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THE DIET OF THICK-BILLED MURRE CHICKS IN THE EASTERN CANADIAN

A. J. GASTON CanadianWildlife Service, Environment , Ottawa, Ontario KIA OE7,Canada

ABSTRACT.--Idescribe the diet of Thick-billed Murre (Uria lornvia) chicks at three colonies in the low Arctic of easternCanada. These are comparedwith diets describedpreviously for in the same area and Prince in the high Arctic. Diets varied more amongcolonies than amongyears at the samecolony. Indices of similarityshowed the diet of chicks at Hantzsch Island to be the most distinct. The diets of all low-arctic colonies were more diverse than those reported at . Some factors governing differencesamong coloniesare discussed.Received 26 November1984, accepted 9 April 1985.

THICK-BILLEDMurres (Uria lomvia) breed over another than to the diet at Prince Leopold Is- a great range of latitude in easternCanada, from land in the high Arctic? (3) Is the diet more 47øNat Cape St. Mary's, Newfoundland to 76øN diverse at low-arctic colonies than at Prince at , , all LeopoldIsland? within waters classified as either low arctic or STUDY AREA AND METHODS high arctic (Salomonsen 1972, Brown et al. 1975). The speciesis the most numerous sea- Thick-billed Murres feed their chicksfor 15-30 days bird in , , and as- at the nest site before the young birds depart to sea sociated channels in the high Arctic and in at about 20% of adult weight (Birkhead and Nettle- and adjacentwaters in the low ship 1981, Gaston and Nettleship 1981). Food is car- Arctic (Brown et al. 1975, Gaston 1982). ried to the site in the adult'sbill; normally, only one North of 60 ø latitude Thick-billed Murres item is brought at a time. The meal is delivered to breed at only 10 coloniesin the eastern Cana- the chick immediately after the incoming adult has landed, and it is therefore difficult to observe the dian Arctic, all of more than 10,000pairs (Fig. type of food brought. Somemeals are dropped, how- 1). Five of these are situated between 60ø and ever, and such discarded items can be collected on 64øN, all in low-arctic waters, and 4 between the breeding ledgesduring the chick-rearingperiod 72ø and 76øN in high-arctic waters. The remain- to determine the diet of the chicks. ing colony is closeto the high-arctic/low-arctic At Prince Leopold Island, Gaston and Nettleship boundary.The samplesdescribed here include (1981)showed that the proportionof different types the mostwesterly and the most easterlyof the and sizes of food identified in the diet of Thick-billed low-arcticgroup. Hence, they should provide Murre chicksvaried with the collectingmethod. Fish a good indication of the range of variation in found on ledges may be those brought to eggs not the diet of Thick-billed Murre chicksthrough- yet hatched, or to chicks that have left the colony. out their range in Canadian low-arctic waters. Consequently,the type of food fed either to very The diet of Thick-billed Murre chicks at young or to very old chicksmay be overrepresented. Where comparisonsare made amongcollections made Prince Leopold Island, in Barrow Strait, was by the same method, however, such biases should describedby Gastonand Nettleship (1981). In not greatly affect the considerationof diversity or this paper I describetheir diets at three colo- overlap. nies in or near Hudson Strait [Coats Island Collections and observations of food delivered to (25,000 pairs), Digges Island (with adjacent Thick-billed Murre chicks were made at Digges Is- mainland cliffs, 300,000 pairs), and Hantzsch land in 1980, 1981, and 1982; at Coats Island in 1981 Island (50,000 pairs)] and comparethese with and 1984;and at HantzschIsland in 1982.Specimens previouslypublished data from the south col- were collected during visits made to the breeding ledgesto weigh and measurechicks or to band them. ony at Akpatok Island (Tuck and Squires1955) At Coats Island and Hantzsch Island these collections and with data from Prince Leopold Island. I were augmentedby observationsmade at closerange addressthree principal questions:(1) Is vari- (<10 m), when food items were identified visually. ability amongcolonies greater than year-to-year All fresh and complete specimenswere weighed variation at individual colonies? (2) Are diets to the nearest 1 g and their overall length measured at the low-arctic colonies more similar to one (+1 mm). They then were preservedin 70%ethanol, 727 The Auk 102: 727-734. October 1985 728 A.J.GASTON [Auk,Vol. 102

