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TRANSCRIPT
Foods Utilized by Bowhead Whales NearBarter Island, Alaska, Autumn 1979
LLOYD F. LOWRY and JOHN J. BURNS
Introduction
Bowhead whales, Salaena mysticetus, spend the late spring, summer, andearly autumn in the Beaufort Sea (Sergeant and Hoek, 1974). On their northward spring migration they usuallybegin to pass Point Barrow, Alaska, inthe latter part of April (Durham, 1979;Braham and Krogman l
; Marquette2).
Their penetration of the spring ice sheetfrom the central Bering Sea is as muchas 3,000 km (Tomilin, 1957). DuringSeptember and October they again passPoint Barrow on their way into theChukchi Sea, some at least havingmoved westward along the Alaskancoast (Braham and Krogman, footnoteI). The latest sighting of a bowheadnear Point Barrow in the autumn, ofwhich we are aware, was of one seen on14 November 1977 (Burns, pel's. obs.).
The Beaufort Sea is a region of greatseasonal extremes. A cover of ice andsnow largely prevents photosynthetic primary production for at least 4months of the year. A brief period ofopen water occurs each summer, theduration and geographical extent ofwhich vary greatly from year to year. Inspite of such conditions, which wouldappear not to favor regular productionof large plankton stocks, several species of animals, including the bowheadwhale, use the Beaufort Sea as a summer-early autumn feeding area.
'Braham, H., and B. D. Krogman. 1977. Population biology of the bowhead (Sa/aena mysticetus)and beluga (De/phinapterus /eucas) whale in theBering, Chukchi, and Beaufort Seas. Processedrep., 29 p. Natl. Mar. Mammal Lab., Natl. Mar.Fish. Serv., NOAA, 7600 Sand Point Way N.E.,Bldg. 32, Seattle, WA 98115.2Marquette, W. M. 1977 The 1976 catch ofbowhead whales (Sa/aena mysticetus) by Alaskan Eskimos, with a review of the fishery, 197376, and a biological summary of the species.Processed rep., 80 p. atl. Mar. Mammal Lab ..Natl. Mar. Fish. Serv., NOAA, 7600 Sand PointWay N.E., Bldg. 32, Seattle, WA 98115.
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It is not known whether bowheadsfeed during winter while they are in theBering Sea. They feed only little or notat all during their spring migration(Johnson et aI., 1966; Marquette, footnote 2; Foote3 ). Intensive feeding occurs during summer and autumn in theeastern Arctic (Scoresby, 1820) and indeed in Alaskan waters the only whalestomachs which contained appreciableamounts of food have been taken during the autumn whaling season (Lowryet aI., 1978). Large aggregations ofbowheads, which appeared to be feeding, have been seen nearshore east ofPoint Barrow in September (Brahamand Krogman, footnote I; Burns, pel's.obs.). Intensive feeding presumablyoccurs while the whales are in theBeaufort Sea. The quantities and kindsof available food in this region must beadequate to meet immediate bioenergetic requirements of whales and toprovide reserves for periods of reducedfeeding.
There are no published accounts ofwhat foods are utilized by bowheads inthe Beaufort Sea east of Point Barrow.We have recently acquired and examined samples of stomach contentsfrom fi ve of these animals taken between 20 September and 11 October1979 near Barter Island, Alaska.
Methods
In the field, samples ranging involume from about 200 to 2,500 mlwere randomly collected from thewhales' stomachs. From the whaledesignated as 79-KK -5, an additionalnonrandom sample was taken whichincluded primarily large, unusual appearing items. With the exception of
3Foote, D. C. 1964. Observations of the bowheadwhale at Point Hope, Alaska. Unpubl. manuscr.,78 p. Geogr. Dep .. McGill Univ., Montreal,Que.. Can.
Lloyd F. Lowry and John J. Burns are with theAlaska Department of Fish and Game, 1300 College Road, Fairbanks, AK 99701.
whale 79-KK-I, which was recovered 2days after it was struck, samples werecollected within a few hours of the timethe whale was killed and landed. Sampleswere labeled, preserved in a 10 percentFormalin4 solution, and delivered to Fairbanks, Alaska.
