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Common Tern

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Common Tern
Conservation status
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Aves
Order: Charadriiformes
Family: Sternidae
Genus: Sterna
Species: S. hirundo
Binomial name
Sterna hirundo
Linnaeus, 1758

The Common Tern (Sterna hirundo) is a seabird of the tern family Sternidae. This bird has a circumpolar distribution, its four subspecies breeding in temperate and subarctic regions of Europe, Asia and North America. It is strongly migratory, wintering in coastal tropical and subtropical regions. Breeding adults have light gray upperparts, white to very light gray underparts, a black cap, orange-red legs, and a thin, sharp, bill. Depending on the subspecies, the bill may be black-tipped red or all-black.

Contents

[edit] Taxonomy

The terns, family Sternidae, are small to medium-sized seabirds closely related to the gulls, skimmers and skuas. They are gull-like in appearance, but typically have a lighter build, long pointed wings (which give them a fast, buoyant flight), a deeply forked tail and long bulgy legs. Most species are grey above and white below, and have a black cap which is reduced or flecked with white in the winter.[2]

The Common Tern was first described by Linnaeus in his Systema Naturae in 1758 under its current scientific name, Sterna hirundo.[3] Its closest relatives appear to be the other Eurasian members of Sterna, to which it may be ancestral.[4]

The genus name, Sterna, is a latinised version of an old word for the terns; "stearn" appears in the poem The Seafarer, written around 1000 AD. Hirundo is the Latin for "swallow", referring to the similar long, forked tails of the two birds.[5] This resemblance also leads to the informal name Sea Swallow.[6] The Scottish names pictar, tarrock and their many variants are believed to be onomatopoeic, derived from the distinctive call.[5] Due to the difficulty in distinguishing the two species, all the common names are shared with the Arctic Tern.[7]

Four subspecies are generally recognized, although S. h. minussensis is sometimes considered to be just an intergrade between S. h. hirundo and S. h. longipennis.[8][9]

Subspecies Breeding range Distinctive features
S. h. hirundo
(Linnaeus, 1758).		 Europe, North Africa, Asia east to western Siberia and Kazakhstan, North America.[10] The nominate subspecies, only limited variation within this form.
S. h. minussensis
(Sushkin, 1925).		 Lake Baikal east to northern Mongolia and southern Tibet.[11] Paler upper body and wings than longipennis, black-tipped crimson bill.[11]
S. h. longipennis
(Nordmann, 1835)		 Central Siberia to China, Alaska.[10] Darker gray than nominate, with shorter, black bill, darker red-brown legs and longer wings.[10]
S. h. tibetana
(Saunders, 1876)		 Himalayas to southern Mongolia and China.[10] Like nominate, but bill shorter with broader black tip.[10]

Differences between American and Eurasian populations of the nominate subspecies are small. American birds have a slightly shorter wing length on average, and the extent of the black tip on the upper mandible tends to be less than in birds from Scandinavia and further east in Eurasia. The proportion of black in the bill is least in the west of Europe, so British breeders are very similar to American birds in this respect.[10]

[edit] Description

This medium-sized tern is 32–39 cm long (including a 6–9 cm fork in the tail) with a 72–83 cm wingspan. It weighs 97–146  g.

Breeding adults have light gray upperparts, white to very light gray underparts, a black cap, orange-red legs, and a thin, sharp, orange-red bill. The Common Tern's upperwings show a dark primary wedge, unlike the Arctic Tern in which the upperwing surface is uniformly grey. Its long tail extends no further than the folded wingtips on the standing bird, unlike Arctic and Roseate Terns in which the tail protrudes past the wingtips. The Common Tern is not as pale as the Roseate Tern and has longer wings. It is also smaller than the South American Tern.

In winter, the forehead and underparts are white, the bill is all black or black with a red base, and the legs are dark red or black.

Juvenile Common Terns show extensive ginger coloration and lack the 'scaly' appearance of juvenile Roseate Terns.

The Common tern is an agile flyer, capable of rapid turns and swoops, hovering, and vertical take-off. Flight speed during migration is 12–15 m/s (43–54 km/h, or 27–33 mph).[12] When commuting with fish it flies close to the surface in a strong head wind, but 10–30 m above the surface otherwise.

