Pelicans (genusPelecanus) are agenus of largewater birds that make up the familyPelecanidae. They are characterized by a longbeak and a largethroat pouch used for catchingprey and draining water from the scooped-up contents before swallowing. They have predominantly pale plumage, except for thebrown andPeruvian pelicans. The bills, pouches, and bare facial skin of all pelicans become brightly coloured before the breeding season.
The eight living pelican species have a patchy, seasonally-dependent yet global distribution, ranginglatitudinally from thetropics to thetemperate zone. Pelicans are absent from interiorAmazonian South America, frompolar regions and the open ocean; at least one species is known to migrate to the inland desert ofAustralia'sRed Centre, after heavy rains create temporary lakes. White pelicans are also observed at the American state ofUtah'sGreat Salt Lake, for example, some 600 miles (965 km) from the nearest coastline (the Pacific West Coast). They have also been seen hundreds of miles inland in North America, having flown northwards along theMississippi River and other large waterways.
Long thought to be related tofrigatebirds,cormorants,tropicbirds, andgannets and boobies, pelicans instead are most closely related to theshoebill andhamerkopstorks (although these two birds are not actuallytrue 'storks'), and are placed in the orderPelecaniformes.Ibises,spoonbills,herons, andbitterns have been classified in the same order. Fossil evidence of pelicans dates back at least 36 million years to the remains of atibiotarsus recovered from lateEocene strata of Egypt that bears striking similarity to modern species of pelican.[1] They are thought to have evolved in theOld World and spread into the Americas; this is reflected in the relationships within the genus as the eight species divide into Old World andNew World lineages.[3] This hypothesis is supported by fossil evidence from the oldest pelican taxa.[1]
Pelicans will frequent inland waterways but are most known for residing along maritime and coastal zones, where they feed principally on fish in their large throat pouches, diving into the water and catching them at/near the water's surface. They can adapt to varying degrees ofwater salinity, from freshwater and brackish to—most commonly—seawater. They aregregarious birds, travelling in flocks,hunting cooperatively, and breedingcolonially. Four white-plumaged species tend to nest on the ground, and four brown or grey-plumaged species nest mainly in trees.[4] The relationship between pelicans and people has often been contentious. The birds have been persecuted because of their perceived competition with commercial and recreational fishing.[5] Their populations have fallen throughhabitat destruction, disturbance, and environmental pollution, and three species are of conservation concern. They also have a long history of cultural significance inmythology, and in Christian andheraldiciconography.
Taxonomy and systematics
Etymology
The name comes from theAncient Greek wordpelekan (πελεκάν),[6] which is itself derived from the wordpelekys (πέλεκυς) meaning "axe".[7] Inclassical times, the word was applied to both the pelican and the woodpecker.[8]
The genusPelecanus was first formally described byCarl Linnaeus in his landmark 175810th edition ofSystema Naturae. He described the distinguishing characteristics as a straight bill hooked at the tip, linear nostrils, a bare face, and fully webbed feet. This early definition included frigatebirds, cormorants, andsulids, as well as pelicans.[9]
The family Pelecanidae was introduced (as Pelicanea) by the FrenchpolymathConstantine Samuel Rafinesque in 1815.[10][11] Pelicans give their name to the Pelecaniformes, anorder which has a varied taxonomic history.Tropicbirds,darters, cormorants, gannets, boobies, and frigatebirds, all traditional members of the order, have since been reclassified: tropicbirds into their own order,Phaethontiformes, and the remainder into the Suliformes. In their place, herons, ibises, spoonbills, the hamerkop, and the shoebill have now been transferred into the Pelecaniformes.[12] Molecular evidence suggests that the shoebill and the hamerkop form asister group to the pelicans,[13] though some doubt exists as to the exact relationships among the three lineages.[14]
The eight living pelican species were traditionally divided into two groups, one containing four ground-nesters with mainly white adult plumage (Australian,Dalmatian,great white, andAmerican white pelicans), and one containing four grey- or brown-plumaged species which nest preferentially either in trees (pink-backed,spot-billed andbrown pelicans), or on sea rocks (Peruvian pelican). The largely marine brown and Peruvian pelicans, formerly consideredconspecific,[4] are sometimes separated from the others by placement in thesubgenusLeptopelecanus[15] but in fact species with both sorts of appearance and nesting behavior are found in either.
DNA sequencing of both mitochondrial and nuclear genes yielded quite different relationships; the threeNew World pelicans formed one lineage, with the American white pelican sister to the two brown pelicans, and the five Old World species the other. The Dalmatian, pink-backed, and spot-billed were all closely related to one another, while the Australian white pelican was their next-closest relative. The great white pelican also belonged to this lineage, but was the first to diverge from the common ancestor of the other four species. This finding suggests that pelicans evolved in the Old World and spread into the Americas, and that preference for tree- or ground-nesting is more related to size than genetics.[3]
Length 1.3–1.8 m (4.3–5.9 ft), wingspan 2.44–2.9 m (8.0–9.5 ft), weight 5–9 kg (10–20 lb).[17] Plumage almost entirely white, except for black primary and secondaryremiges only visible in flight.
