D. 130

Taylor, T. T. 63, 76

Thomas, R. E. 67, 68, 77, 78, 79, 80, 81, 82, 83, 84, 85, 86, 87

Thorsteinson, F. V. 131, 132, 133, 134

Villars, C. 74

Wallace, R. L. 105

Williamson, R. W. 44, 113

Wing, B. L. 124

Yancey, R. M. 116

Zimmerman, S. T. 128, 135
RECENT TECHNICAL MEMORANDUMS

Copies of this and other NOAA Technical Memorandums are available from the

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(web site: www.ntis.gov). Paper and microfiche copies vary in price. 

AFSC-

 

2 KAJIMURA, H., and E. SINCLAIR. 1992. Fur seal investigations, 1990, 192 p. NTIS No. PB93-

109080.

1 MERRICK, R. L., D. G. CALKINS, and D. C. MCALLISTER. 1992. Aerial and ship-based surveys

of Steller sea lions (Eumetopias jubatus) in southeast Alaska, the Gulf of Alaska, and Aleutian Islands

during June and July 1991, 41 p. NTIS No. PB92-235928. 

F/NWC216 ZENGER, H. H., JR., and M. F. SIGLER. 1992. Relative abundance of Gulf of Alaska sablefish

and other groundfish based on National Marine Fisheries Service longline surveys, 1988-90, 

103 p. NTIS No. PB92-222843.

215 KINOSHITA, R. K., B. M. K. BROOKE, L. E. QUEIROLO, and J. M. TERRY. 1992. Economic status of

the groundfish fisheries off Alaska, 1990, 99 p. NTIS No. PB92-187699.

214 ANTONELIS, G. A. 1992. Northern fur seal research techniques manual, 47 p. NTIS No. PB92-191824.

213 BAKKALA, R. G., W. A. KARP, G. F. WALTERS, T. SASAKI, M. T. WILSON, T. M. SAMPLE,

A.M. SHIMADA, D. ADAMS, and C. E. ARMISTEAD. 1992. Distribution, abundance, and biological

characteristics of groundfish in the eastern Bering Sea based on results of U.S.-Japan bottom trawl and

hydroacoustic surveys during June-September 1988, 362 p. NTIS number pending. 

212 WILSON, M. T., and C. E. ARMISTEAD. 1991. 1989 bottom trawl survey of the eastern Bering Sea

continental shelf, 212 p. NTIS No. PB92-135607.

211 LOW, L-L. (editor). 1991. Status of living marine resources off Alaska as assessed in 1991, 

95 p. NTIS No. PB92-145713.

210 LOW, L-L. (editor). 1991. Status of living marine resources off the Pacific coast of the United States as

assessed in 1991, 69 p. NTIS No. PB92-145705.

209 KINOSHITA, R. K., D. A. COLPO, and J. M. TERRY. 1991. Oregon, Washington, and Alaska exports of

edible fishery products, 1990, 48 p. NTIS No. PB92-115971.

207 LIVINGSTON, P. (editor). 1991. Groundfish food habits and predation on commercially important prey

species in the eastern Bering Sea from 1984 to 1986, 240 p. NTIS No. PB92-104710.

206 LORENZ. J. M., M. L. MURPHY, J. F. THEDINGA, and K V. KOSKI. 1991. Distribution and abundance

of juvenile salmon in two main channel habitats of the Taku River, Alaska and British Columbia, 16 p.

NTIS No. PB91-234344.

205 DORN, M. W. 1991. Spatial and temporal patterns in the catch of Pacific whiting in the U.S. management

zone during 1978-88, 68 p. NTIS No. PB91-227702.