There are a number of similar species. In the breeding areas, Roseate Tern can be distinguished by its pale plumage, long, mainly black bill and very long tail feathers.[13] In flight, Roseate's heavier head and neck, long bill and faster, stiffer wingbeats are characteristic.[14] In the wintering regions, Antarctic Tern is bulkier, with a thick bill and white rump, White-cheeked Tern has uniform gray upperparts,[15] and South American Tern is larger, red-billed and has a smoother, more extensive black cap.[16] The most difficult species to separate is Arctic Tern, and until the key characteristics were clarified, distant or flying birds of the two species were often jointly recorded as "commie terns". Although similar in size, the two terns differ in structure and flight. Common Tern has a larger head, thicker neck, and more triangular and stiffer wings than its relative, and has a more powerful, direct flight. Arctic has grayer underparts than Common, which make its white cheeks more obvious, whereas the rump of Common can be grayish in winter, compared to the white of its relative. Common Tern develops a dark wedge on the wings as the breeding season progresses, but the wings of Arctic stay white throughout the summer. All the flight feathers of Arctic are translucent in sunshine, only the four innermost of common share this property. The bill of adult Common is orange-red with a black tip, and its legs are bright red, while both features are a darker red in Arctic, which also lacks the black bill tip.[13][17]

Hybrids between Common and Roseate Terns have been recorded, particularly from the US, and the intermediate plumage and calls shown by these birds is a potential pitfall. Such birds may have more extensive black on the bill, but confirmation of mixed breeding may depend on the exact details of individual flight feathers.[10]

[edit] Voice
Common Tern
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The Common Tern has a wide repertoire of calls, which have a lower pitch than the equivalent calls of Arctic Terns. The most distinctive is the alarm kee-yah, stressed on the first syllable, in contrast to the second-syllable stress of the Arctic Tern. The alarm call doubles up as a warning to intruders. Serious threats evoke a kyar, given as a tern takes flight, and quietens the usually noisy colony while its residents assess the danger. A down-slurred keeur is given when an adult is approaching while carrying a fish, and is possibly used for individual recognition (chicks emerge from hiding when they hear their parents giving this call). A common call is a kip uttered during social contact. Other vocalizations include kakakakaka when attacking intruders, and a staccato kek-kek-kek from fighting males.[18]

[edit] Distribution and habitat

Winter plumage adults in Brazil

All populations of the Common Tern are strongly migratory, wintering south of their breeding ranges in the temperate and subarctic Northern Hemisphere. First summer birds usually remain in the wintering quarters, although a few return to breeding colonies after the arrival of the adults.[19] In North America, the Common Tern breeds along the Atlantic coast from Labrador to North Carolina, and inland throughout much of Canada east of the Rocky Mountains. In the United States, some breeding populations can also be found in the states bordering the Great Lakes, and locally on the Gulf coast.[20] There are small colonies in Venezuela and on a few Caribbean islands.[21] New World birds winter along both coasts of Central and South America, to Argentina on the east coast and to Northern Chile on the west coast.[19][20]

This tern breeds across most of Europe, with the highest numbers in the north and east of the continent. There are small populations on the north African coast, and in the Azores, Canary Islands and Madeira. Most winter off western or southern Africa, with birds from the south and west of Europe tending to stay north of the equator, with other European birds moving further south.[22] The breeding range continues across the temperate and taiga of Asia, with scattered outposts on the Persian Gulf and the coast of Iran.[21] Western Asian birds winter in the northern Indian Ocean, those further north and east, such as S. h. longipennis, move through Japan, Thailand and the western Pacific as far as southern Australia.[19] There are small and erratic colonies in West Africa, in Nigeria and Guinea-Bissau, unusual in that they are within what is mainly a wintering area.[21] Only a few Common Terns have been recorded in New Zealand,[23] and this species' status in Polynesia is unclear.[24]

As long distance migrants, Common Terns sometimes occur well outside their normal range. Stray birds have been found inland in Africa (Zambia and Malawi), and on the Maldives and Comoros island,[25] the nominate subspecies has reached Australia,[26] and Asian S. h. longipennis has recent records from western Europe.[27]

