Rails, coots, and moorhens

(Rallidae)

Class Aves

Order Gruiformes

Suborder Grues

Family Rallidae

Thumbnail description
Small to medium-sized birds with short and deep to long and slender bill, moderately long neck, broad wings, short, soft tail, and strong legs

Size
4.7–24.8 in (12–63 cm); 0.7 oz–9.13 lb (20.41 g–4.14 kg)

Number of genera, species
33 genera; 134 species

Habitat
Wetlands, grasslands, forest, and dense scrub

Conservation status
Extinct in the Wild: 1 species; Critically Endangered: 4 species; Endangered: 12 species; Vulnerable: 16 species; Near Threatened: 9 species; Data Deficient: 4 species

Distribution
Worldwide, except for polar regions and waterless deserts; widely distributed on oceanic islands

Evolution and systematics

The Rallidae is by far the largest family in the Gruiformes. In some classifications the family has been assigned to its own order Ralliformes, while others have allied with to the Charadriiformes; DNA evidence suggests that it shares a common ancestor with both gruiform and charadriiform birds. Skeletal morphology suggests a close alliance with the Psophiidae (trumpeters) and the Heliornithidae (sungrebes and finfoots), and a phylogenetic study of the Gruiformes using morphological characters, published by Bradley Livezey in 1998, places the families Psophiidae, Aramidae (limpkin), Gruidae (cranes), Heliornithidae and Rallidae together in the suborder Grues. On the basis of DNA evidence that the Rallidae may have had a distinct lineage for a long time, it has also been proposed that the rails should be elevated to their own suborder, the Ralli, alongside the Grues.

Fossil evidence tells us little about the origins of the Rallidae. The earliest rail fossils are from the Lower Eocene, about 50 million years ago, but the family may have existed earlier than this. DNA-DNA hybridization studies suggest that rails may have diverged from the other gruiform groups as long as 86 million years ago, in the Upper Middle Cretaceous. The first adequate diagnostic material on fossil rail genera comes from the Upper Oligocene and Lower Miocene, 20–30 million years ago, and by then the birds had attained a morphology similar to that of modern rails. Most continental fossil rails from Pliocene and Pleistocene deposits have been assigned to modern genera.

The geographic origins of modern rails have been obscured by the antiquity, cosmopolitan distribution, and inadequate taxonomy of the family. However, Storrs Olson has shown that the greatest number of rail species and peculiar genera, and the most primitive species, are found in the Old World tropics. The New World has fewer groups, most of which are apparently derived from Old World stem groups. A few genera appear to have specialized and radiated in the New World, some of which (e.g. Rallus and Fulica) have reinvaded the Old World.

Several classifications have been proposed for the family. The generally accepted classification, published in 1973 by Olson, listed 35 genera in two subfamilies, the Himantornithinae and Rallinae, the former containing one species, Himantornis haematopus, the nkulengu rail of Africa. Livezey's classification also recognizes these subfamilies. Sibley and Ahlquist have suggested that the flufftails (Sarothrura) of Africa and Madagascar diverged from the rest of the group about 60 million years ago and should be separated into a family Sarothruridae within its own superfamily, the Sarothruroidea.

In this work we follow Olson and Livezey's subfamily treatment and Olson's taxonomic treatment regarding genera—modified to some extent by subsequent studies. Within the family, 33 genera containing 134 extant species and 312 subspecies are recognized, following the list published in 1998 in the book Rails: a guide to the rails, crakes, gallinules and coots of the world by Barry Taylor.

Two "natural groups" within the Rallidae are usually recognized: the crakes, rails, and wood-rails, most of which are terrestrial; and the gallinules (including moorhens) and coots, which are more aquatic. The term "rail" is applied to the whole family and also to longer-billed species in many genera, while "crake" is applied mostly to the smaller, short-billed, species, particularly in the genera Laterallus and Porzana. "Gallinule" can cover all the birds in the second group except the coots, though it is often restricted to Gallinula and Porphyrio.

Physical characteristics

The rails are a relatively homogeneous group of birds, ranging in size from the tiny black rail (Laterallus jamaicensis), 4.7 in (12 cm) long and weighing 0.7 oz (20 g), to the flightless takahe (Porphyrio mantelli) 24.8 in (63 cm) long and weighing up to 9.2 lb (4.2 kg). The sexes are usually similar in size but in a few species the male is markedly larger than the female.

Rail plumage is often cryptic, common colors being somber browns, chestnut, black, blue-gray, or gray, but the Porphyrio gallinules are predominantly iridescent purple, blue, or green. The upperparts are often spotted, barred, or streaked and the flanks strongly barred, while the undertailcoverts may contrast strongly with the rest of the plumage. In most species the sexes are very similar in appearance, exceptions including the flufftails and the New Guinea Rallina forest-rails. Only the watercock (Gallicrex cinerea) shows any significant seasonal change in plumage color or pattern. The juvenal plumage is often a duller, less patterned version of the adult plumage.

The downy plumage of most species is black or dark brown, but the nkulengu rail chick is cryptically patterned with brown, black, and white, closely resembling precocial chicks of other orders. Some chicks, especially those of coots (Fulica), have distinctively colored filoplumes or bristles. Coots, gallinules (Porphyrio), and moorhens (Gallinula) have brightly colored bare skin on the head, and such prominent features act as signals for feeding.

The bodies of rails are often laterally compressed, allowing easy movement through dense vegetation, and the neck can be quite long. The wings are short, broad, and rounded. In some species the alula has a sharp claw, used by the young, and possibly also by adults, when climbing. Short flights are usually low and weak, but some species migrate or disperse over long distances. The tail is short and soft.