Monotypic. Breeds in inland Canada and United States, wintering in southern United States, Mexico and Central America.[18][19] Status: least concern.[18]
Length up to 1.4 m (4.6 ft), wingspan 2–2.3 m (6.6–7.5 ft), weight 3.6–4.5 kg (7.9–9.9 lb).[20] Smallest pelican; distinguished by brown plumage; feeds by plunge-diving.[21]
Five subspecies. Coastal distribution ranging from North America and the Caribbean to northern South America and the Galapagos.[22] Status: least concern.[23]
Length up to 1.52 m (5.0 ft), wingspan 2.48 m (8.1 ft),[24] average weight 7 kg (15 lb).[25] Dark with a white stripe from the crown down the sides of the neck.
Monotypic. Pacific Coast of South America from Ecuador and Peru south through to southern Chile.[22] Status: near threatened.[26]
Length 1.40–1.75 m (4.6–5.7 ft), wingspan 2.45–2.95 m (8.0–9.7 ft), weight 10–11 kg (22–24 lb).[27][28] Plumage white, with pink facial patch and legs.
Monotypic. Patchy distribution from eastern Mediterranean east to Indochina and Malay Peninsula, and south to South Africa.[22] Status: least concern.[29]
Length 1.60–1.90 m (5.2–6.2 ft), wingspan 2.3–2.5 m (7.5–8.2 ft), weight 4–8.2 kg (8.8–18.1 lb).[30] Predominantly white with black along primaries and very large, pale pink bill.
Monotypic. Australia and New Guinea; vagrant to New Zealand, Solomons, Bismarck Archipelago, Fiji andWallacea.[22] Status: least concern.[31]
Length 1.25–1.32 m (4.1–4.3 ft), wingspan 2.65–2.9 m (8.7–9.5 ft),[32] weight 3.9–7 kg (8.6–15.4 lb).[33] Grey and white plumage, occasionally pinkish on the back, with a yellow upper mandible and grey pouch.[32]
Monotypic. Africa, Seychelles and southwestern Arabia;[22] extinct in Madagascar.[34] Status: least concern.[35]
Length 1.60–1.80 m (5.2–5.9 ft), wingspan 2.70–3.20 m (8.9–10.5 ft), weight 10–12 kg (22–26 lb).[27][28] Largest pelican; differs from great white pelican in having curly nape feathers, grey legs and greyish-white plumage.[32]
Monotypic. South-eastern Europe to India and China.[22] Status: near threatened.[36]
Length 1.27–1.52 m (4.2–5.0 ft), wingspan 2.5 m (8.2 ft), weight c. 5 kg (11 lb).[37] Mainly grey-white all over, with a grey hindneck crest in breeding season, pinkish rump and spotted bill pouch.[37]
Monotypic. Southern Asia from southern Pakistan across India east to Indonesia;[22] extinct in the Philippines and possibly eastern China.[37] Status: near threatened.[38]
Fossil record
The fossil record shows that the pelican lineage has existed for at least 36 million years; the oldest known pelican fossil was assigned toEopelecanus aegyptiacus and was found in lateEocene (middle to late part of the earlyPriabonian stage/age) deposits of the Birket Qarun Formation within theWadi Al-Hitan World Heritage Site in Egypt.[1] A more complete fossil pelican ofearly Oligocene age is known from deposits at theLuberon in southeastern France, and is remarkably similar to modern forms.[39] 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 at the time.[39] AnEarly Miocene fossil has been namedMiopelecanus gracilis on the basis of certain features originally considered unique, but later thought to lie within the range of interspecific variation inPelecanus.[39] The Late EoceneProtopelicanus may be apelecaniform orsuliform – or a similar aquatic bird such as a pseudotooth (Pelagornithidae), but is not generally considered apelecanid.[40][1] The supposedMiocene pelicanLiptornis fromPatagonia is anomen dubium (of doubtful validity), being based on fragments providing insufficient evidence to support a valid description.[41]
Fossil finds from North America have been meagre compared with Europe, which has a richer fossil record.[42] SeveralPelecanus species have been described from fossil material, including:[43]
A brown pelican opening mouth and inflating air sac to display tongue and some inner bill anatomyAmerican white pelican with knob which develops on bill before the breeding seasonAn adultbrown pelican with a chick in a nest in Chesapeake Bay,Maryland, US: This species will nest on the ground when no suitable trees are available.[49]Australian pelican displaying the extent of its throat pouch (Lakes Entrance, Victoria)
Pelicans are very large birds with very long bills characterised by a downcurved hook at the end of the upper mandible, and the attachment of a hugegular pouch to the lower. The slenderrami of the lower bill and the flexible tongue muscles form the pouch into a basket for catching fish, and sometimes rainwater,[15] though to not hinder the swallowing of large fish, the tongue itself is tiny.[50] They have a long neck and short stout legs with large, fully webbed feet. Although they are among the heaviest of flying birds,[51] they are relatively light for their apparent bulk because of air pockets in the skeleton and beneath the skin, enabling them to float high in the water.[15] The tail is short and square. The wings are long and broad, suitably shaped for soaring and gliding flight, and have the unusually large number of 30 to 35 secondaryflight feathers.[52]
Males are generally larger than females and have longer bills.[15] The smallest species is the brown pelican, small individuals of which can be no more than 2.75 kg (6.1 lb) and 1.06 m (3.5 ft) long, with a wingspan of as little as 1.83 m (6.0 ft). The largest is believed to be the Dalmatian, at up to 15 kg (33 lb) and 1.83 m (6.0 ft) in length, with a maximum wingspan of 3 m (9.8 ft). The Australian pelican's bill may grow up to 0.5 m (1.6 ft) long in large males,[53] the longest of any bird.[4]
Pelicans have mainly light-coloured plumage, the exceptions being the brown and Peruvian pelicans.[54] The bills, pouches, and bare facial skin of all species become brighter before breeding season commences.[55] The throat pouch of the Californian subspecies of the brown pelican turns bright red, and fades to yellow after the eggs are laid, while the throat pouch of the Peruvian pelican turns blue. The American white pelican grows a prominent knob on its bill that is shed once females have laid eggs.[5] The plumage of immature pelicans is darker than that of adults.[54] Newly hatched chicks are naked and pink, darkening to grey or black after four to 14 days, then developing a covering of white or greydown.[56]