For present purposes,where I have combined data from sightrecords and collections,I have had to lump certain taxonomicgroups that can be distinguished only in the hand. For comparisonof overlap in- volving this type of data, I have lumped data from all samplesinto the samecategories, namely, capelin (Mallotusvillosus), lanternfish (Myctophidae),arctic cod (Boreogadussaida), blennies (Blennioidei), sand lance (Ammodytesspp.), redfish (Sebastesspp.), scul- pins (Cottidae),lumpfish (Cyclopteropsisspp.), lump- suckers(Eumicrotremus spp.), snailfish (Liparisspp.), Greenlandhalibut (Reinhardtiushippoglossoides), squid (Gonatusfabricci), and Crustacea.Comparisons made with Prince Leopold Island are basedon Table 66 of Gastonand Nettleship (1981) and thosewith Akpa- tok Island are basedon Table 1 of Tuck and Squires (1955).Comparisons of diet diversity are basedonly on specimencollections, all of which were identified to species level. Names used follow Hunter et al. Fig. 1. The easternCanadian Arctic, showing the (1984). position of Thick-billed Murre colonies mentioned in the text. RESULTS

Food items fed to Thick-billed Murre chicks and representativespecimens of all taxa were iden- tified by staff of the National Museums of Canada. ranged from lessthan 1 g to 41.5 g in weight. For difficult groups, such as blennies (Blennioidei) The mean weight of items collected ranged and sculpins(Cottidae), all identificationswere con- from 5.6 g at DiggesIsland in 1982 to 11.6 g at firmed by museumpersonnel. Dried specimenswere Coats Island in 1981 (Table 1). All collections measured,and their weights were determined from showed a strongly skewed distribution, with the regressionof weight on length3 determined for the commonestsize classbeing below 5 g. The freshspecimens (Appendix 1). In estimatingthe con- range and variation among weights was similar tribution of different taxato chick diets by weight, I among all samples.The size of individual items used only specimens collected, including those gives a good indication of the size of mealsfed weighedfresh and thoseestimated from length mea- surements. to chicksbecause practically all meals seen de- To describethe similarity between diets at differ- livered consistedof only a single item, except ent colonies,or at the samecolony in different years, at Coats Island in 1984, where 6% (n = 125) of I usedMorisita's Index (CA),as recommended by Wol- deliveries were of two items. Mean weights of da (1981): individual taxa basedon fresh specimensonly are given in Appendix 2. 2 X n•n2• Cx Fish comprisedmore than 90% of items in all (X• + X2)N•N2' samples(Fig. 2). The commonestin all samples where)•i = IX n,(nj,- 1)]/[Nj(Ni - 1)],N i = the num- except those from Hantzsch Island was arctic ber of specimensin samplej, and nj•= the number cod. Other common constituentswere capelin of specimensof speciesi in samplej. To describethe (29% of items at Coats Island in 1984), blennies diversity of the diets, I used the Shannon-Weaver (14-18% of items at Digges Island), sand lance diversity index (H'; Tramer 1969): (up to 20% of items at Digges Island), and scul- pins (48% of items at Hantzsch Island). Full de- H'= - i p,(lnp•), tails of speciesidentified and numbers record- ed in this study are given in Appendix 2. The where p• = the proportion of all specimensin the picture was broadly similar when diets were samplethat belongto speciesi and s = total species considered by weight. The predominant fish in the sample. The magnitude of both these indices is consider- remained the same in all samples(Fig. 3), but ably affectedby the level of identificationemployed. capelin and sand lance were lessimportant and The more categoriesinto which the sampleis split, blennies more important. the greater the potential diversity and the smaller Although arctic cod was the commonestfish the likely overlapbetween samples (Diamond 1983). recorded in all but one sample, ranging from October1985] Foodof Thick-billedMurre Chicks 729

TABLE1. Mean weights (g) and size range of food items fed to Thick-billed Murre chicksat three colonies.