The bill shape is variable: from long and thin to short and fine, straight or slightly down-curved; or short and laterally compressed; or short and heavy. It is often brightly colored. Gallinules and coots have a frontal shield that may be of a contrasting color to the bill. Shield and bill colors often become duller in the nonbreeding season, when the shield may also shrink.

Rails have strong, often long, legs. The toes are often long, for walking on marsh vegetation. In some species the legs and feet are brightly colored. Coots have the pelvis and legs modified for diving, and lateral lobes on the toes aid swimming.

Flightlessness is a well-known feature of the family and all flightless rails occur on islands. Of the 134 extant rail species, 41 are known only from islands, including New Guinea, New Zealand, and Madagascar, and 24 (59%) of these are flightless. Flightlessness has evolved many times within the family, often and repeatedly on predator-free islands. The energetic cost of flight is high and flight muscles and associated structures average 20–25% of body weight in typical birds. Where such costs do not convey the benefits of dispersal and escape from predators it is obviously advantageous to become flightless. The muscles and bones of the wing and pectoral girdle are greatly reduced, the feathers become loosely constructed and the leg muscles usually become better developed.

The strong tendency of rails to become flightless suggests a predisposition to the condition, and rails are pre-adapted to coping with some of its restrictions. Thus many volant species are behaviorally flightless, avoiding predators by running. Many are temporarily flightless during wing molt, and the postnatal development of flight in most species is slow.

Distribution

Rails occur throughout the world, being absent only from polar regions, waterless deserts, and mountains above the snow line. Of the 33 rail genera, four (Porzana, Porphyrio, Gallinula, and Fulica) occur worldwide. The genus Porzana includes 13 species of small rails, one of which, Baillon's crake (Porzana pusilla), ranges from western Europe through Africa and Asia east to Japan and Australasia.

The genus Porphyrio has five species of medium to very large gallinules, including the purple swamphen (Porphyrio porphyrio), which has a similar range to Baillon's crake. In the genus Gallinula, the common moorhen (Gallinula chloropus) occurs from the Americas east through Africa and Eurasia to the Malay Archipelago; farther east it is replaced by the very similar dusky moorhen (Gallinula tenebrosa). Although moorhens are commonly regarded as wetland birds, two species occur in forest: the Samoan moorhen (Gallinula pacifica) and the San Cristobal moorhen (Gallinula silvestris). The center of the coots' species diversity is in South America, where eight of the 11 species occur.

The relatively unspecialized genus Gallirallus, with 10 extant and five recently extinct species, is distributed largely from Indonesia through Australasia to western Pacific islands. Seven extant species are flightless, as were all the recently extinct species. The islands in this region also support 17 other endemic rails, eight flightless, and seven of these in five endemic genera. The genus Gymnocrex contains three species of very distinctive, long-legged forest rails of Indonesia and New Guinea, one of which, the Talaud rail (Gymnocrex talaudensis) was discovered only in 1996.

Africa and Madagascar have 23 endemic rail species, with 15 species in six endemic genera, including Sarothrura (the flufftails). In comparison, 51 rail species occur only in the Americas, including 27 in eight endemic genera, the most diverse of which is Laterallus, with nine species. South and Central America are home to all seven species of the wood-rails (Aramides). Six of the nine Rallus species occur only in the Americas, including the well-known clapper and king rails (Rallus longirostris and R. elegans).

Most Holarctic rails are migratory, and five of the nine species that breed in the western Palaearctic winter in sub-Saharan Africa. All rails that breed in North America are migratory to some extent. Relatively little is known about the migrations of species that breed from India east to Japan and south through the Oriental region, but evidence suggests that many birds that breed in the northern regions of Asia move south after breeding. Even less is known about the movements of South American rails, but most of the species known or suspected to be migratory or dispersive inhabit wetlands or wet grassland. All rails occurring widely in Australia are migratory, dispersive, nomadic, or irruptive. In Africa, some species move away from the equator to breed during the rains.

The widespread occurrence of rails on oceanic islands reflects these birds' powers of dispersal and their tendency to vagrancy. The high degree of long-distance vagrancy in the family is also indicative of the readiness with which rails are blown off course by unfavorable winds as a result of their relatively

poor flight performance. The purple gallinule (Porphyrio martinica) is a vagrant to South Africa, but there are no instances of African rails occurring in the Americas.

Habitat

The cosmopolitan distribution of the family reflects the ability of rails to adapt to a great diversity of habitat types, both natural and artificial, including wetland, grassland, savanna, scrub, and forest.

Although the greatest number of species occurs in wetland habitats, many species occupy forest habitats in tropical regions. The most primitive living rail, the nkulengu rail, is a forest bird, as are the members of other primitive or unspecialized genera such as Aramides, Canirallus, and Gymnocrex. Species in the most specialized or derived genera such as Rallus, Porzana, Gallinula, and Fulica are aquatic or marsh-dwelling. This suggests that forest was the primitive habitat of the family.

Rails occupy all types of vegetated wetlands, plus some open water habitats. Freshwater wetland habitats include swamps, bogs, marshes, floodplains, pans, ponds, ditches, rice fields, and vegetation fringing streams, rivers, canals, and lakes. Some species, such as the white-browed crake (Porzana cinerea) of Asia and Australasia, and Porphyrio species, prefer floating vegetation, on which they search for food. Some rails occur at coastal wetlands such as lagoons, saltmarshes, tidal creeks, and mudflats, while mangroves are an important habitat for species such as some South American Aramides wood-rails. Coots, the most aquatic rails, occupy fresh to saline waterbodies.

Forest habitats range from low to high altitudes and include primary and secondary growth, riverine and swamp forest, overgrown and abandoned cultivation at forest margins, banana groves, cassava plantations, and dense evergreen or deciduous thickets. Substrates may be clear, with leaf-litter, soft earth or mud, or may have dense herbaceous vegetation. Some species, such as the gray-throated rail (Canirallus oculeus) of Africa and Woodford's rail (Nesoclopeus woodfordi) of the

southwest Pacific, occur at forest streams, swamps, or muddy patches, but others are not associated with wet areas.