Air sacs
Anatomical dissections of two brown pelicans in 1939 showed that pelicans have a network ofair sacs under their skin situated across the ventral surface including the throat, breast, and undersides of the wings, as well as having air sacs in their bones.[57] The air sacs are connected to the airways of the respiratory system, and the pelican can keep its air sacs inflated by closing itsglottis, but how air sacs are inflated is not clear.[57] The air sacs serve to keep the pelican remarkably buoyant in the water[58] and may also cushion the impact of the pelican's body on the water surface when they dive from flight into water to catch fish.[57] Superficial air sacs may also help to round body contours (especially over the abdomen, where surface protuberances may be caused by viscera changing size and position) to enable the overlying feathers to form more effective heat insulation and also to enable feathers to be held in position for good aerodynamics.[57]
Distribution and habitat
Modern pelicans are found on all continents except Antarctica. They primarily inhabit warm regions, although breeding ranges extend to latitudes of 45° South (Australian pelicans inTasmania) and 60° North (American white pelicans in western Canada).[4] Birds of inland and coastal waters, they are absent from polar regions, the deep ocean, oceanic islands (except the Galapagos), and inland South America, as well as from the eastern coast of South America from the mouth of theAmazon River southwards.[15]Subfossil bones have been recovered from as far south as New Zealand'sSouth Island,[59] although their scarcity and isolated occurrence suggests that these remains may have merely been vagrants from Australia (much as is the case today).[60]
Behaviour and ecology
An Australian pelican gliding with its large wings extended
Pelicans swim well with their strong legs and their webbed feet. They rub the backs of their heads on theirpreen glands to pick up an oily secretion, which they transfer to theirplumage to waterproof it.[4] Holding their wings only loosely against their bodies, pelicans float with relatively little of their bodies below the water surface.[32] They dissipate excess heat by gular flutter – rippling the skin of the throat and pouch with the bill open to promoteevaporative cooling.[15] They roost and loaf communally on beaches, sandbanks, and in shallow water.[15]
A fibrous layer deep in the breast muscles can hold the wings rigidly horizontal for gliding and soaring. Thus, they usethermals for soaring to heights of 3,000 m (10,000 ft) or more,[61] combined both with gliding and with flapping flight inV formation, to commute distances up to 150 km (93 mi) to feeding areas.[4] Pelicans also fly low (or "skim") over stretches of water, using a phenomenon known asground effect to reducedrag and increaselift. As the air flows between the wings and the water surface, it is compressed to a higher density and exerts a stronger upward force against the bird above.[62] Hence, substantial energy is saved while flying.[63]
Adult pelicans rely on visual displays and behaviour to communicate,[64] particularly using their wings and bills.Agonistic behaviour consists of thrusting and snapping at opponents with their bills, or lifting and waving their wings in a threatening manner.[65] Adult pelicans grunt when at the colony, but are generally silent elsewhere or outside breeding season.[32][66][67][68] Conversely, colonies are noisy, as chicks vocalise extensively.[64]
Pelicans aregregarious 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.[4] The location of the breeding colony is constrained by the availability of an ample supply of fish to eat, although pelicans can use thermals to soar and commute for hundreds of kilometres daily to fetch food.[55]
The Australian pelican has two reproductive strategies depending on the local degree of environmental predictability. Colonies of tens or hundreds, rarely thousands, of birds breed regularly on small coastal and subcoastal islands where food is seasonally or permanently available. In arid inland Australia, especially in the endorheicLake Eyre basin, pelicans breed opportunistically in very large numbers of up to 50,000 pairs, when irregular major floods, which may be many years apart, fill ephemeralsalt lakes and provide large amounts of food for several months before drying out again.[61]
In all species, copulation takes place at the nest site; it begins shortly after pairing and continues for three to ten days before egg-laying. The male brings the nesting material, in ground-nesting species (which may not build a nest) sometimes in the pouch, and in tree-nesting species crosswise in the bill. The female then heaps the material up to form a simple structure.[4]
The eggs are oval, white, and coarsely textured.[15] All species normally lay at least two eggs; the usual clutch size is one to three, rarely up to six.[15] Both sexes incubate with the eggs on top of or below the feet; they may display when changing shifts. Incubation takes 30–36 days;[15] hatching success for undisturbed pairs can be as high as 95%, but because of sibling competition orsiblicide, in the wild, usually all but one nestling dies within the first few weeks (later in the pink-backed and spot-billed species). Both parents feed their young. Small chicks are fed byregurgitation; after about a week, they are able to put their heads into their parents' pouches and feed themselves.[56] Sometimes before, but especially after being fed the pelican chick may seem to "throw a tantrum" by loudly vocalizing and dragging itself around in a circle by one wing and leg, striking its head on the ground or anything nearby and the tantrums sometimes end in what looks like aseizure that results in the chick falling briefly unconscious; the reason is not clearly known, but a common belief is that it is to draw attention to itself and away from any siblings who are waiting to be fed.[4]
