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* Atlantic Puffin (
Atlantic Puffin. Photo: omarrun
: This species winters largely offshore, sometimes beyond the continental shelf into pelagic zones. Post-breeding patterns of dispersal are varied and complex, more work is needed to identify the main wintering areas and comprehensively describe dispersal patterns . This species is exclusively marine, occurring along rocky coasts and offshore islands, breeding variably on grassy maritime slopes, flattish ground, sea cliffs and boulder fields. In winter, birds are wide ranging in offshore and pelagic habitats, usually within the bounds of the continental shelf . Its diet consists primarily of fish throughout the year, supplemented in spring and summer by crustaceans, polychaetes and squid. Individuals feed by underwater pursuit diving . This species nests mostly in burrows but also in rock crevices and cavities, the nest being a scrape, often lined with grass and feathers .
: Max 200 km, mean max 62.2 km, mean 30.35 km.
: Max 70 m, mean 37.03 m.
FORAGING TRIP DURATION
: Mean 2.46 h (n=2)
: Shallow waters, tidal fronts.
KEY PREY ITEMS
: Mid-sized schooling mid-water fish, especially sandeels Ammodytes sp. (NE Atlantic) and capelin Mallotus villosus (Canada).
KEY FORAGING ASSOCIATIONS
: None significant.
General feeding behaviour
Atlantic Puffins are pursuit-divers that use their wings to propel themselves through the water. Birds searching for prey commonly dip their head in the water and surface-dive once prey is spotted 15.
Foraging habitat - breeding season, migration and wintering period
Atlantic Puffins are primarily found in pelagic waters, except when breeding, when they forage in fairly shallow waters [9, 26], and are considered relatively little-studied at sea other than in low-arctic and boreal zones of the North Atlantic [11, 24, 30].
Tidal fronts are believed to be important foraging zones for puffins (and many other seabirds) with prey being brought to the surface by flow gradients. Puffins breeding on the Isle of May, Scotland were found to be associated with tidal fronts (or ‘rips’ ) and are abundant at the Aberdeen Front during both the breeding and non-breeding seasons . Transect surveys around the Isle of May found that puffins were found mostly close to the island, but were also associated with sand banks further offshore e.g. the Wee Bankie, although to a lesser extent than other auks .
Important foraging associations with other species
Atlantic Puffins feed predominantly on small to mid-sized (5–15 cm length), schooling mid-water fish. A wide range of prey items are taken, but when available high calorific value species are preferred. Several studies have found marked differences in diet between colonies and between years [14, 17, 19, 27] usually linked to reduced availability of principal prey items causing a switch in foraging behaviour [4, 29]. Principal prey items may also vary between the early and late chick-rearing periods.
Typical prey species in U.K. include sandeel, sprat
, capelin, whiting
, Five-bearded Rockling
, Northern Rockling
, and Three-bearded Rockling
. Sandeel are generally the preferred food items where available and can make up to 75-100% of diet in some locations . Sprat (up to 70%), capelin (up to 80%), rockling (up to 42%) and herring (up to 40%) are also important, with a wide range of other species forming smaller proportions of the diet, including a variety of demersal fish in their young pelagic stages. On the Isle of May, for instance, sandeels comprised 50-93% of all prey brought to chicks [21, 23]. The remainder was mainly sprat with the exception of 1980-1982 and 1984 when herring made up 14-38% of the diet. Similarly, sandeels comprised more than 79% of puffin diet by weight on Hermaness, U.K. from 1973-1986, but then declined to 19% in 1987 and 36% in 1988 during a period of low sandeel abundance in which most chicks starved . Over twelve other species of fish were also recorded but only rockling and Haddock formed more than 10% of the diet by weight in any year prior to 1987. In 1988, rockling formed 42% of the diet and other gadoids (mostly Saithe) 21%. The differences in diet between studies and years shows that Atlantic Puffins will switch prey, readily taking fish of reduced calorific value and of smaller size in years when the preferred species are less available.
In Norway, Atlantic Puffins generally harvest small pelagic species (c. 3-8 cm in length) such as sandeels, capelin and herring, or the youngest pelagic stages of demersal fish such as cod
, Saithe and Haddock [1, 4, 6, 7].
In the north-west Atlantic, puffins prey on a variety of species, including juvenile pelagic fishes, such as herring, juvenile and adult capelin and sandeels . At times, they prey on juvenile demersal fishes, such as gadoids . In eastern Canada, capelin has a key position in all marine food webs, and comprised 80% or more of the Atlantic Puffin and common guillemot chick diets in years when capelin stocks were high [12, 28].