A few species occur in dense grassland habitats, which may be wet to predominantly dry, the latter including savanna, pampas, meadows, and crop fields. Grassland habitats may be permanently or seasonally occupied; for example, the African crake (Crex egregia) occurs in seasonally moist to wet grassland, which is frequently burned during the dry season, forcing the birds to emigrate after breeding.

Most rails do not have specialized diets and this enables many to exploit ephemeral or atypical habitats. They are able to colonize islands where, in the absence of competitors and predators, they can radiate to occupy almost any available terrestrial niche. They are capable of adapting to harsh conditions on remote oceanic islands. The recently extinct Ascension rail (Atlantisia elpenor) lived on Ascension Island, where the terrestrial environment consists of bare, waterless tracts of lava and ash. It apparently obtained its food and water from the eggs and regurgitated prey of the seabirds, which formerly nested on Ascension in great numbers. The spotless crake (Porzana tabuensis) of the Pacific region, has successfully colonized islands. It normally occurs in a great variety of wetland and scrub habitats, but on some islands it occupies dry rocky habitats with no water.

For rails that feed chiefly on invertebrates, the structure of the vegetation and the nature of the substrate are the most important factors influencing habitat choice. The Virginia rail (Rallus limicola) avoids marshes with high stem densities or large amounts of residual vegetation—features that impede movement—whereas vegetation height is not important if adequate overhead cover is present. It needs shallow water and a substrate with a high invertebrate abundance, and is most common in wetlands with 40–70% upright emergent vegetation interspersed with open water, mudflats, or matted vegetation. The buff-spotted flufftail (Sarothrura elegans) occupies natural forests and thickets, but also areas dominated by alien vegetation. Its invertebrate food is equally abundant on substrates below exotic vegetation and those below indigenous plants.

Behavior

Most rails are solitary or occur in pairs, family parties, or small groups. The most gregarious species are the coots, most of which associate in large monospecific flocks outside the breeding season. Some gallinules and moorhens also associate in loose flocks when not breeding. The black-tailed native-hen (Gallinula ventralis) of Australia may occur in flocks of up to 20,000 birds during its periodic irruptions.

Wetlands and lush grasslands are structurally simple and may be highly productive, with food concentrated in a narrow spatial range. In such conditions it may be possible for males to control territories in which two or more females can breed, relegating less successful males to suboptimal territories, or to none at all. This strategy may apply to the yellow rail and the corncrake, while polyandry in the striped crake may have evolved in response to great variability in breeding conditions and the availability of abundant food in the breeding habitat.

In the promiscuous Porphyrio and Gallinula species, social structure and mating systems are complex. The common moorhen is normally monogamous, but immatures from earlier broods, and sometimes other mature birds, often help care for chicks. Polyandrous trios and cooperative nesting also occur, while intraspecific brood parasitism is regular. Monogamy prevails in most races of the purple swamphen, but in two races birds often live in communal groups. In New Zealand, stable groups, usually of kin, hold permanent territories and are polygamous, usually with two to seven breeding males, one to two breeding females, and one to seven nonbreeding helpers (offspring from previous matings). Unstable groups are usually non-kin and are promiscuous, with much aggression and many male members, and are largely unsuccessful. Within a stable group mate-sharing is total and multiple paternity prevails. Dominant females lay in a common nest and all group members care for the young. Habitat saturation and a shortage of prime breeding territories appear to be responsible for this breeding strategy.

Agonistic behavior in rails is common and often conspicuous. In the purple swamphen, the position of the tail and wings is important in agonistic display, while differing body postures indicate aggression or anxiety. Moorhens and coots share similar agonistic displays, in which the degree of prominence of the frontal shield is often an important component.

Rails are territorial, many species defending territories only while breeding. Winter feeding territories are maintained by the water rail (Rallus aquaticus), the spotted crake (Porzana porzana), and the African crake, and this phenomenon is probably more widespread than is known.

Most rails are very vocal, with an extensive repertoire, as is to be expected in birds that inhabit dense cover where visual contact is often very limited and communication by sound is important. Calls include screams, squeals, trills, whistles, whines, hoots, moans, booms, rattles, clicking and ticking notes, snoring noises, humming and buzzing sounds, trumpets, roars, grunts, barks, frog-like croaks, and snake-like hisses; calls of some small species may be very insect-like. The advertising and territorial calls of many species are given in a repetitive series, are often loud, and are given most commonly in the early morning, the evening, and at night.

Most rails normally keep within dense cover and are adept at moving around without causing any noise or disturbance of the vegetation. When alarmed, most run rather than fly, and they can melt quietly and rapidly into cover, compressing the body laterally for easy passage through vegetation. Rails often walk with bobbing head and flicking tail. Tail jerking is used in visual orientation and signaling between con-specific individuals, but in the common moorhen and the purple swamphen it is also directed toward potential predators as an alertness signal and pursuit deterrent.

Many rails are predominantly crepuscular. Some terrestrial and marsh species forage at night as well as by day and it is possible that nocturnal activity is largely confined to species of open habitats where visibility at night is relatively good. Rails that forage in tidal areas are often active during low tides at night. Most rails normally roost singly, in pairs, or in family groups, generally on the ground in dense cover, but sometimes above ground in dense vegetation such as bushes and trees.

Feeding ecology and diet

Rails are characteristically omnivorous, generalized feeders, often opportunistic and able to take advantage of new food sources. In general the most aquatic species, such as the gallinules and coots, are largely herbivorous, while those that inhabit terrestrial and marshy habitats are either omnivorous or take predominantly animal food, at least in the breeding season.