The Common Tern breeds over a wider range of habitats than any of its relatives, nesting from the taiga of Asia to tropical shores,[28] and at altitudes up to 2000 m (6500 ft) in Armenia, and 4,800 m (15,500 ft) in Asia.[22] It avoids areas which are frequently exposed to excessive rain or wind, and also icy waters, so it does not breed as far north as the Arctic Tern, and . This species breeds close to freshwater or the sea on almost any open flat habitat, including sand or shingle beaches, firm dune areas, salt marsh, or, most commonly, islands. Flat grassland or heath, or even large flat rocks may be suitable in a island environment.[28] In mixed colonies, Common Terns will tolerate somewhat longer ground vegetation than Arctic Terns, but avoid the even taller growth acceptable to Roseate Terns; the relevant factor is the different leg lengths of the three species.[29] Common Terns adapt readily to artificial floating rafts, and may even nest on flat factory roofs. Outside the breeding season, all that is needed is access to fishing areas, and somewhere to land. Boats, buoys and piers are often used, as well as natural beaches and rocks.[28]

[edit] Behaviour

On land it can walk or run. It perches on rocks, posts, rails, boats, or buoys.

Head-scratching with the foot is direct (not over the wing) and can be performed during flight. Anting and sunbathing have not been reported. Water bathing is done in shallow waters close to shore. Sleep is done with the bill tucked into the scapular (shoulder) feathers.

[edit] Breeding

Nest site, Elliston, Newfoundland and Labrador

The Common Tern breeds in colonies which do not normally exceed 2000 pairs,[22] but may occasionally number more than 20,000 pairs[30] Colonies inland tend to be smaller than on the coast. Common Terns often nest alongside other coastal species, such as Arctic,[31] Roseate and Sandwich Terns, Black-headed Gulls,[32][33] and Black Skimmers.[34]

On their return to the breeding sites, the terns may loiter for a few days before settling into a territory. Terns defend only a small area, with distances between nests sometimes being as little as 50 cm (20 in), although 150–350 cm (60–130 in) is more typical. As with many birds, the exact site is used year after year, with one pair returning for 17 successive breeding seasons. Around 90 percent of experienced birds reuse their former territory, so young birds must nest on the periphery, find a bereaved mate, or move to another colony.[35]

Males select a nesting territory a few days after their arrival in the spring. There a male is joined by his previous partner, unless she is more than five days late, in which case the pair may separate.[36] The defence of the territory is mainly by the male, who repels intruders of either sex. He gives an alarm call, opens his wings, raises his tail and bows his head to show the black cap. If the intruder persists, the male stops calling and fights by bill grappling until the intruder submits by raising its head to expose the throat. Aerial trespassers are simply attacked, sometimes following a joint upward spiralling flight.[35] Despite the aggression shown to adults, wandering chicks are usually tolerated, whereas in a gull colony they would be attacked and killed.[37]

Hovering and calling to deter intruders Great Gull Island

Aerial courtship displays then occur, in which a male and a female climb in wide circles up to 200  (600 ft) or more, calling all the while, before the birds descend together in zigzag glides. The male may carry a fish, in which case he may attract the attention of other males too.[38] On the ground, males offer fishes to females. Once the pair is established or confirmed, both male and female scratch depressions in the ground, one of which will eventually become the nest. After the eggs are laid, some lining material is added throughout the incubation period, such as grass, reeds, or even rubbish.

Clutch size is normally three eggs, fewer in bad food years. Eggs are normally 42 × 30.5 mm, 20 ml, and 21 g. They are cream, buff, or brown, and finely marked with streaks or spots of black, brown or grey. Incubation is by both sexes and lasts 22–33 days, depending on disturbances at the colony which may leave the eggs unattended. On very hot days the incubating parent may fly to water to wet its belly feathers before returning to the eggs, thus affording the eggs some cooling. At hatching the young are precocial, eyes open, covered in thick down, and capable of standing and taking food within 1–3 h. Young are brooded and fed fishes by both sexes. Chicks remain on nesting territory, forsaking the nest but seeking refuge in vegetation. They fledge after 22–29 days.