Parents of ground-nesting species sometimes drag older young around roughly by the head before feeding them. From about 25 days old,[15] 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 six to eight weeks, they wander around, occasionally swimming, and may practise communal feeding.[4] Young of all speciesfledge ten to 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.[15] Overall breeding success is highly variable.[4] Pelicans live for 15 to 25 years in the wild, although one reached an age of 54 years in captivity.[55]
Feeding
The diet of pelicans usually consists of fish,[55] but occasionallyamphibians, turtles,crustaceans,insects, birds, and mammals are also eaten.[69][70][71] The size of the preferred prey fish varies depending on pelican species and location. For example, in Africa, the pink-backed pelican generally takes fish ranging in size fromfry up to 400 g (0.9 lb) and the great white pelican prefers somewhat larger fish, up to 600 g (1.3 lb), but in Europe, the latter species has been recorded taking fish up to 1,850 g (4.1 lb).[71] In deep water, white pelicans often fish alone. Nearer the shore, several 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.[72] Although all pelican species may feed in groups or alone, the Dalmatian, pink-backed, and spot-billed pelicans are the only ones to prefer solitary feeding. When fishing in groups, all pelican species have been known to work together to catch their prey, and Dalmatian pelicans may even cooperate withgreat cormorants.[71]
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 to distraction, and grab a fish from the open bill.[73] Pelicans in their turn sometimessnatch prey from other waterbirds.[4]
The brown pelican usually plunge-dives head-first for its prey, from a height as great as 10–20 m (33–66 ft), especially foranchovies andmenhaden.[74][72][71] The only other pelican to feed using a similar technique is the Peruvian pelican, but its dives are typically from a lower height than the brown pelican.[75] The Australian and American white pelicans may feed by low plunge-dives landing feet-first and then scooping up the prey with the beak, but they—as well as the remaining pelican species—primarily feed while swimming on the water.[71] Aquatic prey is most commonly taken at or near the water surface.[54] Although principally a fish eater, the Australian pelican is also an eclectic and opportunisticscavenger andcarnivore that forages inlandfill sites, as well as takingcarrion[76] and "anything from insects and small crustaceans to ducks and small dogs".[76] Food is not stored in a pelican's throat pouch, contrary to popular folklore.[55]
Globally, pelican populations are adversely affected by these main factors: declining supplies of fish through overfishing or water pollution, destruction of habitat, direct effects of human activity such as disturbance at nesting colonies, hunting and culling, entanglement in fishing lines and hooks, and the presence of pollutants such asDDT andendrin. Most species' populations are more or less stable, although three are classified by theIUCN as being at risk. All species breed readily in zoos, which is potentially useful for conservation management.[83]
The combined population of brown and Peruvian pelicans is estimated at 650,000 birds, with around 250,000 in the United States and Caribbean, and 400,000 in Peru.[a] TheNational Audubon Society estimates the global population of the brown pelican at 300,000.[85] Numbers of brown pelican plummeted in the 1950s and 1960s, largely as a consequence of environmental DDT pollution, and the species was listed as endangered in the US in 1970. With restrictions on DDT use in the US from 1972, its population has recovered, and it was delisted in 2009.[84][86]
The Peruvian pelican is listed as near threatened because, although the population is estimated byBirdLife International to exceed 500,000 mature individuals, and is possibly increasing, it has been much higher in the past. It declined dramatically during the 1998El Niño event and could experience similar declines in the future. Conservation needs include regular monitoring throughout the range to determine population trends, particularly after El Niño years, restricting human access to important breeding colonies, and assessing interactions with fisheries.[87]
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. Numbers declined substantially during the 20th century, one crucial factor being the eradication of the importantSittaung valley breeding colony in Burma through deforestation and the loss of feeding sites.[88] The chief threats it faces are from habitat loss and human disturbance, but populations have mostly stabilised following increased protection in India and Cambodia.[38]
The pink-backed pelican has a large population ranging over much ofsub-Saharan Africa. In the absence of substantial threats or evidence of declines across its range, its conservation status is assessed as being of least concern. Regional threats include the drainage of wetlands and increasing disturbance in southern Africa. The species is susceptible tobioaccumulation of toxins and the destruction of nesting trees by logging.[89]
The American white pelican has increased in numbers,[5] with its population estimated at over 157,000 birds in 2005, becoming more numerous east of the continental divide, while declining in the west.[90] However, whether its numbers have been affected by exposure to pesticides is unclear, as it has also lost habitat through wetland drainage and competition with recreational use of lakes and rivers.[5]