Many rails appear to feed largely on the most abundant foods available at any time. A few species take only a restricted range of prey or plant material, and such specialization is usually a reflection of the limited variety of suitable food available. Thus rails that forage in mangroves feed largely on crabs, examples being the chestnut rail (Eulabeornis castaneoventris) and the rufous-necked wood-rail (Aramides axillaris).

Invertebrates are the principal diet of many species, and commonly include worms, mollusks, crustaceans, spiders, and insects. Some rails take vertebrate prey, including small fish, amphibians and their tadpoles, small reptiles (lizards, snakes, and turtle eggs and hatchlings), and the eggs and young of other birds, while a few will eat carrion. Forest-dwelling

rails probably eat fewer plant foods than those in other habitats.

Many types of plant foods are eaten by rails, including seeds, fruits, shoots, stems and leaves, tubers, bulbs, rhizomes and roots, as well as marine and filamentous algae, fungi, lichens, and ferns. Cultivated plants such as vegetables, cereal and fodder crops, fruit, and taro are taken by some species. Coots are almost entirely herbivorous, but some aquatic insects, mollusks, and crustaceans are taken, and coots will sometimes eat eggs, fish, carrion, duck-food pellets, and even food scraps from campsites.

Although most rails drink fresh water, some species are able to survive on oceanic islands with no fresh water. These rails may drink salt water or may obtain most of their water from their food. Thus the white-throated rail (Dryolimnas cuvieri) drinks salt water on Aldabra, and the buff-banded rail (Gallirallus philippensis) can exist on islands with no fresh water. Some rails, such as the saltmarsh-dwelling clapper rail, possess well-developed supra-orbital (nasal) glands, which function in the excretion of salt.

Bill size and shape provide a good indication of a rail's foraging habits. Species with long thin bills probe for invertebrate food in shallow water, soft ground, and litter, while those with small, fine bills take small invertebrates and seeds from the substrate, shallow water, and low vegetation. Those

with straight bills of moderate length and depth take a wide variety of small to large food items, chiefly by probing, gleaning, digging, sifting leaf-litter, stabbing at large prey, and raking in earth and mud. Thick-billed species tear and slice vegetation, and dig or pull up the underground parts of plants.

Some gallinules use the foot to grasp and manipulate food. The purple swamphen uses its bill to pull out emergent plants and then grasps them in the foot while eating the bases.

Only coots regularly dive for food, but coots and Gallinula species regularly up-end when feeding. The white-browed crake often feeds while swimming, floating with the neck extended parallel with the surface and reaching out to capture insects.

Some species show seasonal variations in the proportions of animal and plant food taken, and this may reflect seasonal changes in the availability of food, the use of different habitats when birds are on migration or in wintering areas, or the need for a greater consumption of protein in the breeding season to satisfy the requirements for egg-laying. Many species increase their intake of animal food in the spring and summer, and of plant food in the autumn and winter. Most rail chicks, even those of herbivorous species, are fed primarily on animal food.

Reproductive biology

With the exception of some coots, moorhens, and gallinules, little is known about the breeding of most rails. The nest, eggs, and young of 23 species remained undescribed in 2001.

Monogamy is the predominant mating system in the family. This is to be expected because, although rail chicks are precocial or semi-precocial, they need intensive care at an early age, when they are fed, guarded and brooded by their parents. A non-monogamous mating system occurs in the wild in only five rails: the corncrake (Crex crex), the purple swamphen, the common moorhen, the dusky moorhen and the Tasmanian native hen (Gallinula mortierii), and in captivity in the yellow rail (Coturnicops noveboracensis) and the striped crake (Aenigmatolimnas marginalis).

Most species appear to breed seasonally, during the spring and summer in temperate regions and during the wet seasons in the tropics. Most exceptions to this pattern involve tropical or subtropical species that may have extended or ill-defined breeding periods.

Some species may breed throughout the year if conditions remain suitable, examples from Africa being the common moorhen and the red-knobbed coot (Fulica cristata). Studies of some rails in southwest Australia have shown that in most species the laying period is correlated with peak rainfall, day length, and temperature. The timing of vegetation development is often important to the initiation of nesting in rails of marshy habitats. Thus in the state of Ohio, peak nest initiation of the common moorhen occurs when vegetation height is 18–40 in (45–100 cm) and its growth rate is greatest. The breeding season of the Tasmanian native-hen is determined by rainfall, as it depends on fresh young plant growth.

Courtship feeding and allopreening are common, and aggressive-looking courtship chases often lead to copulation. In some species the male's courtship display involves bowing, and may involve the display of bold flank patterns or contrastingly colored undertail-coverts. Gallinules, moorhens, and coots show the most complex courtship and mating behavior. In the purple swamphen, courtship usually starts with allopreening but the male may also present aquatic plant material in his bill to the female, and the female solicits copulation by adopting the arch-bow posture. Coots and moorhens share similar components in their sexual displays, including a bowing-and-nibbling ceremony (in which one bird is submissive while the other preens it), a greeting and passing ceremony, and a courtship chase.

Nests are usually concealed in thick ground vegetation, often near or over water, but some species nest in dry areas and some in trees. Nest materials are often gleaned from the vegetation closest to the nest site, and nests are often built by both sexes. The nest is usually cup-shaped, but is domed in some species, including most Laterallus crakes. Nests in grass and emergent vegetation often have surrounding vegetation woven into a concealing canopy over the bowl, while nests in wetlands often have ramps up to the bowl. Some species build nests that float or are attached to aquatic vegetation. The giant coot (Fulica gigantea) and horned coot (Fulica cornuta) build enormous permanent nests of aquatic vegetation which, in the horned coot, is usually placed on a conical mound of stones, up to 13 ft (4 m) in diameter at its base, about 2 ft (60 cm) high and about 3 ft (1 m) in diameter at the top. The structure may weigh about 1.5 tons; each stone weighs up to 1 lb (450 g) and both adults collect stones and carry them to the nest in the bill.

