caprimulgiform

▪ order of birds

Introduction

also called  nightjars 

  any of about 120 species of soft-plumaged birds (bird), the major groups of which are called nightjars (nightjar), nighthawks (nighthawk), potoos (potoo), frogmouths (frogmouth), and owlet-frogmouths (owlet frogmouth). The order also includes the aberrant oilbird of South America. Most are twilight- or night-flying birds. Many produce sounds that are startling, strange, or weirdly beautiful. The calls of caprimulgiforms are surrounded by an aura of mystery richly endowed to elicit interest and sometimes fear from humans. The name of the type genus Caprimulgus, goatsucker, derives from an ancient belief that the birds seen flitting about the goats at dusk were taking milk from the goats' udders, a misconception no doubt fortified by the birds' uncommonly large mouths. In actuality, caprimulgiforms prey on the insects disturbed or attracted by the goats. There is now a tendency to replace the name goatsucker with the more appropriate term nightjar, derived from the birds' voices.

      The caprimulgiform birds are sparrow- to raven-sized birds (14–55 cm [about 5.5–22 inches]) with enormous gapes. They are cryptically coloured and have a patterned plumage, short legs, and (for the most part) long wings.

      All caprimulgiform birds are rather similar in general appearance, but each family has certain peculiar characteristics both in form and in habits. Caprimulgiforms resemble owls (owl) in many ways; however, there are numerous differences, mostly internal, between the two groups. Externally, caprimulgiforms possess bills and feet that are not raptorial, a flatter head with eyes placed laterally rather than in a frontal facial disk, relatively shorter tarsi, and longer tails. A closer inspection of caprimulgiforms and owls reveals a dissimilarity in the number of primary feathers in the wing and, usually, of secondaries and tail feathers.

      Although the true nightjars (Caprimulgidae) are amply distributed throughout the world, the other families are more restricted. The order is absent from New Zealand and some oceanic islands.

Natural history

 The caprimulgiform birds (bird) are primarily crepuscular, their activity being largely limited to the periods of dawn and dusk, although they are also nocturnal when there is sufficient illumination, especially by moonlight. Some species may become active on dark, cloudy days, and a few are somewhat diurnal. Oilbirds (oilbird) possess a system of echolocation that permits them to fly freely in total darkness, an adaptation related to their roosting and nesting in caves, and are thus implicitly well equipped for nocturnal life. They leave their caves at dusk and return at dawn and are highly gregarious in their foraging behaviour as well as in roosting and nesting. The other members of the order are more or less solitary. Some nightjars (nightjar) that are migratory behave gregariously during migration and to some extent while in their wintering regions.

      Although aerial feeders, most of the true nightjars roost on the ground, rocks, or fallen trunks, but some prefer horizontal branches of trees, in which case they usually perch lengthwise along the branch. Some ground roosters, however, will seek higher perches as singing or foraging posts; these are often slender branches or vines and the birds sit on them crosswise. Some species may even roost so perched. Unlike the nightjars, the frogmouths (frogmouth), potoos (potoo), and owlet-frogmouths (owlet frogmouth) are arboreal (tree-dwelling). The last normally sit crosswise on a branch and fairly upright, both when active and at rest, resembling small, long-tailed owls (owl). Potoos and frogmouths frequently sit crosswise and upright when actively foraging and also appear very owl-like, although at rest they may perch quite differently. The potoos are noted for a peculiar stance, perching usually at the top of a stump, broken branch, or at a knob on an upwardly inclined limb. When alarmed in daylight they slowly flatten their plumage and stretch their bills upward in a stiff posture with the eyes nearly closed and the bill slightly open. Frogmouths adopt a similar broken-branch alarm posture.

      During the day the oilbird perches horizontally on ledges inside caves, usually on its nest. Ledges are limited and mostly occupied by nests. On its forays outside the oilbird has been observed to perch on the bare parts of tree branches. Its mien is rather hawklike (hawk), but its crouching stance, unique in some ways, is more like that of a nightjar.