Weight Colony Year Mean SD n Median Range Coats Island 1981 11.6 8.74 23 11.0 < 1-35.0 1984 10.1 9.01 42 6.0 0.7-41.5 Digges Island 1980 8.9 8.25 59 5.7 0.7-31.8 1981 8.6 6.66 36 7.2 1.5-30.0 1982 5.6 4.17 17 4.1 1.4-16.4 Hantzsch Island 1982 10.1 7.23 35 9.0 1.0-29.0

14% (Hantzsch Island) to 62% (Digges Island, 178 specimenscollected at Prince Leopold Is- 1982) of the diet by number, it never formed land, comparedwith 11 speciesat CoatsIsland as great a proportion as at Prince Leopold Is- (n = 231), 14 at Digges Island (n = 194), and 9 land, where it constituted78% of itemsbrought at Hantzsch Island (n = 73). At Akpatok Island over three years. Capelin and sand lance, both Tuck and Squires'svery muchlarger sample of frequent at the low-arctic sites, were not re- 2,530 fish included 20 taxa of species-levelor cordedat all at Prince Leopold Island. Sculpin higher. were the predominantgroup of fishat Hantzsch Island, but elsewherethey did not exceed27% DISCUSSION of items (at Akpatok Island). Similarity indices calculatedfrom frequen- Comparisonof similarity indicescalculated cies for each combination of years at Digges for samplesfrom the samecolony in different Island, Prince Leopold Island, and CoatsIsland yearswith thosecalculated for samplesfrom were all closeto the expectedmaximum value differentcolonies strongly suggests that differ- of C• of "about 1" (Wolda 1981; Table 2), indi- encesin diet among coloniesare greater than cating that variation among years at the same those causedby annual variation at any one colony is insignificant.Consequently, I com- colony.Because not all sampleswere taken in bined samplesfrom different yearsto increase the sameyears, the differencesamong colonies samplesizes for intercolony comparisons(Ta- could be the result of fluctuationsin the pop- ble 3). All colonysamples differed significantly ulationsof certainfish species, which took place from every other colony in the frequency of over the entire eastern Canadian Arctic. How- differentfood types,using a Chi-squaretest for ever,this is unlikely to applyto differencesbe- heterogeneity(Table 4). tween Coats, Digges, and Hantzsch islands, Chicks at Hantzsch Island had the most dis- where all samplingwas carriedout within four tinct diet, with a mean similarity index of 0.53 years.It seemsreasonable to concludethat there (Table 4). The group of colonieswithin Hud- are real differencesamong colonies in chick sonStrait (Coats, Digges, and Akpatokislands) diets. showedthe greatestsimilarity to one another, Comparisonsof the samples from the low all three pairingshaving indicesgreater than Arctic with those from Prince Leopold Island 0.8. show that the diversity of the diet is consid- Diversityof speciesin the sampleswas great- erably greater at the low-arctic colonies,pre- est at Coats Island in 1984 (H'= 1.91) and sumablyreflecting the greaterdiversity of prey HantzschIsland (H' = 1.81).At DiggesIsland speciesavailable to the Thick-billed Murres. diversity indices ranged from 1.19 in 1982 to Hunter et al. (1984) record48 speciesof marine 1.41 in 1981 and 1.54 in 1980. It was not pos- fishes from Hudson Strait, , and sible to calculatea comparableindex for Tuck's southeastBaffin Island, comparedwith only 27 Akpatok samples because certain families of speciesfrom LancasterSound, Barrow Strait, fisheswere not identifiedby speciesor genus. and adjacentchannels, although this may, to a At PrinceLeopold Island diversityindices were certain extent, reflect differencesin sampling lower than any of thosecalculated for the low- intensity. arctic colonies (1975, 0.47; 1976, 0.91; 1977, 0.62). Differencesamong colonieswithin the low- Only 5 speciesof fish were identified among arctic zone suggestthat there is considerable 730 ^. J.GASTON [Auk,Vol. 102