Rail eggs are usually approximately oval, smooth, and fairly glossy. The ground color is white to dark tan, usually blotched or spotted with red-brown, gray, mauve, or black. Clutch size

varies from one to 19 (most frequently five to 10), and dumping or laying by more than one female in the same nest may occur. Incubation is by both sexes in most species, and incubation periods are 13–31 (usually 15–19) days per egg. Hatching may be synchronous or asynchronous.

Rail chicks hatch covered in down and are precocial or semi-precocial, usually leaving the nest after one to three days. Chicks are usually fed bill-to-bill at first and are normally tended by both parents, in some species also by helpers. The fledging period is four to eight weeks but in the giant coot it is about four months. Chicks' legs and feet grow rapidly, reaching full size before the rest of the body, but the growth of the wings is generally much retarded. The young usually become independent as soon as they are fully fledged. First breeding usually occurs when the birds reach their first year.

One or two broods are usually reared, and many species will re-lay several times after failure. Nesting success varies widely in the few species for which figures are available, often depending on factors such as food supply, predation, and flooding. In general, early nests are often more successful than later ones. In North American species, nesting success is given as: 10–100% for the clapper rail, 81% for the king rail, 53% for the Virginia rail, 49–91% for the purple gallinule and over 80% for the American coot (Fulica americana). In many species hatching success is often high, whereas chick survival may be much lower.

Conservation status

The IUCN Red List of birds, published in 2000, shows that, of the 134 living rail species, 33 (almost 25%) are Threatened, including four that are Critically Endangered, 12 Endangered, 16 Vulnerable and one, the Guam rail (Gallirallus owstoni) Extinct in the Wild. A further nine species are listed as Near Threatened and four as Data Deficient. Thus the survival of 62 species (46%) of rails gives cause for concern.

Of 20 rail taxa (16 species and four subspecies) that have become extinct since 1600, 17 (85%) were flightless. The extinction of these rails provides a classic example of the particular vulnerability of island endemics. The principal causes of extinctions among island rails have been introduced mammalian predators such as cats, dogs, rats, mongooses, and pigs, indiscriminate hunting by the first people to visit the islands, and habitat destruction by introduced goats, rabbits, and fire. Introduced predators have probably been responsible for more extinctions than any other cause. Several extant island species are still at risk from the possible accidental introduction of mammalian predators to their islands.

Habitat destruction does not seem to be a critical problem for any threatened island rail at present, but is certainly a major threat to many continental species. For example, the plain-flanked rail (Rallus wetmorei) has a very restricted distribution in coastal Venezuela, where its mangrove and lagoon habitats are being destroyed by housing development, oil exploration, and diking.

Many rails are probably undergoing a continual population decline, largely through habitat loss. The wholesale and enormous destruction of indigenous forests is a severe threat to some species, especially in southeast Asia and South America. Palustrine wetlands are under threat worldwide and are disappearing at an alarming rate. Small crake species, such as the black rail, which inhabit the edges of marshes, are generally more threatened by habitat destruction than are other rails, which live in the interiors of marshes or alongside open water.

Great efforts have been made to save some threatened species, involving captive breeding and reintroduction of birds into the wild, habitat management, and predator control. The takahe and the Guam rail are two good examples.

Significance to humans

Rails have had little association with humans and have no significant place in art, literature, or legend. This is presumably because most rails are unobtrusive, cryptic, and hard to see; many people are not even aware that they exist.

However, one rail did excite the interest of ancient civilizations. The purple swamphen is depicted climbing on papyrus stems in the Egyptian wall paintings at Medum. The Greeks and Romans refrained from eating the "Porphyriõn" but imported the birds and placed them in palaces and temples, where they walked around freely as worthy guests by virtue of the nobleness of their bearing, the graciousness of their nature, and the beauty of their plumage.

Local superstitions about rails include those held by some African peoples to explain the strange calls heard from forest or marsh. The song of the buff-spotted flufftail, one of the most evocative sounds of the African rainforest, is sometimes believed to be the wail of a banshee, or the sound of a chameleon mourning for its mother, whom it killed in an argument over some mushrooms. The extinct Kosrae crake (Porzana monasa) of the Caroline Islands remains a legend among the islanders, since it was regarded as a sacred bird before the arrival of Christian missionaries.

In the Cocos-Keeling Islands, the buff-banded rail (Gallirallus philippensis) is apparently used to hatch chicken eggs in place of domestic hens. In Bangladesh the watercock (Gallicrex cinerea) is used as a fighting bird, as in cockfighting. In South America the giant wood-rail (Aramides ypecaha) is often kept in captivity and individuals are sold in village shops.

Rails have long been hunted for food and sport in many parts of the world and the Eurasian coot (Fulica atra) is still shot in Mediterranean countries for these purposes. In Europe the corncrake (Crex crex) was commonly hunted for food in the past, and is still caught in Egypt during the ancient practice of quail netting. In the United States the larger rails may still be hunted legally and in Audubon's time soras (Porzana carolina) and clapper rails (Rallus longirostris) were heavily hunted. People in Africa, Asia, and South America often trap forest rails, while marsh rails are widely hunted in Asia. Rail eggs are regarded as highly palatable, and common moorhens were formerly extensively exploited for their eggs in Asia. The giant coot is also exploited for its eggs.

The larger rails may occasionally damage crops or pasture. The Tasmanian native-hen has been falsely accused of this, and was at one stage declared vermin. The purple swamphen is said to do considerable damage to growing rice crops in India and Bangladesh, but such damage must be highly localized. The purple gallinule is regarded as a pest in rice fields in some Neotropical areas.