Habitat

      The caprimulgiform birds exploit ample food sources that are almost uniquely theirs. There are no other primarily frugivorous nocturnal birds to compete with the oilbird, its distribution being limited by the availability of suitable caves in regions providing the proper fruit for food.

      The nightjars, by far the largest and most successful group, have no effective avian competitors for the great numbers of night-flying insects except among themselves. Different nightjars have come to occupy almost all habitats, from semidesert to very humid regions and from sea level to 4,000 metres (about 13,100 feet) in altitude. They have a further advantage by preferring the more open areas: savannas, grasslands, and forest clearings. When nightjars are woodland dwellers, they live in the more open forests or brushwood growths; many use the woodlands only for diurnal concealment or nesting, doing most of their feeding at the edge or in clearings.

      Competition between the nightjars and potoos is reduced because the latter hunt mostly from higher perches in open country or exploit the space above the great forests, a vast niche occupied by practically no other nocturnal bird.

      Frogmouths and owlet-frogmouths inhabit forested savannas, forest edge, second growth, and the more open forests. Thus, their chief competitors are the insectivorous owls. As with the nightjars, differences in food preferences and methods of hunting minimize this competition. The hole-nesting owlet-frogmouths, however, have to contend not only with owls but also many diurnal species for suitable nesting sites, and this may explain why they are apparently the least successful family in the order, as suggested by their limited distribution and paucity of numbers.

Food habits

      Except for the oilbird, caprimulgiform birds are insectivorous, with some of the larger species occasionally carnivorous. Many of the nightjars and nighthawks (nighthawk) feed in continuous hawking flight, which may be rather erratic as they pursue their prey. Others of the order make short sallies from arboreal or terrestrial perches in the manner of flycatchers (flycatcher), frequently taking advantage of the increased visibility afforded by roads. Many nightjars take crawling insects by swooping from arboreal perches or by running along the ground, being far more agile on their feet than is generally believed. Almost all insects are taken, but beetles predominate in the food of many species. Also important are moths, the winged forms of termites and ants, and mosquitos; even small birds are taken. There seems to be little difference between the general kinds of food items taken by the New World and Old World nightjars (subfamily Caprimulginae), but the Old World species rely more on foraging in continuous flight than do most in the New World, where aerial foraging is largely limited to the nighthawks (Chordeilinae).

      Potoos sally from exposed arboreal perches, sometimes fairly close to the ground, to feed on essentially the same types of insects as the nightjars. The prey of frogmouths is mostly terrestrial. In short sallies from elevated perches, frogmouths capture large, crawling arthropods on the ground or on branches. Snails, frogs, mice, small birds, and occasionally fruit also are taken. Owlet-frogmouths feed mostly on terrestrial prey, much in the manner of the frogmouths, but also sally from a perch to take flying insects.

      Aberrant in so many ways, oilbirds are the only nocturnal, exclusively frugivorous birds. The fruits eaten are primarily of various trees of the palm, laurel, and bursera families and have large hard seeds. The firm, fleshy pericarp of the fruit is exceedingly rich in protein and oily fats. The fruits are swallowed whole and the seeds later regurgitated. Oilbirds possess an efficient olfactory sense; they employ scent in locating certain of the fruit-bearing trees. The fruits of palms, however, are not aromatic and are no doubt located by sight. The actual collection of the fruit seems to depend on sight. In feeding, the birds reach out with their bills and pluck the individual fruits while hovering. When feeding on large compact bunches of fruit, they sometimes cling briefly with their feet.

      Males of most if not all nightjars have patches of white feathers (feather) that serve as signals in mating. These patches are concealed when the bird is at rest but flashed in courtship displays. Males of several species in Africa and South America develop elongated wing or tail feathers that function in courtship and are lost following the breeding season.

