Pelicans are large water birds in the family Pelecanidae. They are characterised by a long beak and large throat pouch, used in catching, and draining water from, their prey. The eight living pelican species have a sometimes patchy global distribution, ranging latitudinally from the tropics to the temperate zone, though they are absent from much of interior and southern South America as well as from polar regions and the open ocean. Fossil evidence of pelicans dates back at least 30 million years, from the remains of a beak very similar to modern species recovered from Oligocene strata in France.

Pelicans frequent inland and coastal waters where they feed principally on fish. Gregarious birds, they breed colonially and often hunt cooperatively. They have a long history of cultural significance in mythology, and in Christian and heraldic iconography.

Taxonomy and systematics

The closest living relatives of pelicans are the Hammerkop...

...and the Shoebill

The name is derived from the Ancient Greek word πελεκυς pelekys meaning “axe” and applied to birds that supposedly cut wood with their bills or beaks. Pelicans give their name to the order Pelecaniformes, an order which has had a varied taxonomic history; darters, cormorants, gannets, boobies and frigatebirds are traditional members which have since been reclassified as Suliformes, while tropicbirds now have their own order, the Phaethontiformes. Evidence points to the clade consisting of the Shoebill and the Hamerkop being the sister group of the pelicans.^[1]

Cladogram based on Hackett et al. (2008).^[2]

Fossil record

The fossil record shows that the pelican lineage has been in existence for at least 30 million years; the oldest known fossil was found in Early Oligocene deposits in France and is remarkably similar to modern forms.^[3] Its beak is almost complete and is morphologically identical to that of present day pelicans showing that this advanced feeding apparatus was already in existence 30 million years ago.^[3] An Early Miocene fossil has been named Miopelecanus gracilis on the basis of certain features originally considered unique but later thought to lie within the range of inter-specific variation in Pelecanus.^[3] The Late Eocene Protopelicanus may be a pelecaniform or suliform – or a similar aquatic bird such as a pseudotooth (Pelagornithidae).^[4] The supposed Miocene pelican Liptornis from Argentina is a nomen dubium, being based on hitherto indeterminable fragments.^[5]

Fossil finds from North America have been meagre, compared with Europe, which has a richer fossil record.^[6] Several Pelecanus species have been described from fossil material, including:^[7]

• Pelecanus cadimurka Rich & van Tets, 1981 (Late Pliocene, South Australia)^[8]
• Pelecanus cautleyi Davies, 1880 (Early Pliocene, Siwalik Hills, India)^[7]
• Pelecanus fraasi Lydekker, 1891 (Middle Miocene, Bavaria, Germany)^[7]
• Pelecanus gracilis Milne-Edwards, 1863 (Early Miocene, France) (see: Miopelecanus)^[7]
• Pelecanus halieus Wetmore, 1933 (Late Pliocene, Idaho, USA)^[9]
• Pelecanus intermedius Fraas, 1870 (Middle Miocene, Bavaria, Germany)^[7] (transferred to Miopelecanus by Cheneval in 1984)
• Pelecanus odessanus Widhalm (1886) (Late Miocene, near Odessa, Ukraine)^[10]
• Pelecanus schreiberi Olson, 1999 (Early Pliocene, North Carolina, USA)^[11]
• Pelecanus sivalensis Davies, 1880 (Early Pliocene, Siwalik Hills, India)^[7]
• Pelecanus tirarensis Miller, 1966 (Late Oligocene to Middle Miocene, South Australia)^[12]

Living species

The eight living pelican species can be divided into two groups, four with mostly white adult plumage, which nest on the ground (Australian, Dalmatian, Great White, and American White Pelicans), and four with grey or brown plumage which nest in trees (Pink-backed, Spot-billed and Brown Pelicans), or on sea rocks (Peruvian Pelican). The largely marine Brown and Peruvian Pelicans, formerly considered conspecific,^[13] are sometimes separated from the others by placement in the subgenus Leptopelicanus.^[14] The Dalmatian Pelican has sometimes been considered a subspecies of the Spot-billed, though it differs in both nesting habits and morphology and is now accepted as a full species.^[15]