Species accounts

List of Species

Buff-spotted flufftail

Sarothrura elegans

subfamily

Rallinae

taxonomy

Gallinula elegans A. Smith, 1839, Durban, South Africa. Two subspecies.

other common names

French: Râle ponctué; German: Tropfenralle; Spanish: Polluela Elegante.

physical characteristics

6–6.7 in (15–17 cm); 1.4–2 oz (39–61 g). Male has orange-chestnut foreparts and buff-spotted upperparts; female golden brown with buff-spotted upperparts and barred underparts. Juvenile gray-brown.

distribution

S. e. reichenovi: Guinea east to Democratic Republic of Congo (Zaire) and Uganda, south to north Angola; S. e. elegans: southern Sudan and Ethiopia south to South Africa.

habitat

Forest, thickets, and abandoned cultivated areas.

behavior

Territorial when breeding. Diurnal, but breeding males sing mostly at night, giving a loud, repeated, hollow hoot "oooooo," sometimes for 12 hours or more. Some populations sedentary, others have seasonal movements.

feeding ecology and diet

Takes terrestrial invertebrates.

reproductive biology

Monogamous. Breeds during rains. Lays three to five white eggs in domed nest of dead leaves or grass on ground. Incubation 15–16 days; young independent at 19–21 days.

conservation status

Not threatened. Widespread, locally common. Probably holds its own because it colonizes degraded forest habitats and exotic vegetation.

significance to humans

Its hooting vocalization has given rise to many local legends.

Forbes's forest-rail

Rallina forbesi

subfamily

Rallinae

taxonomy

Rallicula forbesi Sharpe, 1887, Owen Stanley Range, New Guinea. Four subspecies recognized.

other common names

English: Forbes's chestnut rail; French: Râle de Forbes; German: Nymphenralle; Spanish: Polluela de Forbes.

physical characteristics

8–10 in (20–25 cm); 3–3.2 oz (87–91 g). Foreparts chestnut; rear upperparts and wings blackish-brown, spotted buff in female; rear underparts barred. Juvenile duller and browner.

distribution

R. f. steini: central New Guinea; R. f. parva: northeastern New Guinea (Adelbert range); R. f. dryas: northeastern New Guinea (Huon Peninsula); R. f. forbesi: southeastern New Guinea.

habitat

Montane forest.

behavior

Poorly known and secretive

feeding ecology and diet

Invertebrates, small vertebrates, and seeds.

reproductive biology

Roosting nest a football-sized domed structure of leaf skeletons and moss on the ground. One breeding nest was a platform in a tree. Eggs probably four to five, white.

conservation status

Not threatened. Not uncommon locally in the east, probably scarce or rare in west.

significance to humans

Regularly hunted for food.

Black rail

Laterallus jamaicensis

subfamily

Rallinae

taxonomy

Rallus jamaicensis Gmelin, 1789, Jamaica. Four subspecies recognized.

other common names

French: Râle noir; German: Schieferralle; Spanish: Polluela Negruzca.

physical characteristics

4.7–6 in (12–15 cm); 0.7–1.6 oz (20.5–46 g). Small and dark, nape to mantle orangy- to reddish brown, upperparts and rear underparts barred or spotted white. Undertail-coverts cinnamon in two races. Female paler on foreparts; juvenile browner, plainer. Hatchlings covered with black down.

distribution

L. j. coturniculus: California; L. j. jamaicensis: eastern United States and eastern Central America, winters from coastal southern and eastern United States to Guatemala and Greater Antilles; L. j. murivagans: coastal central Peru; L. j. tuerosi: lower Junin, Peruvian Andes; L. j. salinasi: southern Peru, central Chile and western Argentina

habitat

Marshes and wet grassland.

behavior

Territorial when breeding. Some populations migratory, others sedentary. Male's breeding "kic-kic-kerr" call distinctive.

feeding ecology and diet

Eats mainly small invertebrates; also fish, tadpoles, and seeds.

reproductive biology

Monogamous; occasional polygyny possible. Breeds in summer in United States, during rains in South America. Nest a bowl of grasses or rushes with a woven canopy, low in marsh vegetation. Eggs two to 13; color is buffy to pinkish-white, with brown speckling concentrated at larger end. Incubation 17–20 days.

conservation status

L. j. tuerosi is Endangered and is known from only two sites at lower Junin, where it is at risk from pollution and water level fluctuations. Other races are Lower Risk/Near Threatened. Most United States populations declined drastically in twentieth century.

significance to humans

None known.

Guam rail

Gallirallus owstoni

subfamily

Rallinae

taxonomy

Hypotaenidia owstoni Rothschild, 1895, Guam. Monotypic.

other common names

French: Râle de Guam; German: Guamralle; Spanish: Rascón de Guam.

physical characteristics

11 in (28 cm); 6–10.7 oz (170–303 g). Nearly flightless. Upperparts olive-brown; foreneck to breast gray; underparts and remiges barred black and white. Juvenile has less gray; chick covered with black down.

distribution

Formerly on Guam, Mariana Island. Reintroduced to Rota, northern Mariana Island, and Guam.

habitat

Forest, woodland, scrub, grassland, and agriculture.

behavior

Territorial; secretive and wary.

feeding ecology and diet

Opportunistic and omnivorous, taking mollusks, insects, geckos, seeds, fish, and carrion. Often forages at edges of fields and roads.

reproductive biology

Monogamous. Breeds all year. Nest a cup of grass, on dry ground in dense grass; eggs one to four (usually three to four); white to pinkish in color with small spots of pink or blue concentrated at the large end. Incubation 19 days; young sexually mature at 16 weeks.

conservation status

Formerly abundant throughout Guam, despite being hunted; 1960s population estimated at 80,000. After 1968 it declined rapidly due to predation by the accidentally introduced brown tree snake (Boiga irregularis), and by 1987 it was Extinct in the Wild. It survives in captive-breeding facilities on Guam and at 14 zoos in the United States (about 180 birds in 1999). From 1987 birds were introduced to the snake-free island of Rota, where they bred from 1999. It was reintroduced to Guam in 1998, into a protected area.

significance to humans

None known.