In the laboratory, a 100 ml aliquot ofeach sample was taken for possiblefuture fatty acid content analysis. Theremainder of the samples were drainedand gently washed on a 1.00 mm meshsieve. The stomach contents were thensorted macroscopically into major taxonomic groups, and the volume of eachgroup was determined. The items ineach group were examined microscopically when necessary and identified tospecies if possible. The number ofindividuals and total volume of eachprey species were determined and thelengths of representative specimenswere measured. In the case of abundant, small organisms such as copepods. the volume and number of thevarious species were estimated fromsubsamples.
Results
Based on the quantities and condition of the stomach contents, all fivewhales were feeding at the time theywere killed. Estimated total volume ofstomach contents varied from 19 to 45 I(5-12 gallons). A summary of the species present, approximate abundance ineach sample, and those previously reported as food items in whales taken offnorthern Alaska are presented in Table1. Copepods were found in all fivesamples and were the dominant fooditem in three. Calanus hyperboreuswas by far the most commonly eatenspecies. The stomach contents of whale79-KK-4 were in particularly goodcondition. In that whale most of the C.hyperboreus eaten were copepod idStage Y. Euphausiids, almost exclu-
"Reference to trade names or commercial firmsdoes not imply endorsement by the NationalMarine Fisheries Service, NOAA.
Marine Fisheries Review
Table 1.-Food items from bowhead whale stomach samples collected at Kaktovik, Barter Island, Alaska, September-October 1979.
Whale specimen number Whale specimen number
Food item 79·KK-l 79-KK-2 79-KK-3 79-KK·4 79-KK-5 Food item 79-KK-l 79-KK-2 79-KK-3 79-KK-4 79-KK-5
Copepods Monoculoides c.f. M. schneider; X XCalanus finmarchicus XX Munnopsis c.t. M. typica XCalanus hyperboreus XXX XXX XX XXX XX Onisimus glacialis XChiridius obtusifrons XX Onisimus Jiloralis -XHeterorhabdus sp. XX XX Rozinante fragifis 1 XMetridia lucens XX XX Weyprechtia heulgini X
Family Lysianassidae XEuphausiids
Thysanoessa raschii' XX XXX XX XXX ShrimpsThysanoessa inermis X Euafus gaimardii X
Sabinea septemcarinata XMysids Family Crangonidae X
Mysis litora/is XX XX XX Unidentified X
Hyperiid amphipods IsopodsHyperia medusarum X Saduria entomon XHyperia sp. XParathemisto abyssorum 1 XXParathemisto libel/ula I X X X X Cumaceans
Oiastylis sp. X
Gammarid amphipodsAcanthostepheia behringiensis' X FishesAcanthostepheia incarinata X Boreogadus saida XApherusa glacialis X Myoxocephalus quadricornis XAtyJus carinatus X X Pungitius pungitiusGammaracanthus loricatus X Unidentified XGammarus Sp.1 X
Pebbles X X
XXX = dominant item in sample.XX = more than 10 individuals in sample.X = 1·10 individuals in sample .. = present in qualitative sample only.'Indicates previously reported food items of bowhead whales from northern Alaska (Lowry et al .. 1978; Marquette. 1979).
1The overall mean percent of contents was calculated based on the volume and percent composition of each sample and theestimated total contents of stomachs from which the samples were taken.
Table 2.-Quanlitative composition of stomach contents from bowhead whales collected at Kaktovik, autumn 1979.For each whale, numbers indicate percent of the sample volume comprised of each prey type.
Whale specimen number Overall meanpercent of
Prey type 79-KK-l 79-KK·2 79-KK-3 79-KK-4 79-KK-5 contents1
Copepod 99.7 99.0 23.4 88.3 <0.1 598Euphausiid 0.3 67.8 4.9 97.9 37.2Mysid 0.3 7.0 0.8 0.2Hyperiid amphipod <0.1 0.1 0.5 0.4 0.1Gammarid amphipod 0.1 0.1 0.3 2.4 01 0.4Other invertebrate <0.1 ,'_0.1 2.3 1.1 0.6Fish <0.1 0.1 1.0 1.7 0.4Sample volume (ml) 2,406.2 545.2 399.7 131.3 357.9Estimated total volumeof contents (gallons) 12 6 5 10
sively Thysanoessa raschii, occurredin four samples and were dominant intwo. Mysis litoralis was common inthree samples. Although at least 22other prey species were identified,none were major components of thefood. The majority of identified specieswere amphipods, most of which arebenthic forms.