Great white pelicans range over a large area of Africa and southern Asia. The overall trend in numbers is uncertain, with a mix of regional populations that are increasing, declining, stable, or unknown, but no evidence has been found of rapid overall decline, and the status of the species is assessed as being of least concern. Threats include the drainage of wetlands, persecution and sport hunting, disturbance at the breeding colonies, and contamination by pesticides and heavy metals.[91]
The Dalmatian pelican has a population estimated at between 10,000 and 20,000 following massive declines in the 19th and 20th centuries. The main ongoing threats include hunting, especially in eastern Asia, disturbance, coastal development, collision with overhead power lines, and the over-exploitation of fish stocks.[92] It is listed as near threatened by the IUCN Red List of Threatened Species as the population trend is downwards, especially in Mongolia, where it is nearly extinct. However, several European colonies are increasing in size and the largest colony for the species, at theSmall Prespa Lake in Greece, has reached about 1,400 breeding pairs following conservation measures.[36]
Widespread across Australia,[5] the Australian pelican has a population generally estimated at between 300,000 and 500,000 individuals.[93] Overall population numbers fluctuate widely and erratically depending on wetland conditions and breeding success across the continent. The species is assessed as being of least concern.[94]
Culling and disturbance
Pelicans have been persecuted by humans for their perceived competition for fish, despite the fact that their diet overlaps little with fish caught by people.[5] Starting in the 1880s, American white pelicans were clubbed and shot, their eggs and young were deliberately destroyed, and their feeding and nesting sites were degraded by water management schemes and wetland drainage.[5] Even in the 21st century, an increase in the population of American white pelicans in southeastern Idaho in the US was seen to threaten the recreationalcutthroat trout fishery there, leading to official attempts to reduce pelican numbers through systematic harassment andculling.[95]
Great white pelicans on Dyer Island, in the Western Cape region of South Africa, were culled during the 19th century because their predation of the eggs and chicks ofguano-producing seabirds was seen to threaten the livelihood of the guano collectors.[78] More recently, such predation at South African seabird colonies has impacted on the conservation of threatened seabird populations, especially crowned cormorants, Cape cormorants, andbank cormorants. This has led to suggestions that pelican numbers should be controlled at vulnerable colonies.[78]
Apart fromhabitat destruction and deliberate, targeted persecution, pelicans are vulnerable to disturbance at their breeding colonies by birdwatchers, photographers, and other curious visitors. Human presence alone can cause the birds to accidentally displace or destroy their eggs, leave hatchlings exposed to predators and adverse weather, or even abandon their colonies completely.[96][97][98]
Oiled brown pelican being washed at a rescue center in Fort Jackson, 2010
DDT pollution in the environment was a major cause of decline of brown pelican populations in North America in the 1950s and 1960s. It entered the oceanicfood web, contaminating and accumulating in several species, including one of the pelican's primary food fish – thenorthern anchovy. ItsmetaboliteDDE is a reproductivetoxicant in pelicans and many other birds, causing eggshell thinning and weakening, and consequent breeding failure through the eggs being accidentally crushed by brooding birds. Since an effective ban on the use of DDT was implemented in the US in 1972, the eggshells of breeding brown pelicans there have thickened and their populations have largely recovered.[74][99]
In the late 1960s, following the major decline in brown pelican numbers in Louisiana from DDT poisoning, 500 pelicans were imported from Florida to augment and re-establish the population; over 300 subsequently died in April and May 1975 from poisoning by the pesticide endrin.[100] About 14,000 pelicans, including 7,500 American white pelicans, perished frombotulism after eating fish from theSalton Sea in 1990.[5] In 1991, abnormal numbers of brown pelicans andBrandt's cormorants died atSanta Cruz, California, when their food fish (anchovies) were contaminated withneurotoxicdomoic acid, produced by thediatomPseudo-nitzschia.[101]
As waterbirds that feed on fish, pelicans are highly susceptible tooil spills, both directly by being oiled and by the impact on their food resources. A 2007 report to the California Fish and Game Commission estimated that during the previous 20 years, some 500–1,000 brown pelicans had been affected by oil spills in California.[98] A 2011 report by theCenter for Biological Diversity, a year after the April 2010Deepwater Horizon oil spill, said that 932 brown pelicans had been collected after being affected by oiling and estimated that ten times that number had been harmed as a result of the spill.[102]
Where pelicans interact with fishers, through either sharing the same waters or scavenging for fishing refuse, they are especially vulnerable to being hooked and entangled in both active and discarded fishing lines.Fish hooks are swallowed or catch in the skin of the pouch or webbed feet, and strongmonofilament fishing line can become wound around bill, wings, or legs, resulting in crippling, starvation, and often death. Local rescue organisations have been established in North America and Australia by volunteers to treat and rehabilitate injured pelicans and other wildlife.[103][104][105]
Parasites and disease
As with otherbird families, pelicans are susceptible to a variety ofparasites.Avian malaria is carried by themosquitoCulex pipens, and high densities of these biting insects may force pelican colonies to be abandoned.Leeches may attach to thevent or sometimes the inside of the pouch.[106] A study of the parasites of the American white pelican found 75 different species, includingtapeworms,flukes,flies,fleas,ticks, andnematodes.
