pelecaniform

pelecaniform [pel΄ ə kan′ə fôrm΄]

adj.

of, or having the nature of, an order (Pelecaniformes) of swimming birds having all four toes connected in a webbed foot, including pelicans and cormorants

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▪ bird

Introduction

   any of the relatively large and diverse group of aquatic birds (bird) that share the common characteristic of webbing between all four toes. Order Pelecaniformes contains the three suborders Fregatae, Pelecani, and Phaethontes and the six families Ahingidae (ahingas or snakebirds (snakebird)), Phalacrocoracidae (cormorants (cormorant)), Phaethontidae (tropic birds (tropic bird)), Fregatidae (frigate birds (frigate bird)), Sulidae (gannets (gannet) and boobies (booby)), and Pelecanidae (pelicans (pelican)). All pelecaniforms are relatively large birds: they range in length from about 40 cm (about 16 inches), excluding the elongated central tail feathers, in the white-tailed tropic bird (Phaethon lepturus) to 1.8 metres (6 feet) in the Dalmatian pelican (Pelecanus crispus).

General features

 In terms of their way of life, the pelecaniform birds (bird) fall into four adaptive groups: the frigate birds (frigate bird) are long-winged masters of piracy and aerial pursuit of surface-living marine prey; the tropic birds (tropic bird) and boobies (booby) are wide-ranging flyers that capture prey underwater by plunging from a height; the pelicans (pelican) are large, large-billed, long-necked, buoyant birds that fish mostly by reaching down while swimming at the surface; and the cormorants (cormorant) and anhingas (snakebird) are heavy-bodied, long-necked, underwater swimmers, respectively pursuing and lying in wait for their prey below the surface. The anhingas are largely confined to fresh water; pelicans and cormorants occur in both freshwater and marine habitats; and the other groups are entirely marine.

Importance to humans

      Most pelecaniform birds are of rather little significance to humans, but the guano (excrement) of cormorants, boobies, and pelicans is an important fertilizer. Exploitation of old accumulations of guano reached its peak in the mid-19th century, and since then only the current production of guano has been available in most areas, but even this provides a substantial resource where the bird populations are large. There were an estimated 18 million guano birds on the coast of Peru early in the 1960s. Of these, about 15 million were guanay cormorants (Phalacrocorax bougainvillii), and the remainder were Peruvian boobies (Sula variegata) and brown pelicans (Pelecanus occidentalis). The harvest of guano at that time amounted to about 180,000 tons per year. The guano birds in the area feed largely on the Peruvian anchovy, and, now that this fish is directly exploited on a large scale for fish meal and fish oil, guano-producing birds have declined to fewer than five million animals. In Walvis Bay, Namib., artificial platforms have been constructed in coastal lagoons and on an offshore reef, greatly facilitating the collection of guano.

      Some pelecaniforms are thought to compete with humans for fish. Cormorants are often accused of reducing sport fish populations, which engenders a strong negative backlash by anglers in both Europe and North America. Typically, cormorants are convenient scapegoats for poor fishing due to other causes. Increasingly, however, they have a negative impact on the fish populations of aquaculture ponds in the southeastern United States. Conversely, cormorants are used by fishermen in some countries (such as China and Japan) to catch fish, which they retrieve to a waiting boat; a collar prevents the cormorants from swallowing the fish for themselves.

      Pelicans, feeding on fish from inland and coastal waters, are among the animals whose diet tends to ensure that they will accumulate residues of insecticides (insecticide) (especially DDT) in their bodies. Among the physiological effects of these substances on birds are changes in calcium metabolism that result in their laying eggs with abnormally thin shells or no shells at all; these eggs usually break before hatching. These effects prevented the successful reproduction of brown pelicans during the early 1970s on the coasts of California and the Gulf of Mexico in the United States, and their populations became endangered as a result. Since the ban on DDT went into effect in the United States at the end of 1972, brown pelicans have been able to rebound in substantial numbers.