A quantitative summary of the majorprey types found in the samples isshown in Table 2. In addition to theproportion of each sample which wascomprised of each prey type, an overallmean percentage of contents was calculated for each prey type based on thevolume of subsamples, the proportionof prey types in the subsamples, and theestimated total contents of each stomach. Copepods and euphausiids in combination made up 91.2-99.7 percent ofthe contents of all subsamples. Therelative importance of these two preytypes varied markedly from over 99percent copepods in whales 79-KK-Iand 79-KK -2, to almost 98 percenteuphausiids in 79-KK-5. In the overallcomposition of the food, cope podswere more important in terms of vol-
Sept.-Oct. 1980
ume than were euphausiids. All otherfood types were insignificant in termsof overall volume consumed.
Discussion
The samples of stomach contents justdescribed constitute the only data available on bowhead whale foods in theeastern portion of the Alaskan BeaufortSea. Two whales taken at Barrow inSeptember 1976 had eaten mostly euphausi ids (Lowry et aI., 1978). Theprincipal euphausiid species (Thysa-
noessa raschii) found in whales takenat Kaktovik was the same as that foundin the whales taken at Barrow. Thisspecies is widely distributed in Arcticwaters (Geiger et aI., 1968) and is ofgreatest abundance in the nearshoreneritic zone (Nemoto, 1966). A whaletaken at Barrow in May 1977 had eatenmainly copepods (Marquette, 1979).None of the copepod species eaten atKaktovik were found in the whale takenat Barrow. This is particularly interesting since the main species eaten at
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Figure I -Diagrammatic representation of major trophic connections in the pelagicfood web of the western Beaufort Sea.
bowheads ----..
phytoplankton ~e:::.:::'~d~J/ ?'WC cod~ ',"ged seal, ~meroplankton -------7- !/ belukhas humans
bacteria hyperl;d amph,pods ~ /
myslds \ ~detritus r.mged seals ~ polar bears ]
~<lrctlc foxes
birds -------=- ~
Barrow (Calanus glacialis and Metridia longa) and Kaktovik (Calanushyperboreus) are all considered to bewidely distributed in surface waters ofthe Arctic Ocean (Grainger, 1965). It isprobable that copepods and euphausiids are, overall, the most importantbowhead foods in the Beaufort Sea.Mysids and hyperiid amphipods alsoare concentrated in some regions andmay be locally important foods.
A large number of prey species wereidentified from the Kaktovik whales.Mysids, isopods, and cumaceans werenot found among the Barrow specimens. The only previous indication thatfishes might be eaten by bowheads wasa fish scale in specimen 77-B-5 (Marquette, 1979). Fishes representingthree species were found in the whalestaken at Kaktovik. In addition, at least10 species of amphipods and 2 speciesof shrimp were first recorded as bowhead prey in these samples.
Although organisms such as gammarid amphipods, fishes, and isopodsare apparently not major foods of bowheads in terms of quantities consumed,their presence in the stomach contentsis of some significance. The presenceof pebbles and bottom dwelling speciesindicates that all the whales taken atKaktovik had fed at least partially nearthe sea floor. Pelagic prey far outnumbered the benthic organisms. Therefore, a feeding dive probably involvesswimming obliquely from surface tobottom and back, feeding the entiretime. Saduria entomon has not beenfound deeper than 44 m in the BeaufortSea (McCrimmon and Bray, 1962).Myoxocephalus quadricornis and Pungitius pungitius are characteristic ofcoastal fresh and brackish water (Andriyashev, 1954; Walters, 1955). Therefore, at least whales 79-KK-3 and 79KK-5 had fed in relatively shallow,nearshore waters.
Although exploration and development of Beaufort Sea hydrocarbonreserves are underway in Canadianwaters and imminent off Alaska, thepotential effects of such activities onfoods of bowhead whales and foodwebs of the Beaufort Sea are impossibleto predict at this time. The only majorbowhead prey species which has been
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tested for hydrocarbon sensiti vity isCalanus hyperboreus. This specieswas found to be "surprisingly resistantto all of the oils tested" (Percy andMullin, 1975). Hydrocarbon sensitivity data for other key species in theBeaufort Sea food web such as othercopepods; Arctic cod, Boreogadussaida; euphausiids; mysids; and hyperiid amphipods are urgently needed.