Like many terns, this species is very defensive of its nest and young, and will harass humans, dogs, muskrats and most diurnal birds, but unlike the more aggressive Arctic Tern, it rarely hits the intruder, usually swerving off at the last moment. Adults can discriminate between individual humans, attacking familiar people more intensely than strangers.[39] Nocturnal predators do not elicit similar attacks; colonies can be wiped out by rats, and adults desert the colony for up to eight hours when Great Horned Owls are present.[40] Common Terns usually breed once a year. Second clutches are possible if the first one is lost. Rarely, a second clutch may be laid and incubated while some chicks from the first clutch are still being fed.[41] The first breeding attempt is usually at four years of age, sometimes at three years. The maximum documented lifespan in the wild is 23 years.[42][43]

[edit] Food and feeding

The head and bill point down during a search for fish

Like all Sterna terns, the Common Tern feeds by plunge-diving for fish, from a height of 1–6 m (3–18 ft), either in the sea or in freshwater lakes and large rivers. It may submerge for a second or so, but to no more than 50 cm (18 in) below the surface.[44] When seeking fish, it flies with its head down and its bill held vertically.[29] It may circle or hover before diving, and then plunges directly into the water, whereas the Arctic Tern favours a "stepped-hover" technique,[45] and the Roseate Tern dives at speed from a greater height, and submerges for longer.[46] The Common Tern typically forages up to 5–10 km (3–6 mi) away from the breeding colony, sometimes as far as 15 km (9 mi).[1] It will follow schools of fish, and its west African migration route is affected by the location of huge shoals of sardines off the coast of Ghana;[44] It will also track groups of predatory fish, waiting for their prey to rise to the sea surface.[1] Terns often feed in flocks, especially if food is plentiful, and the fishing success rate in a flock is typically about one-third higher than for individuals.[44]

Terns have red oil droplets in the cone cells of the retinas of their eyes. This improves contrast and sharpens distance vision, especially in hazy conditions.[47] Birds that have to see through an air/water interface, such as terns and gulls, have more strongly coloured carotenoid pigments in the cone oil drops than other avian species.[48] The improved eyesight helps terns to locate shoals of fish, although it is uncertain whether they are sighting the phytoplankton on which the fish feed, or observing other terns diving for food.[49] Tern's eyes are not particularly ultraviolet sensitive, an adaptation more suited to terrestrial feeders like the gulls.[50]

This tern preferentially hunts fish between 5–15 cm (2–6 in) long.[44][51] The species caught depend on what is available, but if there is a choice, terns feeding several chicks will take larger prey than those with smaller broods.[52] The proportion of fish fed to chicks may be as high as 95 percent in some areas, but invertebrate prey may form a significant part of the diet elsewhere. This may include worms, leeches, mollusks such as small squid, and crustaceans (prawns, shrimp and mole crabs). In freshwater areas, large insects may may caught, such as beetles, cockchafers and moths. Adults may be caught in the air, and larvae picked from the ground or from the water surface. Prey is caught in the bill and either swallowed head-first, or carried back to the chicks. Occasionally, two or more small fish may be carried simultaneously.[44]

The Common Tern may attempt to steal fish from Arctic Terns,[53] but may itself be harassed by kleptoparasitic jaegers,[54] Laughing Gulls,[55] Roseate Terns,[56] or by other Common Terns while bringing fish back to its nest.[53] In one study, two males whose mates had died spent much time stealing food from neighboring broods.[57]

[edit] Predators and parasites

Rats will take tern eggs, and may even store large numbers in caches,[43] and the American mink is a serious predator of hatched chicks, both in North America, and in Scotland where it has been introduced.[58] The red fox can also be a local problem.[59] Because Common Terns nest on islands, the most common predators are normally other birds rather than mammals. The Ruddy Turnstone will take eggs from unattended nests,[60][61] and gulls may take chicks.[62][63] Great Horned Owls and Short-eared Owls will kill both adults and chicks, and Black-crowned Night Herons will also eat small chicks.[64][65] Merlins and Peregrine Falcons may attack flying terns; as with other birds, it seems likely that one advantage of flocking behaviour is to confuse fast-flying predators.[37]

The Common Tern hosts feather lice, which are quite different from those found in the Arctic Terns, despite the close relationship of the two birds.[66] It may also be infected by parasitic worms, such as the widespread Diphyllobothrium species, the duck parasite Ligula intestinalis, and Schistocephalus species carried initially by fish. Tapeworms of the family Cyclophyllidea are also a possibility. The mite Reighardia sternae has been found in Common Terns from Italy, North America and China.[67] A study 75 breeding Common Terns found that none carried blood parasites.[68] It is possible that the Common Tern may be threatened in the future by outbreaks of avian influenza to which it is susceptible.[1]

[edit] Status

Common Tern96.ogg
Asian subspecies S. h. longipennis wintering in Mooloolabah, Australia