The brown pelican has a similarly extensive range of parasites. The nematodesContracaecum multipapillatum andC. mexicanum and thetrematodeRibeiroia ondatrae have caused illness and mortality in thePuerto Rican population, possibly endangering the pelican on this island.[107]
Many pelican parasites are found in other bird groups, but severallice are veryhost-specific.[108] Healthy pelicans can usually cope with their lice, but sick birds may carry hundreds of individuals, which hastens a sick bird's demise. The pouch lousePiagetiella peralis occurs in the pouch and so it cannot be removed bypreening. While this is usually not a serious problem even when present in such numbers that it covers the whole interior of the pouch, sometimes inflammation and bleeding may occur from it and harm the host.[108]
In May 2012, hundreds of Peruvian pelicans were reported to have perished inPeru from a combination of starvation androundworm infestation.[109]
The pelican (henet inEgyptian) was associated inAncient Egypt with death and theafterlife. It was depicted in art on the walls of tombs, and figured infunerary texts, as a protective symbol against snakes.Henet was also referred to in thePyramid Texts as the "mother of the king" and thus seen as a goddess. References in nonroyal funerarypapyri show that the pelican was believed to possess the ability to prophesy safe passage in theunderworld for someone who had died.[110]
Anorigin myth from theMurri people ofQueensland, cited byAndrew Lang, describes how theAustralian pelican acquired its black and white plumage. The story tells that the pelican was once a black bird. During a flood, he made a canoe to save drowning people. He fell in love with a woman and decided to save her, but she and her friends tricked him and escaped. The pelican consequently began preparing to go to war against them by daubing himself with white clay as war paint. Before he had finished, another pelican, on seeing such a strangepiebald creature, killed him with its beak, and all such pelicans have been black and white ever since.[113]
TheMoche people of ancientPeru worshipped nature.[114] They placed emphasis on animals and often depicted pelicans in their art.[115]
Alcatraz Island was given its name by the Spanish because of the large numbers ofbrown pelicans nesting there. The wordalcatraz is itself derived from the Arabical-caduos, a term used for a water-carrying vessel and likened to the pouch of the pelican. The English namealbatross is also derived by corruption of the Spanish word.[116][117]
Christianity
Statue of pelican wounding its breast to feed its chicksWWII 1944 Scottishblood donation poster
ThePhysiologus, a didactic Christian text from the 3rd or 4th century, claims that pelicans kill their young when they grow and strike their parents in the face, but then the mother laments them for three days, after which she strikes her side and brings them back to life with her blood.[118] ThePhysiologus explains this as mirroring the pain inflicted on God by people'sidolatry, and theself-sacrifice of Jesus on the cross whichredeems the sinful (see theblood and water gushing from the wound in his side).[118] This text was widely copied, translated, and sometimes closely paraphrased during theMiddle Ages, for instance by 13th-century authorsGuillaume le Clerc andBartholomaeus Anglicus.[118]
In a newer, also medieval version of the European myth, the pelican was thought to be particularly attentive to her young, to the point of providing them with blood by wounding her own breast when no other food was available. As a result, the pelican came to symbolise thePassion of Jesus and theEucharist,[119][120] supplementing the image of thelamb and the flag.[121] This mythical characteristic is referenced in the hymn "Adoro te devote" ("Humbly We Adore Thee"), where in the penultimate verse,Saint Thomas Aquinas describes Christ as the loving divine pelican, one drop of whose blood can save the world.[122] Similarly, the 1678Christian allegorical novelThe Pilgrim's Progress describes how "the pelican pierce[s] her own breast with her bill … to nourish her young ones with her blood, and thereby to show that Christ the blessed so loveth his young, his people, as to save them from death by his blood."[123]