Vocalizations

      The primary signals for mating and territorial purposes in most caprimulgiforms, however, are vocal and mechanical sounds. The voice is usually well developed, often having such distinctive patterns that the birds are named onomatopoeically ( whippoorwill, chuck-will's-widow, etc.). The almost human lament of the common potoo (Nyctibius griseus) in South America has in some places earned this species the name “poor-me-one,” and both it and the grotesque bawl of the great potoo (N. grandis) have been the source of many superstitious beliefs and legends. Frogmouths sing with various nasal booming, hooting, or croaking sounds, and the little-known owlet-frogmouths are reported to give owl-like whistles, churring calls, and a loud hissing note. Among the true nightjars, some species produce slowly or rapidly repeated churring sounds, in some cases rather toadlike, while others produce a warbled or whistled song, not at all disagreeable. Whistled phrases may be repeated hundreds of times without a break. Many different types of calls are used in different contexts, but the primary song, different for each species, appears to be the most important mechanism for species recognition. Besides vocal sounds, many species produce mechanical sounds in displays, such as claps and whirrs of the wings. In some species the vocalizations are given primarily in flight, in others primarily when perched; still others have different calls or songs when flying than when perched.

      Oilbirds produce a variety of sounds, from clucking calls, apparently used to maintain contact, to harsh snarls and screams, when excited. They seem to have no primary song, which may be superfluous to their gregarious and specialized mode of life. Of greatest interest are the rapidly pulsated clicks that are emitted in shorter or longer bursts and that are used by the oilbird for navigation in total darkness on the principle of echolocation. These clicks are in the range of 7,000 cycles per second and thus audible to humans, in contrast to the ultrasonic pulsations used by bats in their system of echolocation. So far as is known, the only other birds to employ echolocation are some of the cave swiftlets (swiftlet) (Collocalia). To what extent oilbirds use echolocation outside the caves is not known. Like their relatives in the order, they have large, sensitive eyes and probably navigate visually outside their caves.

Reproduction

      Nesting habits are diverse among caprimulgiform families. Nightjars make no nest, depositing their eggs directly on the ground or on the leaf-covered floor of woodland. Occasionally they use a slight natural depression or scratch out a place among the debris. Populations of a few species (notably the North American nighthawks) have adapted to urban life and nest on flat gravel-covered rooftops. The one or two slightly glossy eggs may be cryptically coloured (profusely marbled with pink, buff, brown, or gray) or contrast markedly with their surroundings (white to buff, usually with slight lines or blotches of colour). Incubation lasts about 19 days and may be by both adults or by only the female; there is considerable variation in this behaviour even within some species.

      In contrast to the simplified nesting of nightjars, frogmouths construct nests on the horizontal forks of trees. In the genus Podargus the nest is of twigs and other plant matter and the two or three eggs are white. In Batrachostomus the nest is a pad of the birds' own down, bound and camouflaged externally with cobwebs and lichens, one white egg being laid. Both sexes are believed to incubate, the period being about 30 days. Owlet-frogmouths nest mostly in hollow trees but also in tunnels in banks. They lay three or four white eggs, usually on a mat of leaves or fur. The incubation period is as yet unknown. The potoos are the most highly restricted nesters, for they must find a branch or stub with a suitable depression or crevice of just the right size to accommodate the single egg they lay. The egg is white, marked with brown and gray, and is incubated by both parents for 30 to 35 days.

      On ledges in the caves they inhabit, the oilbirds build up a shallow nest rim of viscous, regurgitated fruit matter, which sets to a firm structure. As the accumulation of seeds raises the level inside the nest, the rim is constantly added to, so as to maintain a slight depression. The nests are used year after year and gradually grow to low mounds. Inevitably, some seeds and the feces of very young nestlings contribute to the fabric of the rim. Two to four eggs are laid (white, subelliptical, slightly rough surface); both sexes incubate, the period being about 33 days.

      The chicks of nightjars are semi-precocial: they are down-covered (buff to brownish, plain or mottled, highly cryptic), fed by both parents, capable of opening their eyes on the first day (though normally keeping them closed for several days), capable of walking on the first day, and able to hop or run very well by the third or fourth day. They usually run with wings raised, resembling young ducklings in gait. From the beginning, chicks may solicit feeding by walking to the front of the adult, reaching up, and pecking at its bill. During the first days they are fed a regurgitated whitish viscous substance but later only partially digested insects. As early as the second day, although sometimes not until a week later, the chicks are led to different resting sites, at first only a metre or so at a time but gradually much greater distances. Some of this moving may result from the chicks' own wanderings, for they are often restless and leave the covering adult.