1981 198• Z980 1981 1682 1982 I COATSIS.NOII OIGGESISLAND I HAOTZSCN[ 1981 1984 1980 1961 1982 1982 S.ND COATSISLAOO I I 01GGESIS.NO I I OANTZSCNI ISLAND • Capelin :•:::• Lanternfish• Arct,cCod • Olennies • San[•Lance • Sculpins • Capelin -.'-'--• Lanternfish•;•'?:r• Arct,cCod • Olennies • SandLance • Sculpms • Lumpsuckers• Snailfpsh• GreenlandHalibut • Non-fish • Non-fish Fig. 2. Proportions of different food types by number in the diets of Thick-billed Murre chicks at Fig. 3. Proportionsof differentfood typesin the three colonies. diets of Thick-billed Murre chicksby weight. regional variation in the availability of differ- (1982) found them to be an important compo- ent fish species.Capelin and sand lance, both nent in the diet of surface-feedingIvory Gulls midwater schooling species,were commonest (Pagophilaeburnea) in , not far from at Coatsand Diggesislands. These fish are typ- Hantzsch Island. Dunbar and Hildebrand (1952) ical of low-arctic waters off eastern Canada mentioned that Ungava natives were familiar (Dunbar and Hildebrand 1952). Stocks fluctu- with lanternfish, which sometimes occurred at ate considerablyover periods of several years the surface,"often deador dying." Clearly,they (Carscadden1984), and this may have account- occur at the surface on occasion, and hence ed for their rarity at Hantzsch and Akpatok is- their appearancein the diet of the tourres tells lands. us nothing about the tourres'foraging. Blenniesand sculpinsare bottom-living fish At Digges Island adult tourreswere collected (Leim and Scott 1968). If their presence indi- from nearby feeding areas during the chick- catesthat the tourres involved were feeding in rearing period in all three years. Analysis of shallow water, then we might have expected stomach contents yielded many fish otoliths, parallel variation in the samples.This was not the majority much smaller than those charac- the case. Only one blenny was found at Hantzsch Island, where sculpins were com- mon. Where either group was numerous, it TABLE2. Values of Morisita's Index (C•) for inter- year comparisonsat a single colony, showing that usually was represented by more than one interyear differenceswere small. species;hence, variation in year-classstrengths seemsunlikely to accountfor the intercolony Colony Years Cx variation observed. Probably, the two groups Digges Island 1980:1981 0.965 favor different water-depths or bottom-types, 1980:1982 0.992 but little information is available on the ecol- 1981:1982 1.034 ogy of noncommercialarctic marine fishes. Prince Leopold Island 1975:1976 1.007 1975:1977 1.000 Lanternfish, recorded only at Hantzsch Is- 1976:1977 1.016 land in 1983, generally are consideredto be Coats Island 1981:1984 1.012 bathypelagicfish. However, Orr and Parsons October1985] Foodof Thick-billedMurre Chicks 731

TABLE3. Numbers of different taxa recorded in the diet of Thick-billed Murre chicks at five colonies.

Prince Coats Digges Akpatok Hatzsch Leopold Taxon Is. Is. Is. a Is. Is.• Capelin (Mallotusvillosus) 59 13 38 0 0 Glacierlanternfish (Benthosema glaciale) 0 0 2 8 0 Arcticcod (Boreogadussaida) 81 I00 931 I0 139 Blennies (Blennioidei) 31 28 536 I 0 Sandlance (Ammodytes spp.) 3 27 36 2 0 Redfish (Sebastesmarinus) 0 0 I 0 0 Sculpins(Cottidae) 51 9 728 35 32 Smoothlumpfish (Cyclopteropsis jordan 0 0 0 0 0 1 Lumpsuckers(Eumicrotremus spp.) 0 2 24 0 0 Snailfish(Liparis spp.) I 0 45 3 1 Greenlandhalibut (Reinhardtiushippoglossoides) I 4 118 7 0 Squid(Gonatus fabricii) 2 I0 70 7 0 Crustacea 2 I 98 0 5 Annelida 0 0 3 0 0 Total 231 194 2,630 73 178 From Tuck and Squires (1955). From Gastonand Nettleship (1981). teristic of the fish delivered to chicks (Gaston only to foraging from a single center, and has and Noble 1985). Comparing only the most not yet been generalized to intercolony com- common fish, arctic cod, the median weights of parisons.The small size of fish at Digges Is- those found in adult stomachs(estimates from land, along with a lower rate of feeding, pre- otolith length) during the chick-rearingperiod sumablyexplains the low growth rate of chicks were 3.0, 4.4, and 3.7 g in 1980, 1981, and 1982, at this colony (Gaston et al. 1983). respectively.Comparable figures from the chick At large colonies,where birds travel long sampleswere 10.6, 7.8, and 4.5 g. This finding distancesto feed, a large proportionof the po- is consistentwith the predictionsof central- tential foragingarea is likely to be far offshore. placeforaging models (Andersson 1978, Orians We thereforemight expectthe diet of birds at and Pearson 1979). large coloniesto containa lower proportionof Gastonet al. (1983)suggested that birds from typicallycoastal fish than the dietsof birds at DiggesIsland foraged at greaterdistances from small colonies. The fact that bottom-living their colony than those at Hantzsch or Coats blennies and sculpins were more common at Island. However, the mean size of fish deliv- Hantzsch (50,000 pairs) and Coats islands ered at Digges Island was smaller than those (25,000 paris) than at Digges Island (with ad- delivered at the other two colonies, although jacent , 300,000 pairs) is not significantlyso. Superficially,this appears consistentwith this expectation.However, the to contradict the predictionsof central-place presentsample of coloniesis insufficientfor foragingtheory. However, the theory applies any firm generalizationto be possible.