Living species of Pelecanus
Common and binomial names[16]	Image	Description	Range and status
Brown Pelican Pelecanus occidentalis Linnaeus, 1766		Length 106–137 cm (42–54 in), wingspan 1.83-2.5 m (6–8 ft), weight 2.75-5.5 kg (6-12 lb). Smallest pelican; distinguished by brown plumage; feeds by plunge-diving.	Five subspecies. Coastal distribution ranging from North America and Caribbean to northern South America and Galapagos. Status: Least Concern.
Peruvian Pelican Pelecanus thagus Molina, 1782		Length 150 cm (60 in), weight 7 kg (15.5 lb). Dark with a white stripe from the crown down the sides of the neck.	Monotypic. Coastal Peru and Chile. Status: Near Threatened.
Spot-billed Pelican Pelecanus philippensis Gmelin, 1789		Length 125–152 cm (49–60 in), weight 4.1–6 kg (9-13.2 lb). Mainly white, with a grey hindneck crest and brownish tail.	Monotypic. Southern Asia from southern Pakistan across India east to Indonesia; extinct in the Philippines. Status: Near Threatened.
Pink-backed Pelican Pelecanus rufescens Gmelin, 1789		Length 125–155 cm (49–61 in), wingspan 2.15–2.9 m (7-9.5 ft), weight 4–7 kg (9-15.4 lb). Grey and white plumage, occasionally pinkish on the back, with a yellow upper mandible and grey pouch.	Monotypic. Africa and southern Arabia; extinct in Madagascar. Status: Least Concern.
American White Pelican Pelecanus erythrorhynchos Gmelin, 1789		Length 130–170 cm (51–67 in), wingspan 2.4–3 m (95–120 in), usual weight range 5–9.1 kg (11-20 lb). Plumage almost entirely bright white, except for black primary and secondary remiges only visible in flight.	Monotypic. Inland North America. Status: Least Concern.
Great White Pelican Pelecanus onocrotalus Linnaeus, 1758		Length 160 cm (63 in), wingspan 2.8 m (110 in), weight 10 kg (22 lb). Plumage pure white, with pink facial patch and legs.	Monotypic. Patchy distribution from eastern Mediterranean east to Vietnam and south to South Africa. Status: Least Concern.
Dalmatian Pelican Pelecanus crispus Bruch, 1832		Length 160–180 cm (63–71 in), wingspan 3 m (118 in), weight 11–15 kg (24-33 lb). Largest pelican; differs from Great White Pelican in having curly nape feathers, grey legs and greyish-white plumage.	Monotypic. South-eastern Europe to India and China. Status: Vulnerable.
Australian Pelican Pelecanus conspicillatus Temminck, 1824		Length 160–180 cm (63–71 in), wingspan 2.3–2.6 m (90–100 in), usual weight range 4.54-7.7 kg (10-17 lb). Predominantly white with black along primaries and very large, pale pinkish bill.	Monotypic. Australia and New Guinea; vagrant to New Zealand, Solomons, Bismarck Archipelago, Fiji and Wallacea. Status: Least Concern.

Description

An Australian Pelican gliding with its large wings extended

Pelicans are very large birds with very long, terminally hooked, bills characterised by the attachment of a huge gular pouch. The slender rami of the lower bill and the flexible tongue muscles form the pouch into a basket for catching fish and, sometimes, rainwater.^[14] They have a long neck and short stout legs with large, fully webbed, feet. Although they appear bulky they are relatively light because of air pockets in the skeleton and beneath the skin so that they float high in the water.^[14] The tail is short and square, with 20 to 24 retrices. The wings are long and broad, suitably shaped for soaring and gliding flight, and have the unusually large number of 30 to 35 secondary flight feathers. They are among the heaviest flying birds.^[17]

Males are generally larger than females and have longer bills.^[14] The smallest species is the Brown Pelican, small individuals of which can be as little as 2.75 kg (6 lb), 106 cm (42 in) long, and can have a wingspan of as little as 1.83 m (6 ft). The largest is believed to be the Dalmatian, at up to 15 kg (33 lb), 183 cm (72 in) long, with a maximum wingspan of 3 m (nearly 10 ft). The Australian Pelican has the longest bill of any bird.^[13]

Distribution and habitat

Modern pelicans are found on all continents except Antarctica. They primarily inhabit warm regions, though breeding ranges extend to latitudes of 45° South (Australian Pelicans in Tasmania) and 60° North (American White Pelicans in western Canada).^[13] Birds of inland and coastal waters, they are absent from polar regions, the deep ocean, oceanic islands, and inland South America, as well as from the eastern coast of South America from the mouth of the Amazon River southwards.^[14]

Behaviour and ecology

Pelicans swim well with their strong legs and their webbed feet. They rub the backs of their heads on their preen glands to pick up an oily secretion, which they transfer to their plumage to waterproof it.^[13] They dissipate excess heat by gular flutter - rippling the skin of throat and pouch with the bill open to promote evaporative cooling.^[14] They roost and loaf communally on beaches, sandbanks and in shallow water.^[14] A fibrous layer deep in the breast muscles can hold the wings rigidly horizontal for gliding and soaring. Thus they use thermals for soaring, combined both with gliding and with flapping flight in V-formation, to commute distances of up to 150 km (93 mi) to feeding areas.^[13]