The pelican is featured in many Christian artworks, especially in Europe. For example, the first (1611) edition of theKing James Bible contains a depiction of a pelican feeding her young in an oval panel at the bottom of the title page.[121] The "pelican in her piety" appears in the 1686reredos byGrinling Gibbons in the church ofSt Mary Abchurch in the City of London. Earlier medieval examples of the motif appear in painted murals, for example, the mural in theparish church ofBelchamp Walter, Essex (c. 1350).[125]
Queen Elizabeth I: thePelican Portrait, byNicholas Hilliard (circa 1573), in whichElizabeth I wears the medieval symbol of the pelican on her chest
The self-sacrificial characterization of the pelican was reinforced by widely read medievalbestiaries. The device of "a pelican in her piety" or "a pelican vulning (fromLatinvulnerō, "I wound, I injure, I hurt") herself" was used in religious iconography andheraldry.[4]
Origin in nature
The legends of self-wounding and the provision of blood occur across cultures.[4] For example, an Indian folktale depicts a pelican that killed her young by rough treatment, but was then so contrite that sheresurrected them with her own blood.[4] Such legends 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 to fully empty the pouch. Another possible derivation is the tendency of the bird to rest with its bill on its breast; theDalmatian pelican has a blood-red pouch in the early breeding season and this may have contributed to the myth.[4]
Heraldry
The arms of the Kiszely family ofBenedekfalva depict a "pelican in her piety" both in thecrest andshield.
Pelicans have featured extensively in heraldry, generally using the Christian symbolism of the pelican as a caring and self-sacrificing parent.[126] Heraldic images featuring a "pelican vulning" refers to a pelican injuring herself, while a "pelican in her piety" refers to a female pelican feeding her young with her own blood.[127] TheKing of PortugalJohn II adopted the pelican as is own personal sygil while he wasInfante, evoking the Christian symbology to equate the sacrifice of his blood to feed the nation. The pelican as a symbol also became synonymous with the increasing charity efforts of theSantas Casas da Misericórdia during his reign and the reconstruction of theHospital das Caldas da Rainha and theHospital Real de Todos-os-Santos, which were mainly patronaged by his wifeD. Leonor.[128]
The medical faculties ofCharles University in Prague also have a pelican as their emblem.[131] The symbol of theIrish 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.[132] The heraldic pelican also ended up as a pub name and image, though sometimes with the image of the shipGolden Hind.[133] SirFrancis Drake's famous ship was initially calledPelican, and adorned the Britishhalfpenny coin.[134]
The pelican is the subject of a popularlimerick originally composed byDixon Lanier Merritt in 1910 with several variations by other authors.[142] The original version ran:[143]
A wonderful bird is the pelican, His bill will hold more than his belican, He can take in his beak Food enough for a week, But I'm damned if I see how the helican.
Notes
^The US government has not accepted the elevation of the two taxa into separate species.[84]
^abcdefghijklmnopNelson, J. Bryan; Schreiber, Elizabeth Anne; Schreiber, Ralph W. (2003)."Pelicans". In Perrins, Christopher (ed.).Firefly Encyclopedia of Birds. Richmond Hill, Ontario: Firefly Books. pp. 78–81.ISBN1-55297-777-3.
^abcSimpson, J.; Weiner, E., eds. (1989). "Pelican".Oxford English Dictionary (2nd ed.). Oxford, United Kingdom: Clarendon Press. p. 1299.ISBN0-19-861186-2.
^Bock, Walter J. (1994).History and Nomenclature of Avian Family-Group Names. Bulletin of the American Museum of Natural History. Vol. 222. New York: American Museum of Natural History. pp. 131, 252.hdl:2246/830.
^abHackett, S.J.; Kimball, R.T.; Reddy, S.; Bowie, R.C.K.; Braun, E.L.; Braun, M.J.; Chojnowski, J.L.; Cox, W.A.; Han, K.-L.; Harshman, J.; Huddleston, C.J.; Marks, B.D.; Miglia, K.J.; Moore, W.A.; Sheldon, F.H.; Steadman, D.W.; Witt, C.C.; Yuri, T. (2008). "A Phylogenomic Study of Birds Reveals Their Evolutionary History".Science.320 (5884):1763–68.Bibcode:2008Sci...320.1763H.doi:10.1126/science.1157704.PMID18583609.S2CID6472805.
^"Brown Pelican"(PDF).Endangered Species Program information sheet. US Fish & Wildlife Service. November 2009. Archived fromthe original(PDF) on 11 November 2012. Retrieved9 June 2012.
^abSnow, David; Perrins, Christopher M, eds. (1998).The Birds of the Western Palearctic concise edition (2 volumes). Oxford, United Kingdom: Oxford University Press. pp. 93–98.ISBN0-19-854099-X.