There is little direct information on the foraging range of Atlantic Puffins during the breeding season, and none for North America . In one study, it is stated that puffins generally forage close (3-5 km) to breeding colonies ; whereas a separate study reports that adults usually forage within 10 km of their colony when feeding chicks, but may range as far as 50-100 km .
Of  foraging trips made by a single puffin fitted with a radio transmitter breeding at the Isle of May,  (64%) were within 2 km, 1 (7%) between 2-10 km and 4 (29%) more than 10 km . All its feeding trips were to the south or east of the colony, but the bird used quite separate areas both on the same and on different days. The bird fed frequently in a tide rip 0.5-1 km from the colony . Transects around the Isle of May found that puffins were most common close to the colony but on some occasions birds occurred at a sandbank, the Wee Bankie, 40 km away. foraging trips studied using GPS loggers at the isle of may found birds made two types of trips: long absences where birds were away overnight (15-41 hrs) and travelled a maximum of 37.8-65.5km from the colony; and shorter excursions (0.7-3.8hrs) reaching a maximum of 9.3-17.5km from the colony .
The maximum distance from St. Kilda, U.K. at which a puffin with a fish has been recorded flying towards the colony is 40 km, with many others seen feeding in the same area . The greatest densities of puffins around the Flamborough Head colony in north east England were recorded 26-28 km away in the morning, whereas later on in the day the highest densities occurred at 6 km and 8 km, and further out at 40 km . Some fish were also carried in from greater distances. Puffins were found within 35-40 km of Skomer and Skokholm, Wales in June 1990 and 1992, with most birds seen after 9.30 a.m. within 10 km of the colonies [34, 35]. Other authors suggest most birds feed within 7 km of colony off Wales [3, 14, 23]. Puffins at Røst Island, Norway, travelled at least 137 km to fishing grounds after a crash of herring stocks nearer to the colony . Studies in Faroe Islands suggested that when feeding conditions were poor birds used more distant feeding areas, some travelling from as much as 250km away  Based on observations from the shore and from a research vessel on foraging activities of puffins in waters adjacent to the breeding colony at Coquet Island, U.K., one study noted that puffins fished up to 20-25 km offshore but tended to forage closer to Coquet Island in July, when chicks were fledging, than in June, before fledging occurred .
On the basis of a mean trip length of 207 min (n=9) during chick-rearing and a flight speed of 80 km/h, it was estimated that puffins breeding on the Farne Islands, north east England could forage at a maximum distance of 137 km from the colony if they spent a negligible amount of time actually fishing (ie giving a theoretical maximum range) . In another study, it was estimated that the theoretical maximum foraging range of puffins from Skomer, Wales was around 32 km on the basis of a mean trip duration of 85.5 min (n=62 trips in 1969) and a flight speed of 48 km/h . However, a boat transect run on one day in 1970 recorded 85% of all puffins within just 3 km of the island, with none of those recorded further afield carrying a fish. However, it is worth noting that mean observed airspeed of puffins recorded using an ornithodolite was actually c. 63 km/h . On the basis of average trip durations at U.K. colonies (various sources) and flight speeds of 48 km/h or 82 km/h one study suggested a maximum range of 49-83 km from Hermaness, Shetland, U.K. 36-61 km from St. Kilda, 57-97 from the Isle of May and 34-58 km from Skomer, and a potential range of over 200 km from Great Island, Newfoundland, Canada .
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Table 1: Showing average value (km) of foraging range in different country/sea areas.
Figure 1: Showing cumulative frequency (with standard deviation) and proportion of birds found foraging at different distances from colony. Source: Birdlife Seabird Foraging Range Database
Diving and depth association
Atlantic Puffins catch most of their prey within 30 m of the water surface, although they are capable of diving to 60 m [13, 26, 33]. Using depth gauges, one study recorded dive depths of 20-30 m (deepest dive 33 m, median 25.5 m, n=6) .
Maximum diving depths of Atlantic Puffins fitted with depth gauges recorded off Newfoundland, Canada were 40–68 m (n=10 birds), and only one bird exceeded 60 m (diving to 68 m) in 17 days of foraging . Records of birds from bycatch mortality in gill-nets most often occurred in the 1 m to 10 m depth-class (144 of 171), and none were caught in nets set deeper than 60 m . Off Norway, dives of 10–45 m were recorded (n=6 birds ).
