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Diving Patterns in Relation to Diet of Gentoo and Macaroni Penguins At The Condor 90:157-167 0 TheCooper Ornithological Society 1988 DIVING PATTERNS IN RELATION TO DIET OF GENT00 AND MACARONI PENGUINS AT SOUTH GEORGIA ’ J. P. CROXALL British Antarctic Survey,Natural EnvironmentalResearch Council, High Cross,Madingley Road, CambridgeCB3 OET, UK R. W. DAVIS Sea World ResearchInstitute, Hubbs Marine ResearchCenter, 1700 South ShoresRoad, San Diego, CA 92109 M. J. O’CONNELL British Antarctic Survey,Natural EnvironmentalResearch Council, High Cross,Madingley Road, CambridgeCB3 OET, UK Abstract. The depthsattained on 1,444dives by 14 GentooPenguins (Pygoscelis Papua) and 6,352dives by eightMacaroni Penguins (Eudyptes chrysolophus) were recorded,together with the timing and durationof the foragingtrip and the amountand type of prey caught. MacaroniPenguins ate only Antarctickrill Euphausiasuperba. When feedingonly at night they madeno divesdeeper than 20 m; on all-day trips 36% of diveswere between 20 to 80 m. GentooPenguins fed during the day.When theycaught krill, 77%of diveswere shallower than 54 m; when fish were taken, 59% of dives were 54 to 136 m, which is consistent with the benthic-demersalhabit of thejuvenile Nototheniaand Champsocephalusfish they eat. The patternof predationon krill by both penguinspecies is consistentwith its vertical migrationto the surfaceat night and dispersalthrough the water column during the day. The food requirements of chick-rearing Macaroni Penguinswould be met by catching at least six adult krill per dive (or 150 juvenile krill or amphipods). For similar Gentoo Penguins,a minimum of 15 adult krill per dive (one every 8 set), or one fish every third dive, is needed. Recorded interannual variations in krill size can treble these rates, which would also be doubled if half the dives were for travelling, not feeding. Key words: GentooPenguin; Macaroni Penguin; PygoscelisPapua; Eudyptes chrysolo- phus;Antarctic krill; Euphausiasuperba; fish; diving;diet; foraging; prey capturerate. INTRODUCTION In none of thesestudies was it possibleto mea- sure accuratelyboth the time that the birds spent Penguins are the most highly adapted of all birds at sea and also the quantity and identity of food for life under water. However, little is known caught. In this study we obtained the first data about their diving patterns and performance, on the diving patterns of Gentoo (Pygoscelispa- which are fundamental aspectsof their feeding pua) and Macaroni (Eudypteschrysolophus) pen- ecology,influencing the prey they encounter and guins and examine these in relation to foraging their efficient exploitation of these. Recent re- trip timing and duration and to the type and views of feeding ecologyand diving performance amount of food obtained. in penguins (Croxall and Lishman 1987, Kooy- Fieldwork was carried out at Bird Island, South man and Davis 1987) indicate that the depths Georgia (54’S, 38”W), as part of an extensive typically reached during diving by free-living study of the feeding ecology and energy budgets penguins are known for only five species.Only ofthese species(Croxall and Prince 1980a; Davis for King Penguins (Aptenodytes patagonicus), et al. 1983; Davis, Croxall, and O’Connell, un- Chinstrap Penguins (Pygoscelisantarctica), and publ.), in the austral summers of 1984 to 1985 JackassPenguins (Spheniscusdemersus) are data and 1985 to 1986. Macaroni and Gentoo pen- other than maximum depth attained available guins are the two commonest speciesof penguin (Kooymanetal. 1982,LishmanandCroxalll983, at South Georgia (with breeding populations es- Wilson 1985). timated at about 5,000,OOOand 100,000 pairs respectively; Croxall et al. 1984) and they are LReceived 20 March 1987. Final acceptance2 Oc- also abundant at Bird Island (breeding popula- tober 1987. tions of ca. 75,000 and 6,000 pairs respectively; [1571 158 J. P. CROXALL, R. W. DAVIS AND M. J. O’CONNELL Croxall and Prince 1980b). Both species are FORAGING TRIP DURATION therefore important consumers of marine re- When DHRs were being deployed, birds bearing sources at South Georgia, Macaroni Penguins thesewere watched until they entered the seaand being preeminent among avian species(Croxall a continuous watch was then maintained during et al. 1984, Croxall et al. 1985, Croxall and Prince daylight hours until the bird returned. Birds were 1987) but there are numerous biological and eco- weighed before and after the foraging trip. In logical differences between the two species. In both study seasonsthe attendancepatterns ashore the present context the most important of these of a sample (n = 10 to 15 birds originally) of are that Gentoo Penguins mainly feed inshore breeding birds of each specieswas recorded con- (within 10 km of land; Croxall and Prince 1987; tinuously throughout most of the chick-rearing J. L. Bengtson, unpubl. data) and attempt to rear period using radio transmitters and automatic two chicks, which are fed on adult Antarctic krill recording apparatus. The radio transmitters (15 (Euphausiu superba) and young stagesof mainly mm