Distribution and abundance

      Pelecaniform birds occur all over the world except in parts of the interior of North America, Africa, Asia, and Australia, in the high Arctic, and in most of Antarctica. Two groups, the tropic birds and frigate birds, are essentially confined to the tropics; the boobies, pelicans, and anhingas are widespread in the tropics but also penetrate far into the temperate zones; and cormorants breed from the Equator to the Arctic Circle and the Antarctic Peninsula.

      The tropic birds are the most pelagic (free-flying over open ocean) of the pelecaniforms and can be seen even in the most unproductive central parts of tropical seas. Some boobies and frigate birds are found far out at sea, but most of them return to land to roost at night. Pelicans, cormorants, and anhingas do not venture far from land but may commute some distance between roosting or breeding places and their feeding grounds.

Natural history

Reproduction

      The breeding of pelecaniform birds (bird) is essentially restricted to places free of mammalian predators. Tropic birds (tropic bird), boobies (booby), and frigate birds (frigate bird) typically breed on oceanic islands or on small islets and stacks off continental coasts. Where man introduced predators (such as cats) to isolated islands, pelecaniforms and other marine birds were often eliminated from their traditional breeding grounds. On Ascension Island and St. Helena in the South Atlantic, for example, tropic birds, frigate birds, and boobies, which used to breed in large numbers on the main islands, have either been exterminated or have been forced to confine their nesting to sites on small offshore islets. Pelicans (pelican) and cormorants (cormorant) are not normally found on islands far from continental land but breed on islands in lakes or offshore or in other protected sites such as trees standing in water or on cliffs. Anhingas (snakebird) breed in trees or bushes close to the sheltered waters where they prefer to feed.

      Pelecaniform breeding, whether on the ground or in trees, is typically colonial, apparently because of the scarcity of safe places. If they are to breed at all, all the birds of an area must crowd into the available space. In practice, most colonies have a fairly definite upper limit of density determined by the distance that an incubating bird can reach out to repel intruders. Nests of the gannet (Morus bassanus) have an average density of about 1 per square metre (about 0.84 per square yard); those of the guanay cormorant average about 3.5 per square metre (about 2.9 per square yard).

      In some species, colonial breeding has become obligatory, and single pairs or small groups do not breed successfully. Other species breed colonially only where there is a shortage of space for nesting. The masked booby (Sula dactylatra), for example, breeds in dense colonies on islets off Ascension Island but in dispersed patterns on Christmas Island (Pacific). Breeding in a number of species is normally dispersed; the red-footed cormorant (Phalacrocorax gaimardi) of South America, for instance, often nests on cliffs where only scattered sites are available. Similarly, tropic birds nest in holes or crevices in cliffs or under the shelter of rocks or bushes. The sites are thus normally well dispersed, and in some places there is intense competition for them.

courtship behaviour

      During courtship (except in tropic birds) the male typically selects a nest site and then displays (display behaviour) from this site to passing females. In the male advertising display of boobies (“sky-pointing”) and of cormorants and anhingas (“wing-waving”), the head and tail are raised and the wings (wing) partially raised. The boobies have species-specific positions of the wings and tail; the cormorants and anhingas wave either their wing tips or their whole wings, to a varying degree. In many species a particular call is associated with the display. In the “rattling” display of frigate birds, the male leans back on his tail, extends his wings along the ground, and with his bill pointing vertically throbs his inflated scarlet throat (gular) sac, vibrates his wings, and claps his bill.

      In most pelecaniforms, when a female has finally joined a male at the nest site, he may go and fetch nest material, which is then added to the nest structure by the female, alone or with the assistance of the male. There is no courtship feeding in the pelecaniforms, but in tropic birds there are a few records of one adult feeding the other during incubation. Copulation normally occurs on the nest site, without any special precopulatory or postcopulatory displays. The pair-bond, once established, is reinforced by such activities as joint nest building and defense and preening of one bird by the other. Certain displays functioning in individual recognition are performed in the air or on the ground when one of a pair returns to the nest and also occur during the “handling” of nest material, which continues during incubation. The boobies have a “head-wagging” recognition display; the cormorants and anhingas have a comparable “kink-throat” display; similarly, the frigate birds use a “rattling” display.