Feeding

Brown Pelicans diving into the sea to catch fish in Jamaica

The diet of pelicans usually consists of fish, but they also eat amphibians, turtles, crustaceans and, at least occasionally, birds.^[18][19] In deep water, white pelicans often fish alone. Nearer the shore, several will encircle schools of small fish or form a line to drive them into the shallows, beating their wings on the water surface and then scooping up the prey.^[20] They catch multiple small fish by expanding the throat pouch, which must be drained above the water surface before swallowing. This operation takes up to a minute, during which time other seabirds may steal the fish. Large fish are caught with the bill-tip, then tossed up in the air to be caught and slid into the gullet head-first. A gull will sometimes stand on the pelican's head, peck it by way of distraction, and grab a fish from the open bill.^[21] Pelicans in their turn sometimes snatch prey from other seabirds.^[13]

The Brown Pelican usually plunge-dives for its prey which is often a type of herring known as menhaden.^[20] Rarely, other species such as the Peruvian Pelican and the Australian Pelican also make use of this fishing method.

The Australian Pelican, although principally a fish eater, is also an eclectic and opportunistic carnivore and scavenger which takes "anything from insects and small crustaceans to ducks and small dogs" ^[22] as well as carrion.^[22]

Consumption of other birds by pelicans is rare. It has been suggested that feeding on other birds is more likely with captive pelicans that live in a semi-urban environment and are in constant close contact with humans,^[19] although this behaviour has also been observed in the wild. On Malgas Island in South Africa, the biologist Marta de Ponte was the first to record Great White Pelicans eating Cape Gannet chicks.^[23] The same species of pelican has been observed swallowing Cape Cormorants, Kelp Gulls, Greater Crested Terns and African Penguins.

Breeding

Spot-billed Pelican feeding juvenile at nest, Garapadu, India

Pelicans are gregarious and nest colonially. Pairs are monogamous for a single season, but the pair bond extends only to the nesting area; mates are independent away from the nest. The ground-nesting (white) species have a complex communal courtship involving a group of males chasing a single female in the air, on land, or in the water while pointing, gaping, and thrusting their bills at each other. They can finish the process in a day. The tree-nesting species have a simpler process in which perched males advertise for females.^[13]

In all species copulation takes place at the nest site; it begins shortly after pairing and continues for 3–10 days before egg-laying. The male brings the nesting material, ground-nesters (which may not build a nest) sometimes in the pouch and tree-nesters crosswise in the bill. The female then heaps the material up to form a simple structure.^[13]

The eggs are oval, white and coarsely textured.^[14] All species normally lay at least two eggs; the usual clutch size is 1-3, rarely up to 6.^[14] Both sexes incubate with the eggs on top of or below the feet; they may display when changing shifts. Incubation takes 30–36 days;^[14] hatching success for undisturbed pairs can be as high as 95 percent but, because of sibling competition or siblicide, in the wild usually all but one nestling dies within the first few weeks (later in the Pink-backed and Spot-billed species). The newly hatched altricial chicks are pink and naked; their skin darkens to grey or black within 4-14 days before developing a covering of white or grey down. Both parents feed their young. Small chicks are fed by regurgitation; after about a week they are able to put their heads into their parent’s pouch and feed themselves.^[24] Sometimes before, or especially after, being fed, they may seem to have a seizure that ends in falling unconscious; the reason is not clearly known.^[13]

Parents of ground-nesting species sometimes drag older young around roughly by the head before feeding them. From about 25 days old,^[14] the young of these species gather in "pods" or "crèches" of up to 100 birds in which parents recognise and feed only their own offspring. By 6–8 weeks they wander around, occasionally swimming, and may practice communal feeding.^[13] Young of all species fledge 10–12 weeks after hatching. They may remain with their parents afterwards, but are now seldom or never fed. They are mature at three or four years old.^[14] Overall breeding success is highly variable.^[13]

Status, threats and conservation

Oiled Brown Pelican being washed at Fort Jackson, Louisiana, in May 2010 following the Deepwater Horizon oil spill

The Dalmatian Pelican is the rarest species with a population estimated at between 10,000 and 20,000. It is listed as "Vulnerable" by the IUCN Red List of Threatened Species as the population trend is downwards, especially in Mongolia where it is nearly extinct. Several colonies however are increasing in size and the colony at the Small Prespa Lake in Greece has nearly 1000 breeding pairs.^[25] The Spot-billed Pelican has an estimated population between 13,000 and 18,000 and is considered to be "Near Threatened" in the IUCN Red List of Threatened Species. The chief threats it faces are from habitat loss and human disturbance but populations have mostly stabilised following increased protection in India and Cambodia.^[26]