TABLE4. Morisita's Index of similarity for samplesof Thick-billed Murre chick diets from five colonies (upper right), and valuesof Chi-squaretest for eachcombination (lower left).

Prince CoatsIs. Digges Is. Akpatok Is. Hantzsch Is. Leopold Is. Coats Is. -- 0.815 0.869 0.601 0.703 Digges Is. 79.34 -- 0.822 0.362 0.862 Akpatok Is. 381.3 210.8 -- 0.739 0.736 Hantzsch Is. 97.2 127.7 64.9 -- 0.424 Prince Leopold Is. 102.5 90.6 133.6 103.3 -- Mean Cx 0.747 0.715 0.791 0.531 0.681 All X2 values significantat P < 0.001. 732 A.J. GASTON [Auk,Vol. 102

ACKNOWLEDGMENTS GASTON,A. J. 1982. On the seabirds of northern . Naturaliste Can. 109: 895-903. I would like to thank everyone who participated , G. CHAPDELAINE,•' D. G. NOBLE. 1983. The in the C.W.S. Hudson Strait Seabird Project for as- growth of Thick-billed Murre chicksat colonies sistingme with collections,particularly David Noble in Hudson Strait: inter- and intra-colonyvaria- and Steven Smith. I am very grateful to Don Mc- tion. Can. J. Zool. 61: 2465-2475. Allister of the National Museums of Canada for iden- , & D. N. NET•LESHIP. 1981. The Thick-billed tifying, or confirmingidentification of, all the fish Murres of Prince Leopold Island. Can. Wildl. specimenmaterial. Hugh Boyd, Dick Brown, Tony Serv. Monogr. No. 6. Diamond, Richard Elliot, and Don McAllister com- ß & D. G. NOBLE. 1985. The diet of Thick- mented on earlier drafts of the paper. Logisticand billed Murres in West Hudson Strait and North- other support was supplied by Petro-Canada,the east Hudson Bay. Can. J. ZooL 63: 1148-1160. BaselineStudies Programme of EnvironmentCana- HUNTER,J. G., S. T. LEACH,D. E. MCALLISTER,& M. B. da, Polar Continental Shelf Project of the Depart- STEIGERWALD. 1984. A distributional atlas of ment of Energy,Mines and Resources,the Depart- records of the marine fishes of arctic Canada in ment of Indian and Northern Affairs, and the the National Museums of Canada and Arctic Bi- Canadian Coast Guard. ological Station. Syllogeus52. LEIM, A. H., & W. B. SCOTT. 1968. Fishes of the At- LITERATURE CITED lantic coast of Canada. Fisheries Res. Bd., Canada Bulletin 155. ANDERSSON,M. 1978. Optimal foraging area: size ORIANS,G. H., & N. E. PEARSON.1979. On the theory and allocationof searcheffort. Theoret.Pop. Biol. of centralplace foraging. Pp. 154-177in Analysis 13: 397-409. of ecologicalsystems (D. J. Horn, R. D. Mitchell, BIRKHFAD,T. R., & D. N. NETTLESHIP.1981. Repro- and G. R. Stairs, Eds.). Columbus, Ohio State ductive biology of Thick-billed Murres Uria lorn- Univ. Press. via, an inter-colony comparison.Auk 98: 258- ORR,C. D., & J. L. PARSONS.1982. IVOry GullsßPa- 269. gophilaeburnea, and ice edgesin Davis Strait and BROWN, R. G. B., D. N. NETTLESHIP,P. GERMAIN, C. E. the . Can. Field-Natur. 96: 323-328. TULL, & T. DAVIS. 1975. Atlas of eastern Cana- SALOMONSEN,F. 1972. Zoogeographicaland ecolog- dian seabirds. Ottawa, Canadian Wildl. Serv. ical problems in arctic birds. Proc. 15th Intern. CARSCADDEN,J. E. 1984. Capelin in the northwest Ornithol. Congr.:25-77. Atlantic. Pp. 170-183 in Marine birds: their feed- TRAMER,E.J. 1969. Bird speciesdiversity: compo- ing ecologyand commercialfisheries relation- nents of Shannon'sformula. Ecology50: 927- ships (D. N. Nettleship, G. A. Sanger, and P. F. 929. Springer,Eds.). Ottawaß Canadian Wildl. Serv. TUCK, L. M., & M. J. SQUIRES. 1955. Food and feed- DIAMOND,A. W. 1983. Feeding overlap in some ing habitsof Brunnich'sMurre (Uria lornvialorn- tropical and temperate seabird communities.Pp. via) on Akpatok Island. J. Fisheries Res. Board, 24-46 in Tropical seabirdbiology (R. W. Schrei- Canada 12: 781-792. ber, Ed.). Stud. Avian Biol. No. 8. WOLDA,H. 1981. Similarity indices, sample size and DUNBAR, M. J., & H. H. HILDEBRAND. 1952. Contri- diversity. Oecologia50: 296-302. bution to the study of the fishesof Ungava Bay. J. Fisheries Res. Board, Canada 9: 83-128.