      In many cases both parents brood the chicks, in others only the female does so, and rarely the male assumes the primary role. By about two weeks the young are well feathered and too large for successful brooding; they rest at first near a parent but later quite alone.

      Undisturbed young birds fly voluntarily at 22 to 28 days of age; disturbed young may fly much earlier. By the time they are able to fly, they begin a certain amount of feeding on their own but are still largely dependent on their parents up to 40 to 50 days. When two young hatch, one of them frequently develops more slowly. If it survives, however, it may require up to 40 days to reach the state of voluntary flight.

      The nestlings of frogmouths, potoos, and owlet-frogmouths are semi-dependent, are covered with whitish down, are fed by both parents, and remain in the nest until fledged. One observer noted that a young common potoo began wandering over the boughs of the nest tree at about four weeks. This same nestling first made trial flights at 47 days and finally left the nest tree when 50 days old. Other reports indicate the nestling period of potoos to be of 40–45 days, that of frogmouths about 30 days. The young are brooded only during the first half of this period, by which time young potoos have attained the juvenile plumage (whitish mottled with brownish) and are already accomplished in assuming the “broken-branch” posture of adults.

      On the whole, the caprimulgiform birds, like most birds, breed solely or mostly at the time of maximum abundance of food supply. Oilbirds may nest to some extent throughout the year, but the maximum nesting takes place during that time of year when the greatest number of appropriate trees are fruiting. Baby oilbirds are helpless (altricial), hatched blind and virtually naked, with only a little sparse down. They have an exceedingly slow rate of development. A thicker coat of down replaces the initial one after three weeks, and the adult plumage begins to appear at five weeks, there being no special juvenile plumage. The nestlings are apparently fed semidigested fruit pulp during the first days. At about two weeks there is a gradual change to whole fruits, regurgitated by the adults into the mouths of the young. As a result of the oil-rich fruit diet, young oilbirds build up extraordinary deposits of fat, which led to economic exploitation. The fat can be boiled down to a high-quality, durable oil, suitable for cooking and lighting. For this reason, oilbirds were once subject to heavy predation by their human neighbours. Today this practice occurs on a much-reduced scale because of a general trend toward conservation. The fat buildup in an oilbird reaches its maximum at about 70 days, at which time a young oilbird may weigh half again as much as an adult. By this time the plumage is well developed but not fully grown. The weight decreases as the feathers grow until the age of 100–120 days, when the young leave the nest. At this time they fly well and look like adults. Both parents participate in feeding and brooding, the latter ceasing after about four weeks.

migration

      A number of tropical caprimulgiforms are sedentary, but the widespread and cosmopolitan nightjars exhibit all degrees of migration. These migrations may be short, even local, or quite long. Some populations of the common nighthawk (Chordeiles minor) of North America and the European nightjar (Caprimulgus europaeus) migrate south to Argentina and South Africa, respectively. Actually, transequatorial migrations in both directions occur, as birds take advantage of the general alternation of seasons, and hence of food supply, on opposite sides of the Equator.

Hibernation

      Some nightjars are able to cope with temporary food shortages by entering into periods of torpidity (torpor), a faculty they share with some swifts, hummingbirds, and a few others. The only known instances of apparently regular, prolonged annual hibernation in these birds, however, are reported for the poorwill Phalaenoptilus nuttallii, a nightjar. A banded individual was observed hibernating in the same small hollow in a rock during several successive winters. The bird was inert, with respiration and heart rates reduced to almost immeasurable levels and body temperature about 22 °C (40 °F) below normal. Subsequent laboratory experiments have shown that several species of nightjar have the ability to relax into a torpid state under abnormal conditions. Such an efficient means of conserving energy may be more generally possessed than is presently realized throughout a family whose habits largely limit the birds to two brief periods of feeding a day, causing them to be especially vulnerable to involuntary fasting if inclement weather should prevent feeding during those periods.

Form and function

 Members of the order Caprimulgiformes are easily recognized by their extremely wide mouths, large eyes, short legs and small, weak feet, and, except in the oilbird, soft plumage, in which browns and grays predominate.

