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Kooyman Et Al: Foraging Patterns of Penguins Kooyman et al: Foraging patterns of Penguins https://www.internationalornithology.org/PROCEEDINGS_Durban/S... In: Adams, N.J. (et al): Proceedings of the 22nd International Ornithological Congress 1998, pp 2021-2039 (1999) S34.2: Foraging patterns of polar penguins G. Kooyman1, C. Hull2, O. Olsson3, G. Robertson4, J. Croxall5 & L. Davis6 1Scripps Institution of Oceanography, University of California, San Diego, La Jolla, California 92093-0204 USA , fax 619 534 1305, e-mail [email protected]; 2University of Tasmania, Hobart 7001, Australia; 3University of Uppsala, Sweden; 4Australian Antarctic Division, Kingston 7050, Tasmania, Australia; 5British Antarctic Survey, Madinghay Road, Cambridge CB3 0ET, UK; 6University of Otago, Dunedin, New Zealand Kooyman, G., Hull, C., Olsson, O., Robertson, G., Croxall, J. & Davis, L. 1999. Foraging patterns of polar penguins. In: Adams, N.J. & Slotow, R.H. (eds) Proc. 22 Int. Ornithol. Congr., Durban: 2021-2039. Johannesburg: BirdLife South Africa. Sub-antarctic and polar penguins have revealed important differences in the distances travelled to foraging areas, the physical and biological characteristics of foraging areas, and foraging patterns. Differences are associated with preferred prey and its abundance. Data were acquired using satellite transmitters and time/depth recorders, the former giving location and rates of travel, the latter diving depths and patterns. Distinctions between travel and feeding dives help to assess foraging success. Data were matched to satellite imagery for determination of sea surface conditions. Sub-antarctic penguins travel further than polar penguins, feed near the Antarctic polar front, and are primarily diurnal feeders. Polar species feed at edges of coastal ice, pack ice, and polynyas. Most locations are neritic. Adelies specialise in krill at shallow depth as do sub-polar Macaroni and Royal Penguins. Emperor Penguins target fish in the mesopelagic zone, which is true also of King Penguins feeding at the Antarctic Polar Front. Hunting with 24h of daylight, polar penguins feed continuously with little hourly variation in depth. Subantarctic penguins show considerable diet differences, with reduced feeding at night. INTRODUCTION Penguins are unique among birds in their degree of adaptation to life in a marine environment possessing more complex structural and physiological adaptations for this than any other group of birds. Given these specialisations for life at sea, coupled with potential restrictions on distribution due to flightlessness and the high energy demands of very small marine homeotherms, we might expect penguins to be particularly good as indicators of habitats where prey are concentrated and available for capture. In this paper, we try to characterise the principal feeding habitats of a range of polar penguin species whose foraging ecology has been the subject of recent study. The data acquired in the last decade on penguin foraging ecology has represented a major breakthrough in the study of marine predators, revolutionising our knowledge of their capabilities and revising our ideas on many aspects of marine science and conservation. We have selected five of the 17 species of penguins to discuss in detail their foraging habits. All penguins are cold temperate or polar in their distribution, but the five chosen are especially connected to the polar or sub-polar environment. This review is based on some common factors among these five: (1)They are some of the most successful of all the species in terms of abundance and biomass (Woehler 1993); (2)They are also some of the best known because of the detailed studies that have been conducted. For our report this specifically includes the use of satellite transmitters to determine geographic distribution, and time depth recorders to determine hydrographic distribution. (3) At some stage in their life cycle all forage at or near the Antarctic Polar Front (APF), also known as the Polar Front Zone (PFZ). These definitions exclude two species that breed below the APF and commonly, if not exclusively, feed in Antarctic waters. These are the Gentoo, Pygoscelis papua, and Chinstrap, P. antarctica, Penguins. The Gentoo Penguin is not considered because its foraging range is typically restricted to the vicinity of its breeding sites at all times of year (Williams 1991; Wilson et al.1998). In contrast, Chinstrap Penguins may meet the above criteria but no satellite-tracking distribution studies have yet been undertaken. A very recent study using geolocation devices suggests a winter-at-sea distribution adjacent to the marginal ice zone (Wilson et al. 1998). Three of the species Adelie, P. adeliae, Royal, Eudyptes schlegeli, and Macaroni, E. chrysolophus, Penguins, are surface feeders seldom descending below 50 m to feed mainly on krill. The King, Aptenodyptes patagonicus, and Emperor, A.forsteri, Penguin frequently feed in the mesopelagic region of 200 to 400 m, catching mainly fish. 1 von 14 28.06.2021, 10:20 Kooyman et al: Foraging