Pelecaniformespelicans, tropicbirds, cormorants, and relatives

Pele­can­i­formes com­prises six fam­i­lies (Phaethon­ti­dae (trop­icbirds), Sul­i­dae (boo­bies and gan­nets), Pha­lacro­co­raci­dae (cor­morants and shags), An­hingi­dae (an­hin­gas), Pele­canidae (pel­i­cans) and Fre­gati­dae (frigate­birds)), six gen­era, and 67 species.

Pele­can­i­form birds are dis­trib­uted world­wide, pri­mar­ily in coastal and ma­rine zones.

Pele­can­i­form birds in­habit ma­rine and in­land wa­ters. Habi­tat varies from pelagic to coastal to in­land fresh­wa­ter en­vi­ron­ments.

In­cu­ba­tion ranges from 25-55 days. Young are al­tri­cial and all chicks, ex­cept trop­icbirds, are born with­out natal down. Fledg­ing age is highly vari­able, rang­ing from 65-210 days. Post-fledg­ing care is pro­nounced and may last for as long as 18 months in some species. Age at first breed­ing varies with species and ranges from two to per­haps eleven years.

Pele­can­i­form birds are medium to large aquatic birds with toti­palmate feet (all four toes joined by web). Most have dis­ten­si­ble gular sac lo­cated be­tween the branches of the lower mandible. In trop­icbirds the small gular re­gion is feath­ered, in pel­i­cans the bare sac is pen­du­lous, and in frigate­birds the large, bare, red gular sac is in­flat­able. Only trop­icbirds have ex­posed ex­ter­nal nares. All pele­can­i­form birds' skulls lack supra­or­bital groove for a nasal gland and dor­sal ver­te­brae are opistho­coelous. Plumage ranges from mostly black with some white to mostly white with some black. Feet and bare re­gions of face and gular sac may be­come brightly col­ored dur­ing breed­ing. Most have desmog­nathous palate, ex­cept trop­icbirds, which have schizog­nathous palate. All ex­cept frigate­birds lack brood patches.

Many pele­can­i­forms breed in mixed colonies with other pele­can­i­forms, gulls, terns, or pen­guins.

Pele­can­i­form birds feed pri­mar­ily on fish and squid. They may also prey on mol­lusks, crus­taceans, am­phib­ians, rep­tiles, and in­ver­te­brates.

Preda­tors of pele­can­i­form birds in­clude hu­mans, birds, rats, cats, dogs, and rep­tiles.

Most pele­can­i­form birds are con­sid­ered sea­son­ally monog­a­mous. Nest-sites and mates may change from year to year, ex­cept per­haps for sulids. Mat­ing dis­plays and pair for­ma­tion be­hav­iors are elab­o­rate and com­plex. Gen­er­ally males dis­play to at­tract fe­males. Fe­males de­fend the nest-site and con­struct the nest from ma­te­ri­als col­lected by the male. Cop­u­la­tions gen­er­ally occur at the nest or dis­play site.

Breed­ing may be an­nual or bi­en­nial, sea­sonal or year round. Pele­can­i­form birds nest in colonies, some­times con­sist­ing of thou­sands of pairs. Nest sites are vari­able, lo­cated on the ground, in trees or shrubs, or on cliff ledges. Nests may be in­sub­stan­tial scrapes or con­struc­tions of twigs and other ma­te­ri­als. Clutch size varies with species and ranges from one to six eggs. Egg color ranges from white to pale green-blue to red-brown.

In pele­can­i­form birds both sexes take ap­prox­i­mately equal stints in­cu­bat­ing eggs. All species ex­cept the frigatids lack brood patches and use foot web­bing to trans­fer heat to eggs. Du­ra­tion of in­cu­ba­tion varies con­sid­er­ably from 24 -57 days. Both par­ents brood and feed chicks re­gur­gi­tated food. Many species have post-fledg­ing care where par­ents con­tinue to feed chicks, in some species post-fledg­ing care is pro­longed and may last for as long as 18 months.

Some pele­can­i­forms are mi­gra­tory whereas oth­ers are mostly seden­tary with wide dis­per­sal of young. All pele­can­i­forms are aquatic birds and as­so­ci­ated with large bod­ies of water. For­ag­ing tech­niques vary from hunt­ing singly to for­ag­ing co­op­er­a­tively in large groups, from plunge div­ing to sur­face div­ing, from un­der­wa­ter stalk­ing am­bushes to aer­ial and un­der­wa­ter pur­suit div­ing. Species with large gular sacs use them in con­junc­tion with for­ag­ing, mat­ing dis­plays and ther­moreg­u­la­tion.

Most pele­can­i­forms are gre­gar­i­ous, gath­er­ing in colonies for breed­ing and roost­ing in large groups.

Pele­can­i­forms are no­tably noisy when in large groups. Vo­cal­iza­tions are vari­able with species, and range from the sharp pierc­ing whis­tle of the trop­icbirds to the gut­tural grunt­ing of cor­morants.

Hu­mans ex­ploit pele­can­i­form birds ex­ten­sively. The eggs, chicks, and/or adults of many species are col­lected for con­sump­tion, pur­ported med­i­c­i­nal pur­poses, or for goods and cloth­ing. Hu­mans col­lect and sell pele­can­i­form guano as fer­til­izer. Some species are trained and used by hu­mans in con­junc­tion with fish­ing.