      In tropic birds, which do not normally have nest sites in the open, courtship display is aerial. Birds fly around in a group, calling, and then two individuals may leave the group and fly together, the upper bird depressing his tail (especially the long central feathers) during gliding or hovering; the flights often end with one or both birds flying into a nest cavity. Tropic birds do not collect nest material but probably sometimes enlarge nest cavities with their bills or feet.

      In all or nearly all pelecaniform families, mating frequently occurs with the same partner in successive years, and in many populations the same nest site may be occupied repeatedly. Since most species nest colonially, fighting over nest sites is common. It is particularly intense in the gannet and in populations of tropic birds when nest sites are in short supply.

Breeding seasons

      Outside the tropics, egg laying by pelicans, boobies, and cormorants occurs typically in the spring, and the laying period extends over one to three months. In the tropics there is greater diversity in both the timing and extent of the laying period. Although many tropical populations have distinct laying periods at the same season in each year, a number of those in relatively seasonless environments have been known to lay in every month of the year. The blue-footed booby (Sula nebouxii) in the Galapagos, for example, is known to breed at intervals of less than a year. In other populations, laying is synchronized but occurs at nonannual intervals; the brown booby (S. leucogaster) on Ascension Island has laying periods roughly every eight months. For the cormorants (P. carbo and P. africanus) of Lake Victoria, conditions are favourable for breeding during about eight months in each year, but the scarcity of islands free of predators forces the birds to occupy the few safe colonies in relays.

Nests, eggs, and young

      Most pelecaniforms build substantial but untidy nests of twigs, grass, algae, or feathers; in many species, guano accumulates on and around the nest as breeding progresses. Many cormorants and pelicans habitually build their nests on the ground, but most species that normally breed where trees or bushes are available build their nests in them. Certain cormorants and boobies sometimes nest on cliff ledges, and tropic birds and most boobies lay their eggs on the open ground in insignificant depressions.

      There is much variation in clutch size. In general, species living in the unproductive tropical seas lay only a single egg. Those living in seasonally productive areas at higher latitudes, or in areas where surface waters are enriched by the upwelling of nutrient laden water from deeper layers, lay larger clutches and raise several young at a time. Tropic birds and frigate birds all have clutches of one egg, but pelicans generally lay one to three eggs, cormorants two to five, and anhingas three to five. The boobies show a more complex picture. The gannet of the temperate zones and the tropical red-footed booby (Sula sula) have clutches of one egg, but the other typical boobies of the tropics—the masked and brown boobies—normally lay two eggs. These two-egg species, however, normally raise only one chick, the older nestling apparently pushing the younger out of the nest soon after hatching. The second egg in these ground-nesting species seems to function as an insurance against infertility, loss, or breakage of the first, risks that may be lower in the tree-nesting red-footed booby. The single-egg clutch of the gannet has not been satisfactorily explained, since pairs have been able to raise two chicks when an extra has been provided experimentally.

      Except in the tropic birds, the eggs of pelecaniforms are small in relation to body size. In the tropic birds they are 9–13 percent of the adult weight, in the frigate birds 6 percent, in the boobies 3–5 percent, and in the pelicans and cormorants 2–3 percent. In species with clutches of more than one, eggs are laid at intervals as short as 24 hours or as long as five to six days in the case of the masked booby. In all the groups, both parents participate roughly equally in incubation and feeding of the young. The lengths of the incubation shifts vary extensively, both within and between species. In some species (such as many cormorants) the shifts last only a few hours, but in tropic birds and some boobies shifts of three or more days are not uncommon; in the great frigate bird (Fregata minor), in the Galapagos, shifts last 10–15 days.

      Incubation generally starts with the laying of the first egg (though not in some cormorants). First and last young may hatch several days apart and often differ substantially in size during the nestling period. The incubation period is just under four weeks in anhingas, four to four and a half weeks in cormorants, about four and a half weeks in pelicans, six weeks in tropic birds, and six to eight weeks in frigate birds.