      The size of the gape is astonishing. When the bird opens its mouth, the opening seems to span the entire head, which is nearly the case. The beak, always somewhat hooked, is large and horny in the oilbird and frogmouths (frogmouth) but reduced to a small projection in the remaining families. In body proportions, caprimulgiforms often appear chunky on account of the fluffiness of their plumage, but their actual bodies are proportionately no stouter than most songbirds (songbird). The tail is of medium length, except in males of those species with ornamental tail features. The wings are medium to long and are rounded in most species, pointed in a few, especially the nighthawks (nighthawk).

      The plumage of all forms presents unexcelled examples of natural camouflage. Coloured in rufescent to ochreous browns, grays, white, and black, the species are variously patterned in greatest accord with their normal surroundings during daytime rest. Those nightjars that roost inside woodlands are streaked and spotted in a way resembling fallen leaves and other detritus on the forest floor. Those that dwell on gravelly terrain are speckled or otherwise patterned. The latter often contrast rather than blend with the soil, appearing like one of the many stones scattered about. The owlet-frogmouths appear as clusters of dead leaves. Potoos and frogmouths are streaked and mottled like bark, so that in the daytime-alarm posture they appear most effectively as dead stubs.

      Caprimulgiform species are frequently dichromatic, having grayish and reddish phases. These colour patterns appear to be randomly distributed in some cases and sex-linked in others, there being several species of nightjars in which males are more grayish, females more reddish.

      The eye's iris colour is brown in nearly all members of the order. Yellow occurs in the irises of some potoos, and yellow, orange, and ruby occur in frogmouths. Contrary to some reports, oilbirds do not have blue irises, nor are any nightjars known to have yellow eyes.

Evolution and paleontology

      The nightjars (nightjar) and related families date back as a group to the Eocene Epoch (about 56–34 million years ago) and are not as old as many other orders of birds. Their closest living relatives are now known to be the owls (owl) and the swifts (swift). The owlet-frogmouths (owlet frogmouth) (Aegithalidae) are the oldest surviving lineage of the early Caprimulgiformes. DNA studies link owlet-frogmouths to the tree swifts (Hemiprocnidae). Fossils of owlet-frogmouths, frogmouths (frogmouth), and potoos (potoo) from Europe suggest much wider global distributions than their present limited distributions in Australasia and South America. Oilbirds (oilbird) have been allied to owls (and many other groups of birds (bird)) in the past, but that relationship requires modern resolution.

Classification

Distinguishing taxonomic characters

      The most important characters used to define the caprimulgiform birds are the type of plumage, structure of the feet and legs, form and structure of the bill, palatal structure, arrangement of the pelvic muscles and flexor tendons, shape of dorsal vertebrae, meristic characters such as the number of primary and secondary feathers and rectrices, presence or absence of intestinal ceca and of an oil gland, location of syrinx, presence and type of rictal bristles, powder down patches, carotid artery relationships, and a few other anatomical details. Variations in some of these same features characterize the different subgroups.

Annotated classification

Order Caprimulgiformes

 Soft plumaged (except Steatornithes), cryptically patterned birds with relatively weak anisodactyle feet and very short tarsi; deeply cleft gape (except Steatornithes); pelvic muscle formula AXY (XY in Steatornithes); flexor tendons fused (synpelmous); 10 primaries; 11–13 secondaries; 10 rectrices; aftershaft small but present; 2 carotid arteries and oil gland present (except in Podargus and Nyctibius).

      Suborder Steatornithes

 Dorsal vertebrae opisthocoelous (concave behind); gape not exceptionally deeply cleft; rostrum movably articulated with skull; plumage firm.

      Family Steatornithidae (oilbirds (oilbird))

 Locally distributed in Guyana, Venezuela, Colombia, Ecuador, Peru, and Trinidad. Desmognathous palate (maxillopalatine bones fused), palatines narrow and not expanded posteriorly; large, strong bill with hard rhamphoteca (horny covering) and subterminal tooth, surrounded by long vibrissae; bronchial syrinx; 15 cervical vertebrae. 1 species, length 40 cm (about 16 inches).