The most abundant species is believed to be the Australian Pelican, with a population generally estimated at around 400,000 individuals. However, estimates for the species have varied wildly between 100,000 and 1,000,000 over the years, and it is possible that the White Pelican, the population of which is more consistently estimated at 270,000 and 290,000 individuals, is more abundant. The Brown Pelican may be even more numerous with estimates of 650,000 birds throughout its range. It has been removed from the endangered species list.^[27]

Mythology and popular culture

Pelicans on a Fifth Dynasty relief at the Abu Gorab temple, Egypt

Moche pelican container from the Larco Museum Collection Lima, Peru

Picture of a vulning pelican from a church in Althofen, Austria

Pelican hood ornament on a 1931 Buick

Ancient Egypt

The pelican (Henet in Egyptian) was associated in Ancient Egypt with death and the afterlife. It was depicted in art on the walls of tombs, and figured in funerary texts, as a protective symbol against snakes. Henet was also referred to in the Pyramid Texts as the 'mother of the king' and thus seen as a goddess. References in non-royal funerary papyri show that the pelican was believed to possess the ability to prophesy safe passage in the underworld for someone who had died.^[28]

Australia

An origin myth from the Murri people of Queensland, cited by Andrew Lang, describes how the Australian Pelican acquired its black and white plumage. The pelican, formerly a black bird, made a canoe during a flood in order to save drowning people. He fell in love with a woman he thus saved, but she and her friends tricked him and escaped. The pelican consequently prepared to go to war against them by daubing himself with white clay as war paint. However, before he had finished, another pelican, on seeing such a strange piebald creature, killed him with its beak, since when all such pelicans have been black and white.^[29]

Pre-Columbian America

The Moche people of ancient Peru worshipped nature.^[30] They placed emphasis on animals and often depicted pelicans in their art.^[31]

Christianity

In medieval Europe, the pelican was thought to be particularly attentive to her young, to the point of providing her own blood by wounding her own breast when no other food was available. As a result, the pelican became a symbol of the Passion of Jesus and of the Eucharist. A reference to this mythical characteristic is contained for example in the hymn by Saint Thomas Aquinas, "Adoro te devote" or "Humbly We Adore Thee", where in the penultimate verse he describes Christ as the "loving divine pelican, able to provide nourishment from his breast".^[32]

The self-sacrificial aspect of the pelican was reinforced by the widely-read mediaeval bestiaries. The device of "a pelican in her piety" or "a pelican vulning (from Latin vulno to wound) herself" was used in heraldry. An older version of the myth is that the pelican used to kill its young then resurrect them with its blood, again analogous to the sacrifice of Jesus. Likewise a folktale from India says that a pelican killed her young by rough treatment but was then so contrite that she resurrected them with her own blood.^[13] The symbol of the Irish Blood Transfusion Service is a pelican, and for most of its existence the headquarters of the service was located at Pelican House in Dublin, Ireland.^[33]

These legends regarding the self-wounding and the provision of blood may have arisen because of the impression a pelican sometimes gives that it is stabbing itself with its bill. In reality, it often presses this onto its chest in order to fully empty the pouch. Another possibility is the fact that the bird often rests with its bill on its breast. The Dalmatian Pelican has a blood-red pouch in the early breeding season and this fact may have contributed to the myth.^[13]

Heraldry

Pelicans have featured extensively in heraldry, generally using the Christian symbolism of the pelican as caring and self-sacrificing parent. Corpus Christi College, Cambridge features a pelican on its coat of arms ^[34] as does Corpus Christi College, Oxford.^[35] The medical faculties of Charles University in Prague also have a pelican as their emblem.^[36] These uses symbolise the bird as a caring mother, representing Christ feeding his followers with his body and blood ('Corpus Christi' means 'body of Christ').^[34]

Modern usage

The pelican is the national bird of Sint Maarten and features on its coat of arms.^[37] It is also used on the Louisiana state flag and Louisiana state seal, as the Brown pelican is the Louisiana state bird. The pelican is featured prominently on the seals of Loomis Chaffee, Louisiana State University, and Tulane University, and is also the mascot of Tulane. A pelican logo is used by the Portuguese bank Montepio Geral.[1] A pelican is depicted on the reverse of the Albanian 1 lek coin, issued in 1996.^[38]

The pelican is the subject of a popular limerick originally composed by Dixon Lanier Merritt in 1910 with several variations by other authors.^[39]

References