      The young, which are helpless and essentially naked at hatching, are brooded continuously for several days until their down grows. The tropic birds are exceptional in that they have thick down at hatching and are sometimes left alone by their parents even on the day of hatching. The rate of development of the young varies dramatically and is correlated with the characteristics of the environments in which the birds live. The fledging period in the pelagic tropical masked booby is double that of cormorants of comparable body size living in the cool, productive waters off the coast of North America (17 weeks versus less than 8 weeks). Brown pelicans in North America fledge in less time (9 weeks) than white-tailed tropic birds weighing only one-tenth as much (10 weeks) and in less than half the time needed by tropical frigate birds weighing about one-third as much (more than 20 weeks).

      The young of all pelecaniforms are fed on regurgitated food that they obtain by inserting their heads into the mouths of their parents. In frigate birds and tropical boobies, the parents continue to feed their young for a long period after they can fly, apparently because the young take a long time to become sufficiently skillful in hunting to survive in tropical areas where food is generally scarce. The most extreme case is that of the great frigate bird, in which the young can fly at about five months of age but sometimes are fed for more than a year after this. In the tropic birds and the gannet, however, the young are not fed by the parents after fledging; they leave their birthplace either by flying straight out to sea or by fluttering down to the water and swimming out to sea.

molt

      In the pelecaniforms, as in most birds, replacement of the feathers (molt) occurs mainly in the intervals between breeding. In a few species molt progresses concurrently with breeding activities, but normally a complete molt commences when breeding is over. In the anhingas the primary flight feathers of the wing are shed simultaneously, but in the other groups molt is gradual. The primary and secondary feathers typically are replaced by means of two or three molt waves that start several feathers apart and move outward in parallel until all feathers have been shed. In most if not all pelecaniforms, feathers of the head and neck and some on the body are replaced a second time shortly before the next breeding season. Special feathers (crests, neck plumes, white patches) acquired at this time are shed around the time of egg laying in some species; in pelicans some head feathers are replaced yet again during incubation.

      In many of the pelecaniforms the colours of the soft parts (especially the bare skin on the face) change as the birds become sexually active. In some pelicans a horny triangular plate grows on the upper mandible toward the tip before the breeding season and is shed after laying.

Ecology

Survival and mortality

      Full-grown pelecaniform birds have few natural enemies, and, although some are taken by marine predators, they are generally long-lived. Nestlings, recently fledged young, and sometimes even adult birds suffer heavy mortality when food shortages occur. In many tropical areas where food is generally scarce, irregular fluctuations in the supply lead to drastic variations in breeding success. Mortality from starvation also occurs among birds of richer seas, most dramatically along the west coast of South America in years when oceanographic changes cut off the normally abundant food supply of the huge bird populations. Even under average conditions, young pelecaniforms in their first year after fledging experience much higher mortality than adults. In the European shag (P. aristotelis), more than half the young die during this period, although among adults annual mortality is only about 15 percent in males and 20 percent in females. In the British population of the gannet, about 80 percent of the fledglings die before reaching breeding age (about five years), most during their first year. Adults die at an average rate of less than 6 percent per year and have a mean expectation of life exceeding 16 years. Age records for individual banded birds include a great frigate bird that was recently found breeding at an age of more than 30 years and a masked booby breeding at about 23 years. Negative evidence suggests, however, that few individuals of these species survive much longer than the two examples.

Population movements

      Members of the pelecaniform populations breeding at high latitudes generally move to lower latitudes in winter but do not perform transequatorial migrations. Adequate analyses of banding results have been carried out on only a few species. Great cormorants (P. carbo) from Britain do not show a well-defined migration but disperse random distances from the colonies. In all populations of gannets, juveniles move further than adults. A number of juvenile gannets from New Zealand have been found in eastern Australia only a week after leaving their natal colonies on their first flight, having made journeys exceeding 2,600 km (1,600 miles) at average speeds of more than 370 km (230 miles) per day. Gannets that survive to breeding age usually return to the same cluster of nests where they were hatched. In a similar way, juvenile lesser frigate birds (F. ariel) from colonies in the central equatorial Pacific disperse on a broad front, many moving over 6,500 km (4,000 miles) from their natal island. Their pattern of dispersal may be related to that of the prevailing winds in the area.