      Suborder Caprimulgi

 Dorsal vertebrae heterocoelous (saddle-shaped); deeply cleft gape; rostrum fixed; plumage soft.

      Family Podargidae (frogmouths (frogmouth))

 Confined to Australasian (except New Zealand) and southern Oriental regions (including extreme southern India). Desmognathous palate; palatines broad throughout, slightly expanded posteriorly; wide, strong bill with hard rhamphotheca; bronchial syrinx; 13 cervical vertebrae; well-developed powder down tufts on either side of rump; oil gland absent in Podargus, very small in Batrachostomus; 1 carotid artery (left). 3 genera, 15 species, length 20–40 cm (about 8–16 inches).

      Family Aegothelidae (owlet-frogmouths (owlet frogmouth))

 Australasian region (except New Zealand). Desmognathous palate; bill similar to Podargidae but shorter, weaker, and largely hidden by forehead feathering; bronchial syrinx; unique in the order in lacking ceca. 1 genus, about 8 species; length 16–25 cm (about 6–10 inches).

      Family Nyctibiidae (potoos (potoo))

 Neotropical. Schizognathous palate (small vomers and separate maxillopalatines); palatines narrow anteriorly, greatly expanded posteriorly; small, weak bill but with prominent horny angular projection on maxillary tomium midway between tip and rictus, and tomium very broad, horny, and strongly convex from that point to rictus; extremely short tarsi; toes unusually wide basally, forming a broad flattened sole; tracheobronchial syrinx; 14 cervical vertebrae; large powder down patches on sides and breast; no oil gland; 1 carotid artery (left). One genus, 7 species, length 20–55 cm (about 8–22 inches).

      Family Caprimulgidae (nighthawks (nighthawk), nightjars (nightjar))

 Worldwide in tropical and temperate zones; absent from northernmost Eurasia and America, southernmost South America, New Zealand, and some oceanic islands; subfamily Chordeilinae (nighthawks) restricted to New World. Schizognathous palate (except Chordeiles and perhaps other Chordeilinae); palatines narrow anteriorly, greatly expanded posteriorly; small, weak bill; well-developed rictal bristles (except Eurostopodus and Chordeilinae); small, weak feet; lateral toes much shorter than middle toe, 4th toe with only 4 phalanges (segments; 5 is normal for this order), middle claw with inner edge pectinated; hallux very short and directed inward; tracheobronchial syrinx; 14 cervical vertebrae. 15 genera, 90 species; 14–40 cm (about 5.5–16 inches) long without ornamental feathers.

Critical appraisal

      The major divisions of the order Caprimulgiformes are clearly defined. Their composition is remarkably homogeneous, so that there are no apparent taxonomic problems above the level of genus. The rather aberrant oilbird has traits similar to owls (owl), but evidence places it in this order. Within the order the main taxonomic problems relate to reevaluation at the generic level and clarification of species limits. These problems are likely to be settled only with the aid of field studies utilizing bioacoustical methods along with other disciplines.

Paul A. Schwartz

Additional Reading

F. Haverschmidt, Birds of Surinam (1968), includes much information on the South American caprimulgiforms. J.P. Chapin, The Birds of the Belgian Congo, vol. 2 (1939), contains readable accounts of the natural history of several tropical nightjars. D.W. Snow, “The Natural History of the Oilbird, Steatornis caripensis, in Trinidad, W.I.,” Zoologica, 46:27–48 (1961) and 47:199–222 (1962), provides an excellent and readable account of the natural history of that species. The information on nightjar hibernation is scattered among a number of short journal articles, but the reader will find interesting summaries in E.C. Jaeger, “Further Observations on the Hibernation of the Poorwill,” Condor, 51(3):105–109 (1949); and J.T. Marshall, Jr., “Hibernation in Captive Goatsuckers,” Condor, 57(3):129–134 (1955). A review of the later work in this field is presented by G.T. Austin and W.G. Bradley, “Additional Responses of the Poorwill to Low Temperatures,” Auk, 86(4):717–725 (1969).Paul A. Schwartz Ed.

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