* Atlantic Puffin (Fratercula arctica)

Atlantic Puffin. Photo: omarrun


GENERAL ECOLOGY: 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 [16]. 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 [16]. 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 [16]. This species nests mostly in burrows but also in rock crevices and cavities, the nest being a scrape, often lined with grass and feathers [16].

FORAGING RANGE: Max 200 km, mean max 62.2 km, mean 30.35 km.

FORAGING DEPTH: Max 70 m, mean 37.03 m.


KEY HABITATS: Shallow waters, tidal fronts.

KEY PREY ITEMS: Mid-sized schooling mid-water fish, especially sandeels Ammodytes sp. (NE Atlantic) and capelin Mallotus villosus (Canada).



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’ [36]) and are abundant at the Aberdeen Front during both the breeding and non-breeding seasons [8]. 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 [37].

Important foraging associations with other species
None significant.

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 Sprattus sprattus, capelin, whiting Merlangius merlangus, Saithe Pollachius virens, Red-fish Sebastes marinus, Haddock Melanogrammus aeglefinus, herring Clupea harengus, Five-bearded Rockling Ciliata mustela, Northern Rockling C. septentrionalis, and Three-bearded Rockling Gaidropsarus vulgaris [26]. Sandeel are generally the preferred food items where available and can make up to 75-100% of diet in some locations [20]. 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 [27]. 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 Gadus morhua, 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 [6]. At times, they prey on juvenile demersal fishes, such as gadoids [19]. 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].

Foraging range
There is little direct information on the foraging range of Atlantic Puffins during the breeding season, and none for North America [26]. In one study, it is stated that puffins generally forage close (3-5 km) to breeding colonies [9]; 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 [18].
Of [14] foraging trips made by a single puffin fitted with a radio transmitter breeding at the Isle of May, [9] (64%) were within 2 km, 1 (7%) between 2-10 km and 4 (29%) more than 10 km [36]. 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 [36]. 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 [39].

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 [25]. 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 [38]. 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 [2]. 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 [39] 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 [10].

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) [31]. 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 [14]. 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 [32]. 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 [9].

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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) [22].
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 [13]. 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 [33]. Off Norway, dives of 10–45 m were recorded (n=6 birds [5]).

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

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Compiled by: Ben Lascelles, Nigel Varty, Kate Tanner, Rory McCann