Feeding ecology

      The various groups of pelecaniform birds are specialized for different ways of life and in particular for different feeding methods. Tropic birds and boobies, though they differ in many ways, are both adapted for catching prey underwater by plunging from the air. They are powerful flyers and buoyant swimmers, but, although their feet have large webs, their legs are not streamlined, and they are not specialized for fast swimming. The feet are probably used underwater more for steering than for propulsion, and the birds depend mainly on the impetus of the dive to enable them to approach their prey at high speed.

      Tropic birds are adept at hovering while locating prey, and probably obtain most of their food near the surface. Many boobies often dive from greater heights and probably go deeper than tropic birds; the gannet sometimes dives from more than 30 metres (about 100 feet). Blue-footed boobies, when hunting in groups, tend to dive almost simultaneously. A disyllabic whistle is often heard from such groups as they start to dive and may be a signal given by the initiator. It has been suggested that the simultaneous plunging of several birds may confuse the fish in a school and so increase each bird's chance of catching one. Red-footed boobies, and perhaps also other boobies, catch flying fish (family Exocoetidae) in the air as well as in the water. These fish, with squid of the family Ommastrephidae, are staple foods of the tropical boobies and the tropic birds. Boobies of upwelling zones and the gannet feed almost entirely on fish.

      Frigate birds are among the most aerial of all sea birds. Their best-known feeding habit involves piracy, in which they harry other sea birds until they disgorge their prey, after which the frigate birds catch the food in the air or pick it from the surface. They also catch prey for themselves, however, pursuing flying fish when they leap through the air and snatching fish (and probably squid) from the surface without alighting.

      Pelicans generally catch their prey while swimming, thrusting their long bills and long necks below the surface to scoop up fish in their distensible throat pouches. When fishing in shallow water, pelicans often cooperate to form a kind of living net. They form a crescent facing the shore, or even a circle, and then close in, splashing and paddling hard. Each bird keeps station until the fish panic and can be captured as they try to escape between the birds. Pelicans have also been observed herding and capturing ducklings in a similar way. The brown pelican, unlike the other species, often forages some distance offshore and habitually fishes by plunging from the air.

      Cormorants and anhingas are adapted for underwater swimming. The cormorants pursue free-swimming or bottom-living fish, and some species also eat mollusks and other invertebrate animals. When feeding on schooling fish, cormorants often engage in mass fishing activities in which a flock advances in a long line stretching at right angles to the direction of movement, apparently with the fish fleeing ahead of them. The birds swim forward while above the surface and also while pursuing prey underwater, and laggards fly forward and land just ahead of the line. Anhingas do not pursue their prey but lie in wait underwater and then stab passing fish. Most of the fish that they eat are slow swimming and laterally flattened.

      The physiological adaptations for diving have been little studied in cormorants and anhingas. It is known, however, that for cormorants feeding on the bottom the mean length of the dives is directly related to water depth. Dives of most species usually last less than half a minute, although dives as long as one minute are not uncommon. The time spent resting after a dive averages between one-half and one-third of the duration of the dive; if dives are very long, the length of the rest periods approaches that of the dives. Anhingas often stay underwater for as long as two minutes, and a dive of nearly seven minutes has been recorded in a captive bird.

Form and function

Distinguishing characteristics

 Few generalizations can be made about the gross morphology of the pelecaniforms, since their external form reflects the diversity of their adaptations. In the larger and more aerial species, the skeleton is extensively pneumatized: nearly all the bones contain air sacs that are connected with the respiratory system. In the frigate birds (frigate bird), pelicans (pelican), and boobies (booby), the major limb bones are no more than thin-walled hollow tubes with some internal struts. Pneumatization is least developed in anhingas (snakebird), which need to have a high specific gravity to remain underwater while motionless.