Figure 2. Showing cumulative frequency (with standard deviation) and proportion of birds found within foraging depths around the colony. Source: Birdlife Seabird Foraging Range Database
1. Anker-Nilssen, T. (1987) The breeding performance of puffins Fractercula arctica on Røst, northern Norway in 1979–1985. Fauna Norv Ser C Cinclus 10:21–38.
2. Anker-Nilssen, T. and Lorentsen, S.-H. (1990) Distribution of Puffins Fratercula arctica feeding off Røst, northern Norway, during the breeding season, in relation to chick growth, prey and oceanographical parameters. Polar Res. 8:67–76.
3. Ashcroft, R.E. (1976) Breeding, biology and survival of Puffins. D.Phil. diss., Oxford Univ., Oxford, U.K.
4. Barrett, R.T. (2002) Atlantic Puffin Fratercula arctica and common guillemot Uria aalge chick diet and growth as indicators of fish stocks in the Barents Sea. Mar. Ecol. Prog. Ser. 230:275-287.
5. Barrett, R.T. and Furness, R.W. (1990) The prey and diving depths of seabirds at Hornøy, north Norway after a decrease in the Barents Sea capelin stocks. Ornis Scand. 21:179–186.
6. Barrett, R.T., Anker-Nilssen, T., Rikardsen, F., Valde, K., Røv, N. and Vader, W. (1987) The food, growth and fledging success of Norwegian puffin chicks Fractercula arctica in 1980-1983. Ornis Scand 18:73–83.
7. Barrett, R.T., Becker, P.H., Furness, R.W., Hunt, G.L.Jr., Latrouite, D., Montevecchi, W.A., Olsen, B., Skov, H., Tasker, M.L. and Wright, P. (1994) Seabird prey harvests in the North Sea. Seabird/Fisheries Interaction Study Group Rep ICES (Copenhagen) CM 1994.
8. BirdLife International (2000). The Development of Boundary Selection Criteria for the Extension of Breeding Seabird Special Protection Areas into the Marine Environment. OSPAR Convention for the Protection of the Marine Environment of the North-East Atlantic. Meeting of the Biodiversity Committee (BDC) Vlissingen (Flushing): 20 – 24 November 2000.
9. Bradstreet, M.S.W. and Brown, R.G.B. (1985) Feeding ecology of the Atlantic Alcidae. Pp. 264–318 In: Nettleship, D.N. and Birkhead, T.R. (Eds.) The Atlantic Alcidae: evolution, distribution and biology of the auks inhabiting the Atlantic Ocean and adjacent water areas. Academic Press, London, U.K.
10. Breakwell, K.J., Denny, M.J., Evans, S.M. Noble-Rollin, D.C. and Redfern, C.P.F. (1996) Foraging distributions of terns and puffins in coastal waters off Coquet Island (Northumberland). Transactions of the Natural History Society of Northumbria 57:13-20.
11. Brown, R.G.B. (1985) The Atlantic Alcidae at sea. Pp. 384–426 In: Nettleship, D.N. and Birkhead, T.R. (Eds.) The Atlantic Alcidae: evolution, distribution and biology of the auks inhabiting the Atlantic Ocean and adjacent water areas. Academic Press, London, U.K.
12. Brown, R.G.B., Nettleship, D.N. (1984) Capelin and seabirds in the northeast Atlantic. Pp 184-194 In: Nettleship, D.N., Sanger, G.A. and Springer, P.F. (Eds.) Marine birds: their feeding ecology and commercial fisheries relationships. Can Wildl Serv Spec Publ, Ottawa.
13. Burger, A.E. and Simpson, M. (1986) Diving depths of Atlantic Puffins and common murres. Auk 103:828-830.
14. Corkhill, P. (1973) Food and feeding ecology of puffins. Bird Study 20:207-220.
15. Cramp, S. and Simmons, K.E.L. (1983). The birds of the Western Palearctic. Vol. 3. Oxford University Press, Oxford.
16. del Hoyo, J., Elliott, A. and Sargatal, J. (1996) Handbook of the birds of the world, Vol 3: Hoatzin to Auks. Barcelona, Spain: Lynx Edicions.
17. Furness, R.W. (1982) Population, breeding biology and diets of seabirds on Foula in 1980. Seabird 6:5-12.
18. Harris, M.P. (1984) The Puffin. Poyser, London.
19. Harris, M.P. and Hislop, J.R.G. (1978) The food of young puffins Fratercula arctica. Journal of Zoology, London 185:213-236.
20. Harris, M.P. and Riddiford, N.J. (1989) The food of some young seabirds on Fair Isle in 1986-88. Scottish Birds 15:119-125.