patterns of Penguins https://www.internationalornithology.org/PROCEEDINGS_Durban/S... Distribution, breeding and diet The congeneric Macaroni and Royal Penguins have very similar breeding biology and ecology (Croxall 1984). The range of the Macaroni Penguin is the sub-Antarctic islands of the Atlantic and Indian Ocean sectors of the Southern Ocean (with the exception of Macquarie Island where the Royal Penguin is endemic), with small outlying populations in South America and on the islands of the Antarctic Peninsula. Breeding commences in September (Royal) and October/November (Macaroni) with males returning to colonies a week or so before females (Fig.1). After the courtship period, incubation (35 days) and brooding (20-25 days) involves three long (10-20 day) shifts, alternately by males and females. There is bi-parental care during chick rearing with average foraging trips usually in the range 10-48h in Macaroni Penguins. The shortest Royal Penguin foraging trips are for three days during the guard stage. Later in creche the trips lengthen again. After the 55-60 day chick-rearing period, adults feed at sea for 2-4 weeks, returning to the breeding colony to moult (which lasts 20-25 days) before departing to sea for the winter. Both species take mainly planktonic crustaceans (especially euphausids) and small epipelagic fish. Royals feed primarily on myctophid fish, dominated by Krefftichthys anderssoni, and euphausiids, particularly Euphausia vallentini. Prey items are those associated with the PFZ. Inter-annual differences in diet, degree of digestion of prey and quantity of food brought ashore suggest differences in distribution of prey between years. Diet is consistent across the breeding season in both prey species. About 51% by mass of the diet is euphausiids, and 32% of that mass is made up of E. vallentini. Another 41% by mass consists of myctophids, of which nearly 24% is K. anderssoni (Hindell 1988). Macaroni Penguins tend to specialise in Antarctic krill Euphausia superba at South Georgia (Croxall & Prince 1980; Croxall et al. 1997) but populations in the Indian Ocean have more diverse diets. King Penguins breed on most sub-Antarctic islands from the Falkland Islands east to Macquarie Island. They are most abundant in the Crozet Archipelago, Indian Ocean sector of the southern ocean. The King Penguin breeding cycle, which was first described by Stonehouse (Stonehouse, 1956; Stonehouse 1960) at South Georgia, starts in early spring with the moult which takes about a month to complete (Fig. 1). Then courtship and egg-laying take place normally during November and December. The single chick hatches in January/February and is fed until the beginning of the winter, by which time it can be as heavy as an adult. However, the chick remains in the colony through the winter. During the overwinter food shortage chicks fast, but feeding is resumed in the spring for some months until the chick becomes independent. Thereafter the parents moult and soon after may begin a new breeding cycle, although considerably later than in the previous year. Consequently, the King Penguin has one of the most unusual breeding time tables among birds (Olsson 1996). The total time required for these activities exceeds that available during a single summer (Olsson 1996), and the result is that a breeding and moult cycle takes more than one year to complete. The diet of King Penguins is very consistent. In all areas for which there is information on food habits the main food item by mass are myctophids. In this group the principal species is K. anderssoni. Diet analyses completed at South Georgia, Marion, Crozet, and Macquarie all show the same result (Adams & Klages 1987; Hindell 1988; Cherel et al. 1996; Olsson & North 1997). In short, myctophids are a basic food item of King Penguins, and probably many other predators in the PFZ. Emperor Penguins are known to breed almost completely around the Antarctic Continent as well as far south into the Weddell and Ross Seas. There are significant gaps in their distribution along the coastline of the Bellingshausen and Amundsen Seas, the east side of the Antarctic Peninsula, and Wilkes Land Coast. Emperor Penguins begin courtship in the fall (April), and lay in late May to early June (Williams 1995) (Fig. 1). The larger male assumes all incubation duties and about 70 days later, and a few days after hatching, the female returns to relieve the male in late July to mid-August. Over the next five months the female and male come and go to the colony about 10 times to feed the chick. Each trip away tends to get shorter, declining from about three weeks to three days (Kirkwood & Robertson 1997). Around summer solstice the chicks fledge by leaving the colony, and the parents depart to moult (Kooyman et al. 1990). The moult lasts about 35 days. The primary foods of Emperor Penguins are krill, fish, and squid, which vary according to colony location and season (Cherel & Kooyman 1998; Kirkwood & Robertson 1997; Klages 1989; Putz 1995; Robertson & Newgrain 1996; Robertson et al.
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