      Those pelecaniforms that plunge into the water from a height (boobies, tropic birds (tropic bird), and the brown pelican) have a system of air sacs under the skin. These air sacs, connected with the lungs, form a spongy mattress that presumably protects the bird as it strikes the water. The appearance of this structure in species of pelicans that do not plunge suggests that the buoyancy it confers may be advantageous in itself.

      The reduction of the external nostrils in most pelecaniforms is probably also connected with diving and swimming underwater. The closure, complete in boobies and in adult cormorants (cormorant) and anhingas, is only partial in frigate birds and pelicans; the nostrils are developed normally in tropic birds. The reduction of the nostrils in the aerial Fregatae as well as in the highly aquatic Pelecani suggests (as does the structure of the foot) that the Fregatae originated from a more aquatic stock. In the birds with nonfunctional nostrils, air enters the mouth cavity directly from the outside and then reaches the lungs in the normal way via the glottis—that is, the muscular opening to the windpipe—at the base of the tongue. In boobies and cormorants entry of air to the mouth when the bill is closed is by means of secondary external nostrils. These are slitlike openings at the angle of the mouth on each side where a horny flap (jugal operculum) at the base of the upper mandible overlaps the lower mandible.

      Tropic birds, pelicans, and boobies have water-repellent plumage and are very buoyant. They swim high in the water and can reach significant depths only by acquiring momentum during aerial dives; their return to the surface is rapid. Frigate birds have wettable plumage, cannot swim or dive, and quickly become waterlogged if they alight on the water. In cormorants and anhingas the large contour feathers are wettable and trap little air when the birds are swimming underwater; this keeps buoyancy low in these birds and enables them to submerge from the surface without benefit of an aerial dive. The compressible but dense body plumage remains dry near the skin, at least in many species. The hind limbs of anhingas, which are not specialized for fast swimming, are relatively efficient for climbing and perching. Anhingas do not normally take flight from the water but climb out and dry their wings (wing) in a characteristic spread-wing posture that is also commonly used by cormorants. Once dry, anhingas show highly maneuverable flight and can also soar well, in contrast to the cormorants, which have a higher wing loading. Many species of pelecaniforms habitually spread their wings in “sun-bathing” postures, even when dry. The function of this behaviour is not well understood.

Physiological adaptations

      Most pelecaniform birds nest in exposed situations, and special adaptations are necessary to prevent overheating of adults and young, particularly in tropical areas. The masked booby in the Galapagos, for example, stands with its back to the sun, thus shading its feet and naked throat pouch and permitting loss of heat by convection and conduction. It elevates its scapular (shoulder) feathers and droops its wings, allowing maximum exposure of shaded feather areas, and it exhibits sustained fluttering of the throat pouch. Throat fluttering, which permits evaporative cooling with minimum expenditure of energy, is used under heat stress by all the pelecaniforms except the tropic birds, which do not have a naked throat pouch. The rate of throat fluttering remains roughly constant under increasing heat stress but becomes continuous instead of intermittent, and there are increases in its amplitude and in the throat area involved. All species make use of panting to increase evaporative cooling. Under heat stress the rate of breathing increases with increasing body temperature, and, when it reaches a maximum, the amplitude of breathing increases.

      Although even the newly hatched young of certain species are capable of throat fluttering, they cannot regulate their temperature when left in the sun, until they have acquired down and in some species not until they are several weeks old. Young tropic birds provide an exception: they have dense down at hatching and are sometimes left alone for long periods within a few days of hatching; small chicks are capable of regulating their body temperature by panting, even in the sun, but most nests are in the shade.