21. Harris, M.P. and Wanless, S. (1986) The food of young razorbills on the Isle of May and a comparison with that of young guillemots and puffins. Ornis Scandinavica 17:41-46.
22. Harris, M.P., Towll, H., Russell, A.F. and Wanless, S. (1990). Maximum dive depths attained by auks feeding young on the Isle of May, Scotland. Scottish Birds 16:25-28.
23. Hislop, J.R.G. and Harris, M.P. (1985) Recent changes in the food of young puffins Fratercula arctica on the Isle of May in relation to fish stocks. Ibis 127:234-239.
24. Huettmann, F. (2000) Environmental determination of seabird distribution patterns off eastern and arctic Canada. Ph.D. diss., Univ. of New Brunswick, Fredericton.
25. Leaper, G.M., Webb, A., Benn, S., Prendergast, H.D.V., Tasker, M.L., and Schofield, R. (1988) Seabird studies around St Kilda, June 1987. Nature Conservancy Council, CSD Report No. 804.
26. Lowther, P.E., Diamond, A.W., Kress, S.W., Robertson, G.J. and Russell, K. (2002) Atlantic Puffin (Fratercula arctica), The Birds of North America Online (A. Poole, Ed.). Ithaca: Cornell Lab of Ornithology; Retrieved from the Birds of North America Online:
27. Martin, A.R. (1989) The diet of Atlantic Puffin Fratercula arctica and northern gannet Sula bassana at a Shetland colony during a period of changing prey availability. Bird Study 36:170-180.
28. Montevecchi, W.A. (2000) Seabirds. In: Bundy A, Lilly G, Sheldon P (Eds.) A bulk-biomass model of the Newfoundland and Labrador Shelves. Can Tech Rep Fish Aquat Sci 2310:15–18.
29. Nettleship, D.N. (1991) The diet of Atlantic Puffin chicks in Newfoundland before and after the initiation of an international capelin fishery, 1967–84. Pp 2263-2271 In: Bell, B.D. (Ed.) Proc 20th Int Ornitholog Congress. New Zealand Ornitholog Congress Board: Wellington
30. Nettleship, D.N. and Evans, P.G.H. (1985) Distribution and status of the Atlantic Alcidae. Pp. 54–154 In: Nettleship, D.N. and Birkhead, T.R. (Eds.) The Atlantic Alcidae: evolution, distribution and biology of the auks inhabiting the Atlantic Ocean and adjacent water areas. Academic Press, London, U.K.
31. Pearson, T.H. (1968) The feeding ecology of sea-bird species breeding on the Farne Islands, Northumberland. Journal of Animal Ecology 37:521-552.
32. Pennycuick, C.J. (1987) Flight of auks (Alcidae) and other northern seabirds compared with southern Procellariiformes: ornithodolite observations. Journal of Experimental Biology 128:335-347.
33. Piatt, J.F. and Nettleship, D.N. (1985) Diving depths of four alcids. The Auk 102:293-297.
34. Stone, C.J., Harrison, N.M., Webb, A. and Best, B.J. (1992) Seabird distribution around Skomer and Skokholm Islands, June 1990. JNCC, Report No. 30.
35. Stone, C.J., Webb, A., Barton, T.R. and Gordon, J.R.W. (1993) Seabird distribution around Skomer and Skokholm Islands, June 1992. JNCC, Report No. 152.
36. Wanless, S., Harris, M.P. and Morris, J.A. (1990) A comparison of feeding areas used by individual common murres (Uria aalge), razorbills (Alca torda) and an Atlantic Puffin (Fratercula arctica) during the breeding season. Colonial Waterbirds 13:16-24.
37. Wanless, S., Harris, M.P. and Greenstreet, S.R. (1998) Summer sandeel consumption by seabirds breeding in the Firth of Forth, southeast Scotland. ICES J. Marine Science 55:1141-1151.
38. Webb, A., Tasker, M.L. and Greenstreet, S.P.R. (1985) The distribution of guillemots (Uria aalge), razorbills (Alca torda) and puffins (Fratercula arctica) at sea around Flamborough Head, June 1984. Nature Conservancy Council, CSD Report No. 590.
39. Harris M.P., Bogdanova M.I., Daunt F. and Wanless S. (2012). using GPS technology to assess feeding areas of Atlantic Puffins F
. Rining and Migration 27: 43-49
Compiled by: Ben Lascelles, Nigel Varty, Kate Tanner, Rory McCann
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