In the Alaskan Beaufort Sea, bowheads depend primarily on a pelagicfood web (Fig. I). Arctic cod feedprimarily on copepods in offshore(greater than 40 m deep) waters duringsummer (Frost et a1. 5 ). During thesummer, ringed seals, Phoca hispida,feed mostly on euphausiids, hyperiidamphipods, and mysids (Lowry et a1. 6 ,
1978). Ringed seals and Arctic cod areprobably the most significant trophiccompetitors of bowhead whales inthe Beaufort Sea. Changes in popUlations of these competing species,whether natural or caused by AlaskaOuter Continental Shelf oil and gas
5Frost, K. J., L. E Lowry, and J. J. Burns. 1978.Offshore delrersal fishes and epibenthic inverte·brates of the northeastern Chukchi and westernBeaufort Seas. Appendix to: Trophic relationships among ice inhabiting phocid seals. AnnuRep. RU 232, 39 p. U.S. Dep. Cammer., Natl.Oceanic Almas. Admin., Outer Continental ShelfEnvironmental Assessment Program, Boulder,Colo.6Lowry, L. E, K. J. Frost. and J. J. Burns. 1979.Trophic relationships among ice-inhabiting phocid seals. Final Rep. Beaufort Sea Activities, RU232,55 p. U.S. Dep. Commer., Natl. OceanicAlmos. Admin .• Outer Continental Shelf Environmental Assessment Program. Boulder,Colo.
development activities, may affect thesize and recovery rate of the bowheadpopulation.
Acknowledgments
This study was part of ProjectWhales (Research Uni t 280), completed by the Naval Arctic ResearchLaboratory, Barrow, Alaska. ProjectWhales was funded by the U.S. Bureauof Land Management/Outer Continental Shelf studies. T. Albert, University of Maryland, coordinated the acquisition of specimens and undertookfield work to acquire some of them. G.Jarrell, M. Nerini, and D. Smullin,with the National Marine FisheriesService, Seattle, Wash., also providedassistance -Jarrell and Smullin by obtaining some specimens at Barter Islandand Nerini by transporting and makingthem available to us. Laboratory facilities were provided by the AlaskaDepartment of Fish and Game, Fairbanks. We are indebted to G. Mueller,A. Adams, and K. Coyle, University ofAlaska, Institute of Marine Sciences,for identification of copepods and amphipods. K. Frost, Alaska Departmentof Fish and Game, Fairbanks, providedinvaluable assistance in all phases ofthis study. Our thanks are also extendedto those whaling captains and crewsat Barter Island who cooperated inthis effort.
Literature CitedAndriyashev, A. P. 1954. Ryby severnykh
morei SSSR (Fishes of the northern seas of the
Marine Fisheries Review
U.S.S.R.). Acad. Nauk SSSR, Opredelitelipo Faune SSSR 53, 566 p. Izd. Akad. NaukSSSR, Moscow-Leningrad. (Translated by Isr.Program Sci. Transl., 1964,617 p.; avail. U.S.Dep. Commer., Natl. Tech. Inf. Serv.,Springfield, Va., as OTS63-11160.)
Durham, F E. 1979. The catch of bowhead whales (Ba/aena mysticetlls) by Eskimos' with emphasis on the western Arctic.Nat. His!. Mus. Los Ang. Cty., Contrib. Sci.314,14 p.
Geiger, S. R., K. Rodriguez, and M. M. Murillo.1968. Euphausiacea of the Arctic Ocean andits peripheral seas. Bull. South. Calif. Acad.Sci. 67:69-79.
Grainger, E. H. 1965. Zooplankton from theArctic Ocean and adjacent Canadian waters.J. Fish. Res. Board Can. 22:543-564.
Johnson, M. L., C. H. Fiscus, B. T. Ostenson,and M. L. Barbour. 1966. Marine mammals. InN. J. Wilimovsky and J. N. Wol fe(editors), Environment of the Cape Thompson
region, Alaska, p. 877-924. U.S. A. E c.,Oak Ridge, Tenn.
Lowry, L. F, K. 1. Frost, andJ. J. Burns. 1978.Food of ringed seals and bowhead whales nearPoint Barrow, Alaska. Can. Field-Nat. 92:67-70.
Marquette, W. M. 1979. The 1977 catch ofbowhead whales (Ba/aena mysticetlls) byAlaskan Eskimos. Rep. Int. Whaling Comm.29:28 I-289.
McCrimmon, H., and J. Bray. 1962. Observations on the isopod Mesidotea entomon inthe western Canadian Arctic Ocean. J. Fish.Res. Board Can. 19:489-496.