      Evaporative cooling requires much water, which is obtained from the food and probably also by drinking seawater. The resulting high intake of salt necessitates special adaptations for elimination of the excess. Marine birds have highly developed salt excretory glands, the lateral nasal glands, which function when the amount of salt taken into the body is more than the kidneys can deal with. The glands secrete a concentrated solution of sodium chloride, which flows through ducts to the nasal cavity. In most sea birds, including the tropic birds, it then flows out through the nostrils and may drip or be shaken off the tip of the bill. In boobies and cormorants, in which the nostrils are closed, the solution trickles from the internal nares in the roof of the mouth and out at the end of the bill. In some species of cormorants that occur in both marine and freshwater habitats, the salt gland is relatively larger in individuals from marine habitats, presumably reflecting higher salt intake.

Evolution and paleontology

      The origin of the pelecaniforms is a matter of some debate. Some authorities assert that pelecaniforms diverged from other birds as late as the early Eocene Epoch (about 54 million years ago); however, available evidence suggests that the stocks leading to the Procellariiformes (procellariiform), typical pelecaniform birds (bird) (Fregatae plus Pelecani), Phaethontes, and Laridae (gulls (gull) and terns (tern)), diverged during the Cretaceous Period, probably at least 70 million years ago. Nevertheless, there is widespread agreement that pelecaniforms diversified into the six extant families by the early Miocene Epoch (some 23 million years ago).

Classification

Distinguishing taxonomic features

      The classification of the order Pelecaniformes is based mainly on morphological features. Recent attempts have been made to clarify relationships by analysis of egg white proteins and by comparative analysis of social behaviour. The parasitic feather lice (Mallophaga) found on members of the different groups also provide clues to relationships.

      The six modern families of pelecaniform birds are readily distinguished, although the relationships among them are not firmly established. The families are internally homogeneous, and in each of them the modern species are often placed in single genera; the Phaethontidae, Fregatidae, Pelecanidae, and Anhingidae are usually treated in this way.

Annotated classification

Order Pelecaniformes

 Aquatic birds with totipalmate feet (webs connecting all 4 toes); 1st toe (hallux) not pointing backward as in most birds but turned inward and joined by a web to 2nd toe. 66 species.

      Suborder Phaethontes

 

      Family Phaethontidae (tropic birds (tropic bird))

 Medium sized (16–19 inches or 40–48 cm long). Plumage satiny white with black markings; central 2 tail feathers much elongated in adult. Sexes similar. Bill laterally compressed, slightly decurved, and with undivided horny sheath (rhamphotheca). Nostrils open in slits; nasal glands each with 2 ducts. No naked gular (throat) pouch. Neck short, 14–15 cervical vertebrae. Pelvic girdle reduced, legs short, set far back. No nest-building behaviour. Single egg heavily marked with brown and black. Chick hatched covered with gray down. 1 genus, Phaethon; 3 living species, worldwide in tropical oceans.

      Suborder Fregatae

 

      Family Fregatidae (frigate birds (frigate bird))

 Large (31–34 inches or 79–86 cm long), light bodied. Plumage blackish, juveniles and some adults with white underparts; juveniles with white heads, sometimes tinged orange. Wings very long, pointed; tail deeply forked. Sexes differ in size (female larger) and in coloration of plumage and soft parts. Bill long, slender, strongly hooked at tip; horny sheath divided by deep grooves. Nostrils almost closed; nasal glands each with single duct. Small gular pouch, highly distensible and brilliant red in courting male. Neck short, 14–15 cervical vertebrae. Furculum (wishbone) fused with coracoids and with sternum (breastbone). Pelvic girdle and leg muscles reduced; legs very short. Feet small, webbing only between bases of toes. One white, thin-shelled egg. Young naked at birth, soon acquire thick white down. 1 genus, Fregata; 5 living species; worldwide in tropical oceans.

      Suborder Pelecani

 Horny sheath of bill divided by deep grooves (largely suppressed in anhingas). Naked gular pouch; facial skin and pouch colourful, especially in breeding season. Nasal glands each with a single duct. Neck long, with 17–20 cervical (neck) vertebrae; 8th and 9th have articulations permitting sharp S-bend, especially in anhingas. Pelvic girdle not reduced, legs strong. Feet large, fully webbed. Eggs thick shelled, with chalky white outer layer. Young naked at hatching.