White-throated rail

Dryolimnas cuvieri

subfamily

Rallinae

taxonomy

Rallus cuvieri Pucheran, 1845, Mauritius. Two subspecies recognized.

other common names

French: Râle de Cuvier; German: Cuvierralle; Spanish: Rascón de Cuvier.

physical characteristics

12–13 in (30–33 cm); 5–8 oz (138–223 g) (aldabranus), 9–9.7 oz (258–276 g) (cuvieri). Orangy- to reddish brown foreparts, prominent white chin and throat, greenish olive upperparts, barred flanks and white lateral undertail-coverts. Juvenile duller and browner.

distribution

D. c. cuvieri: Madagascar; D. c. aldabranus Aldabra Atoll.

habitat

Forest, marsh, and mangroves (cuvieri); coral scrub (aldabranus).

behavior

Permanently territorial. Often confiding. Vocal, with grunts, clicks, squeals, and a song of loud whistles; often calls at night. The Aldabra race is flightless.

feeding ecology and diet

Eats mainly invertebrates; also turtle eggs and hatchlings, and scraps at campsites (Aldabra).

reproductive biology

Monogamous; pair bond permanent. Breeds during rains in Madagascar (October through March); on Aldabra lays in Dec. Nest a bowl of leaves and grass on the ground, or a few twigs and leaves among rocks. Eggs three to six (usually three to four); young independent at 12–15 weeks.

conservation status

Not threatened. Nominate race common. In 2001 3,500–8,000 aldabranus individuals occurred naturally on three islands. Reintroduced to Picard Island in 1999, where it breeds and has a predicted population of 2,500 birds. Its survival depends on preventing the spread of feral cats and the introduction of populations to other islands to safeguard against extinction by catastrophic events.

significance to humans

None known.

Corncrake

Crex crex

subfamily

Rallinae

taxonomy

Rallus crex Linnaeus, 1758, Sweden. Monotypic.

other common names

French: Râle des genêts; German: Wachtelkönig; Spanish: Guión de Codornices.

physical characteristics

10.6–12 in (27–30 cm); 4.6–7.4 oz (129–210 g). Streaked upperparts, tawny upperwing-coverts, barred flanks and blue-gray face, foreneck, and breast. Juvenile duller, with no gray.

distribution

Breeds in Europe and central Asia, east to western China and central Siberia; winters in Africa, mainly from Democratic Republic of Congo (Zaire) and southern Tanzania south to eastern South Africa.

habitat

Breeds mainly in grass meadows; winters in grassland and savanna.

behavior

Most active at dawn and dusk. Breeding male's rasping "krekkrek" call given all night.

feeding ecology and diet

Many invertebrates; also seeds and grass blades. Normally forages within cover.

reproductive biology

Serial polygyny regular, males mating with two or more females. Breeds April and August. Nest a cup of vegetation on the ground in dense vegetation. Eggs six to 14 (usually eight to 12); incubation 16–20 days, by female only; chicks independent at 10–20 days, fledged at 34–38 days. One to two broods per season; breeding success low on agricultural land.

conservation status

In 1999 a total of 1.7–3 million singing males estimated. Western European populations declined rapidly during the twentieth century due to changing grassland management. Considered Vulnerable because of the potential for similar widespread land-use changes in its eastern European strongholds.

significance to humans

Some migrating birds are trapped for food.

Inaccessible rail

Atlantisia rogersi

subfamily

Rallinae

taxonomy

Atlantisia rogersi Lowe, 1923, Inaccessible Island, Tristan da Cunha. Monotypic.

other common names

French: Râle atlantis; German: Atlantisralle; Spanish: Rasconcillo de Tristan da Cunha.

physical characteristics

5–6 in (13–15.5.cm); 1.2–1.7 oz (34–49 g). Smallest flightless bird. Male gray-black, with dark brown back and wings; narrow white barring on upperwings and underparts. Female paler, browner; juvenile black.

distribution

Inaccessible Island.

habitat

All island vegetation types from tussock grass to boulder beaches.

behavior

Territorial, with small territories 0.025–0.1 acres (0.01–0.04 ha). Partly subterranean, using tunnels through vegetation and cavities under boulder beaches.

feeding ecology and diet

Eats invertebrates; also seeds and berries.

reproductive biology

Monogamous; pair bond permanent. Lays October through January. Nest domed, on ground in dense vegetation; of dead

grass or sedges. Eggs: two. May retain immature plumage for two years, suggesting delayed maturity. Fertility possibly low; chick mortality high.

conservation status

Abundant, with a population of 8,400–10,000 birds in 1992; possibly at carrying capacity. Vulnerable: permanently at risk from the accidental introduction of predators and other chance events.

significance to humans

None known.