The Common Tern has a large population of 1,600,000–4,600,000 mature individuals and a huge breeding range estimated at 29,200,000 km2 (113 mi2); it is therefore classed as Least Concern on the IUCN Red List.[1] Breeding numbers have been estimated at 250,000–500,000 pairs.The majority are in Asia, with up to 140,000 pairs in Europe.[69] North America has less than 80,000 pairs, with most on the northeast Atlantic coast;[70] the Great Lakes region holds a declining population of less than 10,000 pairs.[71]

In the nineteenth century, the use of tern feathers and wings in the millinery trade cause large decreases in Common Tern populations in both Europe and North America, especially on the Atlantic coasts and inland. Sometimes entire stuffed birds were used to make hats. Numbers largely recovered early in the following century due to legislation and the work of conservation organizations.[59][65] Although some populations are stable, numbers in North America have fallen by more than 70 percent in 40 years, and there is an overall negative trend in the estimates for this species.[1]

Threats may come from habitat loss through building, pollution or vegetation growth, or disturbance of breeding birds by humans, vehicles, boats or dogs. Local natural flooding may lead to nest losses, and some colonies are vulnerable to predation by rats and large gulls. Gulls may also compete for nest sites. Some birds are hunted in the Caribbean for commercial sale.[1] Breeding success may be enhanced by the use of floating nest rafts, man-made islands or other artificial nest sites, and by preventing human disturbance. Overgrown vegetation may be burned to clear the ground, and gulls can be killed or discouraged by deliberate disturbance.[1]

The Common Tern is one of the species to which the Agreement on the Conservation of African-Eurasian Migratory Waterbirds (AEWA) and the US-Canada Migratory Bird Treaty Act of 1918 apply.[72][73] Parties to the AWEA greement are required to engage in a wide range of conservation strategies described in a detailed action plan. The plan is intended to address key issues such as species and habitat conservation, management of human activities, research, education, and implementation.[74] The North American legislation, although there is a greater emphasis on protection.[75]

  • Three eggs in a nest on the ground

  • Adult incubating eggs in a nest

[edit] References

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  70. ^ Kress, Stephen W; Weinstein, Evelyn H; Nisbet, Ian C; T; Shugart, Gary W; Scharf, William C; Blokpoel, Hans; Smith, Gerald A; Karwowski, Kenneth; Maxwell, George R; Chapdelaine, Gilles; Montevecchi, William A; Lock, Anthony R; Smith, Carol F; Miller, Eileen; Spendelow, Jeffrey A; Gochfeld, Michael; Burger, Joanna; Erwin, R Michael (1983). "The Status of Tern Populations in Northeastern United States and Adjacent Canada". Colonial Waterbirds 6: 84–106. 
  71. ^ Cuthbert (2003) p. 1.
  72. ^ "Annex 2: Waterbird species to which the Agreement applies". Agreement on the conservation of African-Eurasian migratory Waterbirds (AEWA). UNEP/ AEWA Secretariat. http://www.unep-aewa.org/documents/agreement_text/eng/pdf/aewa_agreement_text_annex2.pdf. Retrieved 25 January 2012. 
  73. ^ "List of Migratory Birds". Birds protected by the Migratory Bird Treaty Act. US Fish and Wildlife Service. http://www.fws.gov/migratorybirds/RegulationsPolicies/mbta/mbtandx.html.  Retrieved 25 January 2012.
  74. ^ "Introduction". African-Eurasian Waterbird Agreement. UNEP/ AEWA Secretariat. http://www.unep-aewa.org/about/introduction.htm. Retrieved Retrieved 25 January 2012.. 
  75. ^ "Migratory Bird Treaty Act of 1918". Digest of Federal Resource Laws of Interest to the U.S. Fish and Wildlife Service. U.S. Fish and Wildlife Service. http://www.fws.gov/laws/lawsdigest/migtrea.html.  Retrieved 25 January 2012.