      Family Pelecanidae (pelicans (pelican))

 Very large (50–72 inches or 127–180 cm long). Plumage white, brown, or gray; some species crested. Wings long, broad; tail short. Sexes similar. Bill long, hooked at tip, with enormous gular pouch suspended from highly distensible rami (lateral elements) of lower mandible. Nostrils almost closed. Furcula fused with sternum. Down of nestlings white. 1 genus, Pelecanus; 8 species; seacoasts and freshwater lakes of southern Africa, central and eastern Eurasia, Australia, and North America; north and west coasts of South America.

      Family Sulidae (boobies (booby))

 Large (26–40 inches or 66–102 cm long). Plumage of most species white with black on wings; some brown and white; juveniles largely gray or gray-brown. Wings long and narrow, tail rather long, wedge-shaped. Sexes generally differ in voice, sometimes in colour of bare skin. Bill stout, conical, slightly decurved near the tip and finely serrated on cutting edges. Nostrils closed. Down of nestlings white. 3 genera, 10 living species, widespread but irregular distribution, on seacoasts and oceanic islands.

      Family Phalacrocoracidae (cormorants (cormorant))

 Medium to large (19–40 inches or 48–102 cm long). Plumage usually blackish; some with white underparts, and a few largely gray; some species have ornamental crests or patches of white when breeding. Wings broad, tail long and stiff. Sexes similar, but females up to 30 percent lighter in weight. Bill slender, sharply hooked at tip. Nostrils closed in adults. Bony occipital style projecting upward from back of skull. Down of nestlings dark. At least 1 genus, about 40 species.

      Family Anhingidae (anhingas (snakebird) or darters)

 Large (34–36 inches or 86–91 cm long). Plumage mainly black and brown with pale markings on back and wings; breeding male has light-coloured plumes on head and neck. Tail very long and stiff. In some forms the sexes differ in colour. Bill long, straight, slender, sharply pointed; cutting edges serrated toward tip. Nostrils closed in adults. Occipital style smaller than in cormorants. Head small, neck long and slender. Legs short but strong. Pyloric lobe of stomach with mat of hairlike processes. Down of nestlings brown. 1 extant genus, Anhinga; 2 species. 5 fossil species, in the genera Protoplotus (the earliest from the Eocene of Sumatra), and Anhinga.

Critical appraisal

      Ornithologists continue to debate whether the order Pelecaniformes is a natural evolutionary unit, since the unifying characters, such as the totipalmate foot, may have arisen through convergent evolution. In particular, the tropic birds (Phaethontes) differ in many ways from the other members of the order Pelecaniformes and may well be wrongly placed in this order. They are united to the others primarily by the totipalmate condition, but the first toe in Phaethontes is distinctly more elevated on the tarsometatarsus than in the other groups. Some of the resemblances of the Phaethontes to the Laridae (gulls and terns) in the order Charadriiformes (charadriiform) may not result from evolutionary convergence but may indicate that the Phaethontes arose from forms more closely related to the basal stock of the Laridae than to that of the Pelecaniformes.

      The frigate birds (Fregatae) are generally treated as a distinct suborder, on the basis of a number of morphological characters that they share with members of the order Procellariiformes but not with the Pelecani.

      There is little doubt that the suborder Pelecani is made up of closely related groups. The pelicans are sometimes placed in a separate superfamily, but it is more useful to emphasize the similarities among the modern groups by keeping them together. The anhingas have sometimes been treated as a subfamily of the Phalacrocoracidae, but their morphological and ecological differences from cormorants perhaps justify separation at the family level.

N. Philip Ashmole

Additional Reading

G.F. Van Tets, A Comparative Study of Some Social Communication Patterns in the Pelecaniformes (1966), discusses the behaviour and relationships of the different groups. Bryan Nelson, Galápagos: Islands of Birds (1968), contains a readable and authoritative account of the ecology and behaviour of tropical pelecaniforms, especially boobies and frigatebirds, and his The Sulidae (1978), is an extensive treatment of gannets and boobies.N. Philip Ashmole Ed.

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