diameter, 50 mm length, 25 g weight) were benthic-demersal notothenioid fish (Croxall and attachedto the penguins’ back featherswith epoxy Prince 1980a, Croxall and Lishman 1987). In resin and hose clamps or plastic tie wraps. To contrast, Macaroni Penguins feed mainly off- record the presenceof these birds at the colony shore, rarely within sight of land, and rear one a Yagi 4-element antenna was mounted on the chick on a diet of both adult and juvenile E. roof of a small wooden blind and connected to superba, fish being very rarely recorded (Croxall a radio receiver (164 MHz) inside, set to scan and Prince 1980a; Croxall et al., in press). continuously in sequencethe frequencies of the METHODS deployed transmitters for 10 set each. The re- ceiver was connected to a strip chart recorder. DIVING DEPTH This systemprovided control data against which The depth histogram recorder (DHR) described the foraging trip durations of the experimental by Kooyman et al. (1983) was used. In brief, this birds could be compared. It also confirmed that is cylindrical (95.0 mm long by 23.0 mm di- penguins do not arrive ashore at the breeding ameter) and weighs 95 g (approximately 1.6% of colony during the night. Data on the foraging- adult Gentoo Penguin and 2.3% of adult male attendance patterns during postbrooding chick- Macaroni Penguin body weight). The DHR con- rearing of paint-marked birds (Croxall, unpubl. tains eight electronic counters which respond to data), whose presencewas recorded on three to different presetpressure thresholds, each of which four visits (of 1 hr each) daily, are consistentwith is equivalent to a different depth. When the pen- those of birds with radios. Thus paint-marked guin dives below the surface of the water, the Gentoo Penguinsspent about 12 hr at sea(range = increase in pressure with depth is converted to 6 to 24 hr; n = 2 10) and Macaroni Penguins a voltage shift by a pressure transducer in the either 12 hr (range = 10-l 6 hr; n = 140) or 24 DHR. If the voltage exceedsthe threshold of one hr (range = 18-50 hr; n = 72). or more of the counters, it is recorded in each of them. The settingof thesethresholds was checked FOOD in the laboratory against a pressurestation before The contents of the stomachs of experimental and after each deployment. The DHR was at- birds returning ashorewere collectedusing stom- tached to feathersin the middle of the back with ach lavage techniques (Wilson 1984). Birds were two small metal hose clips. Each bird was also flushed repeatedly until no more food was ob- given a distinctive paint mark (number or sym- tained; we have confirmed that this corresponds bol) on the breast to facilitate recapture on its to an empty stomach. In 1985 five complete return to the colony. Each recorder was used for stomach samples and in 1986 10 such samples a single foraging trip only, removed when the were obtained weekly from both speciesthrough- bird was recaptured, and the number of dives out the chick-rearing period. These samples en- registered in each counter was read through a abled us to check that experimental birds were decoding processor which was attached to the taking broadly similar prey and had a similar DHR in the laboratory. Birds were selectedafter mass of food in the stomach to other individuals they had fed a chick and when it wasjudged that engaged in similar parental duties at the same they were about to leave the colony to return to stageof the chick-rearingperiod. For control birds sea. in both seasonsand for experimental birds in DIVING AND DIET IN GENT00 AND MACARONI PENGUINS 159 1986, complete stomach contents were weighed Gentoo Penguin fresh at the time of collection and sorted on the same day. Krill (and other crustaceans),fish and Krill Fish squid were sorted,weighed, and preservedin 10% formalin. Both sagittal otoliths were removed from intact fish crania and these, together with -z any loose otoliths, counted and stored dry in plastic tubes. All material was then returned to the United Kingdom for further processing.All n=862 dives krill in samples were counted, including speci- mens representedonly by pairs of eyes.A sample of 100 krill from each stomach was measured, 0 25 50 either the straightened standard length of per- Percent of dives fectly intact specimensor the carapacelength of damaged ones and sex and reproductive status FIGURE 1. Dive depth histograms (proportion of was recorded for as many specimensas possible. dives terminating in each stratum) for Gentoo Pen- Fish otoliths were identified to as low a taxo- guins feeding on fish and krill. nomic categoryas possible, length, breadth, and thickness measured (to 0.01 mm) and weighed by weight) and amphipods Themisto gaudichau- (to 0.001 g). These measurements were used in dii (ca. 20 mm long; 2% by weight). Gentoo Pen- regression equations to estimate the standard guins ate either krill(9 1% by weight, occasionally length and weight of the original fish specimens with one or two fish present) or fish (87% by (using principally data from Hecht [ 19871 and weight, usually with a few krill).
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