APPENDIX1. Formulaefor estimatingfresh weights from length for taxa included in Fig. 3. (Specimensof other taxa were estimated from the mean weight of fresh specimensof the same taxon from the same samplein the preparationof Fig. 3.)

Taxon Regressionformula a r2 n Capelin W = (0.616 x L3/100)- 2.509 0.907 22 Arctic cod W = (0.642 x L3/100)- 0.536 0.921 40 Fish doctor W = (0.346 x L3/100) - 1.464 0.778 11 Sand lance W = (0.247 x L3/100) + 0.086 0.813 17 Sculpin spp. W = (0.790 x L3/100) + 0.576 0.904 21 Where W is in g and L in cm. October1985] Foodof Thick-billed Murre Chicks 733 734 A.J. GASTON [Auk, Vol. 102

APPENDIX3. Numbersof different prey taxa identified in six samplesof chick diets. Figuresin parentheses include those identified visually; these were the figures used in calculatingMorisita's indices.

CoatsIsland DiggesIsland HantzschIsland 1981 1984 1980 1981 1982 1982

Capelin (Mallotus villosus) 5 (7) 13 (52) 8 (8) 3 (3) 2(2) 1 (1) Glacier lanternfish (Benthosemaglaciale) (0) (0) (0) (0) (0) 4(8) Arctic cod (Boreogadussaida) 11 (24) 12(57) 40 (40) 47 (47) 13 (13) 9 (10) Fish doctor (Gymnelusviridis) 2 9 7 1 Fourline snakeblenny(Eumesogrammus praecisus) 3 2 2 Stout eelblenny (Anisarchusmedius) 1 Daubedshanny (Leptoclinus maculatus) 3 2 Slendereelblenny (Lumpenusfabricii) 1 1 All blennies (Blennioidei) 2 (6) (25) (13) (11) (2) (1) Northern sand lance (Ammodytesdubius) 2(3) (0) 18 (18) 7(7) 2(2) 1 (1) Arctic staghornsculpin (Gymnocanthustricuspis) 2 Spatulatesculpin (Icelusspatula) 1 Shorthornsculpin (Myoxocephalusscorpius) 4 Moustachesculpin (Triglopsmurrayi) I Bigeyesculpin (Triglopsnybelini) 4 1 18 Ribbed sculpin (Triglopspingelii) 2 9 All sculpins(Cottidae) 7 (10) (41) (2) (5) (1) (35) Leatherfin lumpsucker(Eumicrotremus derjugini) 1 Atlantic spiny lumpsucker(Eumicrotremus spinosus) 1 All lumpsuckers (0) (0) (0) (2) (0) (0) Gelatinoussnailfish (Liparisfabricii) (1) 3 (3) Greenland halibut (Reinhardtiushippoglosoides) (1) 2(2) 2 (2) (0) 5(7) Unidentified fish 1 1 2 Squid (Gonatusfabricii) (0) 1 (2) 3 (3) 6 (6) 1 (1) 5(7) Crustacea (0) (2) 1 (1) (0) (0) (0) Total 27(51) 45(180) 88(87) 85(83) 21(21) 56(73)