Nemoto, T. 1966. Thysanoessa euphausiids,comparative morphology, allomorphosis andecology. Sci. Rep. Whales Res. Inst. 20:109-155.
Percy, 1. A., and T. C. Mullin. 1975. Effectsof crude oils on arctic marine invertebrates.Beaufort Sea Project Tech. Rep. II, 167 p.Dep. Environ., Victoria, B.C.
Scoresby, W., Jr. 1820. An account of theArctic regions, with a history and descriptionof the northern whale-fishery. Vol. 2. Archibald Constable and Co., Edinburgh, 574 p.
Sergeant, D. E., and W. Hoek. 1974. Seasonal distribution of bowhead and whitewhales in the eastern Beaufort Sea. In J. C.Reed and J. E. Sater (editors), The coast andshelf of the Beaufort Sea, p. 705-719. ArcticInst. North Am., Arlington, Va.
Tomilin, A. G. 1957. Zveri SSSR i prilezhashchikh stran (Mammals of the U.S.S.R.and adjacent countries). Vol. 9. Kitoobraznye(Cetaceans). Izd. Akad. Nauk SSSR, Moscow, 756 p. [In Russ.] (Translated by lsI.Program Sci. Trans!., 1967,717 p.; avail. U.S.Dep. Com mer. , Natl. Tech. Inf. Serv.,Springfield, Va., as TT 65-50086.)
Walters, V. 1955. Fishes of westernArctic America and eastern Arctic Siberia,taxonomy and zoogeography. Bull. Am.Mus. Nat. Hist. 106:259-368.
Table 1.-Amino acid content of urine from a bowheadwhale.
Some Observations on UrineFrom a Bowhead Whale
also occasional oxalate crystals as wellas a few triple phosphate crystals.
On further analysis the following results were obtained: Sodium ion 183meq/I, potassium ion 14.4 meq/I,chloride ion 433 meq/I, osmolality1,440 milliosmols/I, creatinine 400mg/dl, urea N 3,000 mg/dl. The aminoacid pattern obtained can be seen inTable I and is similar to that seen inadult mammals such as the dog, The
W. MEDWAY
The analysis of urine is an essentialpart of any examination of an animal todetermine its health status. This is trueespecially when (for purposes of comparison) there is a large store of dataabout urine of the species in question.The urine of the bowhead whale,Balaena mysticetus, has apparently notbeen examined.
On 18 May 1978 a urine sample identified as having originated from abowhead whale, designated by the National Marine Fisheries Service as#78B2, was presented to this laboratory. The urine was obtained by cystocentesis from a male about 8.4 m inlength, taken by Eskimo hunters on 2May 1978 at Barrow, Alaska. The urinewas refrigerated until 6 May, then frozen. On 15 May the frozen specimenwas prepared for shipment. Unforeseen
W. Medway is Chief. Section of ClinicalLaboratory Medicine, School of VeterinaryMedicine, University of Pennsylvania, Philadelphia, PA 19104.
delays during shipment allowed theurine to thaw by the time it reached thislaboratory. Findings are presented herenot only for general information butalso as a small contribution to the dataconcerned with the biology of this endangered species.
Results and Discussion
On routine clinical urinalysis the following observations were made. Theurine was dark amber, clear, and had noodor. The pH was 5.5, and the specificgravity (SG) was 1.032; there was atrace of protein; and the tests forketones, glucose, reducing substances,bile pigments, hemoglobin, andurobilinogen were negative_ Microscopic examination of the sediment revealed a few red and white cells, but nocasts; however, there were myriads ofepithelial cells (some were cornified),and there were some bladder transitional cells and caudate cells. Therewere many unidentified spheroid crystals (probably urates), and there were
Amino acid
ThreonineSerineAsparagineGlutamineProlineGlutamic acidGlycineAlanineex Aminoadipic acida Amino-n-butyric acidValineHalf cystineCystathionineMethionineIsoleucineLeucineTyrosinePhenylalanineOrnithineLysine1-MethylhistidineHistidine3-MethylhistidineArginineTrimethyllysineNN dimethylarginineN'N dimethylarginine
Nanomoles/mgcreatinine
312899326411844314.51718
1152510
Trace182910
Trace17
Trace16
TraceTrace204
3124
Sept.-Ocr. 1980 91