^Olson, Storrs L. (1985). "Faunal Turnover in South American Fossil Avifaunas: the Insufficiencies of the Fossil Record".Evolution.39 (5):1174–77.doi:10.2307/2408747.JSTOR2408747.PMID28561505.
^Widhalm, J. (1886). "Die Fossilen Vogel-Knochen der Odessaer-Steppen-Kalk-Steinbrüche an der Neuen Slobodka bei Odessa".Schriften der Neurussische Gesellschaft der Naturforscher zu Odessa (in German).10:3–9.
^Thomas, Bob (2 June 2011)."Bird Flight Over Water".College of Social Sciences Intranet. New Orleans, Louisiana: Center for Environmental Communication, Loyola University. Archived fromthe original on 12 April 2016. Retrieved1 August 2012.
^Vestjens, W. J. M. (1977). "Breeding Behaviour and Ecology of the Australian Pelican,Pelecanus Conspicillatus, in New South Wales".Wildlife Research.4:37–58.doi:10.1071/WR9770037.
^abAnon (1980).National accomplishments in pollution control, 1970–1980: some case histories. U.S. Environmental Protection Agency, Office of Planning and Evaluation. pp. 183–184.ISBN1236274539.
^Jaramillo, A. (2009). "Humboldt Current seabirding in Chile".Neotropical Birding.4:27–39.
^abHandbook of Australian, New Zealand and Antarctic Birds. Volume 1, Ratites to Ducks. Marchant, S.; Higgins, P.J. (Coordinators). Melbourne, Victoria: Oxford University Press. 1990. p. 742.ISBN0-19-553068-3.{{cite book}}: CS1 maint: others (link)
^abSmith, A.C.M.; U. Munro (2008). "Cannibalism in the Australian Pelican (Pelecanus conspicillatus) and Australian White Ibis (Threskiornis molucca)".Waterbirds: The International Journal of Waterbird Biology.31 (4):632–635.
^Cursach, J.A.; J.R. Rau; J. Vilugrón (2016). "Presence of the Peruvian Pelican Pelicanus thagus in seabird colonies of Chilean Patagonia".Marine Ornithology.44:27–30.
^abDaigre, M.; P. Arce; A. Simeone (2012). "Fledgling Peruvian Pelicans (Pelecanus thagus) attack and consume younger unrelated conspecifics".Wilson Journal of Ornithology.124 (3):603–607.doi:10.1676/12-011.1.S2CID84928683.
^Gunderson, Dan (16 May 2012)."Loving 'em to death".Statewide. MPR News. Retrieved14 February 2017.
^abBurkett, Esther; Logsdon, Randi J.; Fien, Kristi M. (2007).Status Review of California Brown Pelican(PDF). California Fish and Game Commission Reports. U.S. Environmental Protection Agency, Office of Planning and Evaluation. Archived fromthe original(PDF) on 20 December 2011.
^Ehrlich, Paul R.; Dobkin, David S.; Wheye, Darryl (1988)."DDT and Birds". Stanford University. Retrieved6 August 2012.
^Work, Thierry M.; Barr, Bradd; Beale, Allison M.; Fritz, Lawrence; Quilliam, Michael A.; Wright, Jeffrey L.C. (1993). "Epidemiology of domoic acid poisoning in Brown Pelicans (Pelecanus occidentalis) and Brandt's Cormorants (Phalacrocorax penicillatus) in California".Journal of Zoo and Wildlife Medicine.24 (1):54–62.JSTOR20460314.
^"A Deadly Toll"(PDF).Report. Center for Biological Diversity. April 2011. Retrieved6 August 2012.
^Hart, George (2005).The Routledge Dictionary of Egyptian Gods And Goddesses. Routledge Dictionaries. Abingdon, United Kingdom: Routledge. p. 125.ISBN978-0-415-34495-1.
^Lang, Andrew (2005) [1887].Myth, Ritual & Religion, Volume 1. New York, New York: Cosimo Inc. pp. 140–41.ISBN978-1-59605-204-8.
^Benson, Elizabeth (1972)The Mochica: A Culture of Peru New York: Praeger Press.
^Berrin, Kathleen; Larco Museum (1997).The Spirit of Ancient Peru:Treasures from the Larco Museum. New York, New York:Thames and Hudson.ISBN0500018022.
^"College Crest". Cambridge, United Kingdom: Corpus Christi College, Cambridge University. 2011. Archived fromthe original on 19 July 2012. Retrieved2 May 2012.
^"Corpus Christi". Corpus Christi College, Oxford. Retrieved2 May 2012.
_(bird)_in_Mo_(by).jpg)
.jpg)
_at_nest_with_chicks_in_Uppalpadu_W_IMG_2663.jpg)
_feeding_a_juvenile_in_Garapadu%2c_AP_W_IMG_5362.jpg)
.jpg)
_Tortuga_Bay_on_the_Island_of_Santa_Cruz%2c_Gal%25C3%25A1pagos.JPG)
_-_Portrait_of_Queen_Elizabeth_I_-_Google_Art_Project.jpg)