[edit] Cited texts

  • Beaman, Mark; Madge, Steve; Burn, Hilary; Zetterstrom, Dan (1998). The Handbook of Bird Identification: For Europe and the Western Palearctic. London: Christopher Helm. ISBN 0713639601. 
  • Best, E P H; Haeck, J (1984). Ecological Indicators for the Assessment of the Quality of Air, Water, Soil and Ecosystems: Symposium Papers ("Environmental Monitoring & Assessment"). D Reidel. ISBN 9027717087. 
  • Blomdahl, Anders; Breife, Bertil; Holmstrom, Niklas (2007). Flight Identification of European Seabirds. London: Christopher Helm. ISBN 0713686162. 
  • Brazil, Mark (2008). Birds of East Asia. London: Christopher Helm. ISBN 071367040. 
  • Burger, J., and M. Gochfeld, 1991, The Common Tern: Its breeding biology and social behavior, New York: Columbia University Press.
  • Cocker, Mark; Mabey, Richard (2005). Birds Britannica. London: Chatto & Windus. ISBN 0701169079. 
  • Cuthbert, Francesca J; Wires, Linda R; Timmerman, Kristina (2003). Status assessment and conservation recommendations for the Common Tern (Sterna hirundo) in the Great Lakes region. . U.S. Department of the Interior, Fish and Wildlife Service, Fort Snelling, Minnesota. http://www.fws.gov/midwest/eco_serv/soc/birds/pdf/cote-sa03.pdf. 
  • van Duivendijk, Nils (2011). Advanced Bird ID Handbook: The Western Palearctic:. London: New Holland. ISBN 1780090226. 
  • Enticott, Jim; Tipling, David (2002). Seabirds of the World. London: New Holland Publishers. ISBN 1843303272. 
  • Harrison, Peter (1988). Seabirds. London: Christopher Helm. ISBN 0747014108. 
  • del Hoyo, J., A. Elliott, and J. Sargatal, 1996, Handbook of the Birds of the World, Volume 3: Hoatzin to Auks, Barcelona: Lynx Edicions.
  • Fisher, James; Lockley, R M (1989). Sea‑Birds (Collins New Naturalist series). London: Bloomsbury Books. ISBN 1870630882. 
  • Hume, Rob (1993). The Common Tern. London: Hamlyn. ISBN 0540012661. 
  • Linnaeus, C (1758) (in Latin). Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata.. Holmiae:Laurentii Salvii. 
  • Lythgoe, J N (1979). The Ecology of Vision. Oxford: Clarendon Press. ISBN 0198545290. 
  • National Geographic Society, 2002, Field Guide to the Birds of North America, Washington, DC: National Geographic. ISBN 0792268776
  • Olsen, Klaus Malling; Larsson, Hans (1995). Terns of Europe and North America. London: Christopher Helm. ISBN 0713640561. 
  • Robertson, Hugh; Heather, Barrie (2005). The Field Guide to the Birds of New Zealand. Auckland: Penguin Group (NZ). ISBN 0143020404. 
  • Rothschild, Miriam; Clay, Theresa (1953). Fleas, Flukes and Cuckoos. A study of bird parasites. London: Collins. 
  • Sandilands, Allan P (2005). Birds of Ontario: habitat requirements, limiting factors, and status Nonpasserines, Waterfowl Through Cranes: 1. Vancouver: University of British Columbia Press. ISBN 0774810661. 
  • Schulenberg, Thomas S; Stotz, Douglas F; Lane, Daniel F; O'Neill, John P; Parker, Theodore A (2010). Birds of Peru. Princeton, NJ: Princeton University Press. ISBN 069113023X. 
  • Simpson, Ken; Day, Nicolas (2010). Field Guide to the Birds of Australia - 8th Edition. Camberwell, Victoria: Penguin Books. ISBN 0670072311. 
  • Sinclair, Ian; Hockey, Phil; Tarboton, Warwick (2002). SASOL Birds of Southern Africa. Cape Town: Struik. ISBN 1-86872-721-1. 
  • Sinclair, Sandra (1985). How Animals See: Other Visions of Our World. Beckenham, Kent: Croom Helm. ISBN 0709933363. 
  • Snow, David; Perrins, Christopher M (editors) (1998). The Birds of the Western Palearctic (BWP) concise edition (2 volumes). Oxford: Oxford University Press. ISBN 019854099X. 
  • Stephens, David W; Brown, Joel Steven; Ydenberg, Ronald C (2007). Foraging: behavior and ecology. Chicago: University of Chicago Press. ISBN 0226772640. 
  • Watling, Dick (2003). A Guide to the Birds of Fiji and Western Polynesia. Suva, Fiji: Environmental Consultants. ISBN 982-9030-04-0. 
  • Zeigler, Harris Philip; Bischof, Hans-Joachim (1993). Vision, Brain, and Behavior in Birds: a comparative review. Cambridge, Massachusetts: MIT Press. ISBN 026224036X. 

[edit] External links

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