Twenty-two pele­can­i­form species are in­cluded in the IUCN Red List of Threat­ened Species.

One species, Pal­las's Cor­morant (Pha­lacro­co­rax per­spic­il­la­tus), is listed as 'Ex­tinct'; two are listed as 'Crit­i­cally En­dan­gered' (An­drew's Frigate­bird (Fre­gata an­drewsi) and Ab­bott's Booby (Pa­pa­sula ab­botti)); and two species are listed as 'En­dan­gered', (Flight­less Cor­morant (Pha­lacro­co­rax har­risi) and Chatham Is­land Shag (P. on­slowi)). Of the re­main­ing species, eleven are listed as 'Vul­ner­a­ble' and six are 'Lower Risk'. Major threats in­clude habi­tat de­struc­tion, in­tro­duced preda­tors, pes­ti­cide poi­son­ing and oil spills.

The evo­lu­tion­ary re­la­tion­ships of pele­can­i­form birds re­main un­clear and the mono­phyly of Pele­can­i­formes is strongly de­bated. Tra­di­tion­ally, Pele­can­i­formes (or Steganopodes) is a group com­pris­ing six fam­i­lies in­clud­ing:(Phaethon­ti­dae (trop­icbirds), Sul­i­dae (boo­bies and gan­nets), Pha­lacro­co­raci­dae (cor­morants and shags), An­hingi­dae (an­hin­gas), Pele­canidae (pel­i­cans) and Fre­gati­dae (frigate­birds). Analy­ses of mor­pho­log­i­cal and etho­log­i­cal ev­i­dence sug­gest that phaethon­tids are rel­a­tively prim­i­tive whereas frigatids are rel­a­tively most de­rived of the group. In con­trast, fos­sil ev­i­dence ap­pears to sup­port the hy­poth­e­sis that phaethon­tids and frigatids fall basal rel­a­tive to the rest of the group. Some mor­pho­log­i­cal ev­i­dence sup­ports the mono­phyly of Pele­can­i­formes and its sis­ter group re­la­tion­ships with pro­cel­lari­ids (al­ba­trosses, pe­trels, shear­wa­ters) and ci­coni­ids (storks, herons, ibises, spoon­bills). Some mol­e­c­u­lar, etho­log­i­cal, and mor­pho­log­i­cal ev­i­dence re­futes the mono­phyly of Pele­can­i­formes. One hi­er­ar­chy based on DNA hy­bridiza­tion places phaethon­tids sis­ter to a group com­pris­ing sulids, an­hingids, and pha­lacro­co­racids; pele­canids as sis­ter to a Shoe­bill Stork (Bal­aeni­ceps); and frigatids as sis­ter to sphenis­cids (pen­guins) and in­cluded in a group (Pro­cel­lar­i­oidea) along with gavi­ids (loons) and pro­cel­lari­ids (pe­trels, shear­wa­ters, al­ba­trosses).

The old­est pele­can­i­form fos­sils in­clude a phaethon­tid (Prophaethon) from Eng­land dat­ing from the early Eocene and a fre­gatid (Lim­nofre­gata) from Wyoming also dat­ing to the Eocene. Pha­lacro­co­racid fos­sils ex­tend back to the Eocene-Oligocene bound­ary.

Sulid (Sula ron­zoni) and pele­canid (Pele­canus gra­cilis) fos­sils from France date from the early Oligocene and lower Miocene re­spec­tively. Old and New world fos­sil an­hingids date from the early (An­hinga sub­volans) and the late (A. pan­non­ica, A. gran­dis) Miocene.

Camp­bell, B., and E. Lack, ed­i­tors. 1985. A Dic­tio­nary of Birds. Buteo Books, Ver­mil­lion, SD.

del Hoyo, J., El­liott, A. & Sar­gatal, J. (eds.) 1992. Hand­bood of the Birds of the World. Vol. 1. Lynx Edi­cions, Barcelona.

Fe­duc­cia, A. 1999. The Ori­gin and Evo­lu­tion of Birds, 2nd edi­tion. Yale Uni­ver­sity Press New Haven.

Johns­gard, P. A. 1993. Cor­morants, Darters, and Pel­i­cans of the World. Smith­son­ian In­sti­tu­tion Press Wash­ing­ton.

Sib­ley, C. G. & J. E. Ahlquist. 1990. Phy­logeny and Clas­si­fi­ca­tion of Birds, A Study in Mol­e­c­u­lar Evo­lu­tion. Yale Univ. Press.

Siegel-Causey, D. 1988. Phy­logeny of the Pha­lacro­co­raci­dae. Con­dor 90:885-905.

Van Tets, G. F. 1976. Aus­trala­sia and the ori­gin of shags and cor­morants, Pha­lacro­co­raci­dae. Pp. 121-24 in: Proc. 16th Inter. Or­nith. Congr. Can­berra, Aus­tralia, 1974.

Con­trib­u­tors

Laura Howard (au­thor), An­i­mal Di­ver­sity Web.

Glossary

acoustic

uses sound to communicate

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

motile

having the capacity to move from one place to another.

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

tactile

uses touch to communicate

visual

uses sight to communicate