Giant wood-rail

Aramides ypecaha

subfamily

Rallinae

taxonomy

Rallus ypecaha Vieillot, 1819, Paraguay. Monotypic.

other common names

French: Râle ypécaha; German: Ypecaharalle; Spanish: Cotara Ipacaá.

physical characteristics

16–19.3 in (41–49 cm); 1.2–1.9 lb (565–860 g). Olive-brown and vinous-chestnut, with gray face and foreneck, and black rear body. Juvenile paler and duller.

distribution

Eastern and southeastern Brazil, Bolivia, Paraguay, Uruguay, and northeastern Argentina.

habitat

Marshes, swamps, fields, and gallery forest.

behavior

Often bold and inquisitive. Stance upright, gait elegant. Solitary, but congregates in the evening for a communal display, rushing around with a powerful chorus of screams, shrieks, and wheezes.

feeding ecology and diet

Arthropods, mollusks, seeds, and fruit; forages in early morning and evening.

reproductive biology

Monogamous. Breeds September through February (Uruguay). Nest of grass and stems, on ground or in trees, usually near water. Eggs: four to seven. In captivity, incubation 24 days; young independent at eight to nine weeks.

conservation status

Not threatened. Formerly locally common to abundant, it may have suffered less from habitat destruction than its forest-dwelling congeners.

significance to humans

Often kept in captivity. Hunted in Argentina.

Talaud rail

Gymnocrex talaudensis

subfamily

Rallinae

taxonomy

Gymnocrex talaudensis Lambert, 1998, Karakelong I., Talaud Archipelago.

other common names

French: Râle de Talaud; German: Talaudralle; Spanish: Cotara.

physical characteristics

Approximately 13–14 in (33–35 cm). Chestnut foreparts, olive-green upperparts, tawny remiges, blackish underparts and tail, yellow bill and legs, and white facial skin. Only the holotype is described.

distribution

Karakelong Island, Talaud Archipelago, Indonesia.

habitat

Long wet grass and scrub, including at forest edges.

behavior

Extremely shy; seen only once in the four years after its discovery.

feeding ecology and diet

Snails and beetles.

reproductive biology

Not known.

conservation status

Endangered, with a very small range; faces habitat loss and degradation.

significance to humans

Trapped for food.

White-breasted waterhen

Amaurornis phoenicurus

subfamily

Rallinae

taxonomy

Gallinula phoenicurus Pennant, 1769, Sri Lanka. Four subspecies recognized.

other common names

French: Râle à poitrine blanche; German: Weißbrust-Kielralle; Spanish: Gallineta Pechiblanca.

physical characteristics

11–13 in (28–33 cm); 5.8–11.6 oz (165–328 g). Dark upperparts, white face and underparts, tawny-rufous rear underparts, and yellow bill and legs. Juvenile duller. Chick black and fluffy.

distribution

A. p. phoenicurus: Pakistan and India east to Japan, and south through southeastern Asia to Sundas; northern populations winter to the south, reaching Arabia; A. p. insularis: Andaman and Nicobar Island; A. p. midnicobaricus: central Nicobar Island;A. p. leucomelanus: Sulawesi, western Moluccas, Lesser Sundas.

habitat

Marshes, grass, forest, scrub, and mangroves.

behavior

Not particularly shy. Perches, climbs, and swims well. Territorial when breeding. Characteristic calls include roars, grunts, cackles and croaks; often vocal at night.

feeding ecology and diet

Takes invertebrates, small fish and some plant material. Forages on land and in water.

reproductive biology

Monogamous. Breeds all months, mainly during rains. Nest a cup of twigs, stems, and leaves, close to the ground and near water. Eggs three to nine; color is dull brownish to white/gray with reddish spots and marks. Incubation 20 days by both parents.

conservation status

Not threatened. Common to local over much of its range, which is expanding northeast. Uses humanmade habitats, even in built-up areas.

significance to humans

None known.

Laysan rail

Porzana palmeri

subfamily

Rallinae

taxonomy

Porzanula palmeri Frohawk, 1892, Laysan I.

other common names

French: Marouette de Laysan; German: Laysansumpfhuhn; Spanish: Polluela de Laysan.

physical characteristics

5.9 in (15 cm). Flightless. Light brown, streaked darker on upperparts, ashy-gray from face to breast; some white flank markings. Juvenile buff on underparts.

distribution

Recently extinct; occurred on northwestern Hawaiian Islands; naturally on Laysan Island, introduced to Midway Atoll.

habitat

Tussock grass and scrub thickets.

behavior

Was active, restless, pugnacious and presumably territorial. Showed little fear of humans. Called communally after dusk, with a warbling or rattling song.

feeding ecology and diet

Ate principally insects; also spiders, birds' eggs, carrion, and some plant material. Approached people for food.

reproductive biology

Apparently monogamous. Bred mainly April through July. Nest a cup or ball of grass in shelter of tussock or other vegetation. Eggs two to three.

conservation status

Formerly common on Laysan I.; habitat destruction by introduced rabbits led to its extinction between 1923 and 1936. Introduced to two islands on the Midway Atoll in 1891 and 1910, it thrived but was exterminated by rats that came ashore from a U.S. Navy landing craft in 1943.

significance to humans

None known.

Striped crake

Aenigmatolimnas marginalis

subfamily

Rallinae

taxonomy

Porzana marginalis Hartlaub, 1857, Gabon. Monotypic.

other common names

French: Marouette rayée; German: Graukehl-Sumpfhuhn; Spanish: Polluela Culirroja.

physical characteristics

7–8.3 in (18–21 cm); 1.5–2.2 oz (41.5–61 g). Male dark brown with white streaks on upperparts; anterior underparts pale cinnamon; rear underparts orangy- to reddish brown. Female has gray foreparts; juvenile duller and plainer.

distribution

Ivory Coast east to Cameroon and south to Congo; eastern Democratic Republic of Congo (Zaire) to Kenya and south to northeast South Africa. In south, largely a wet season visitor, retreating towards equatorial regions after breeding.

habitat

Seasonally inundated grassland, pans, and marsh edges.

behavior

Diurnal and secretive. Territorial when breeding; female gives ticking advertising call, often at night.

feeding ecology and diet

Invertebrates, small fish, and frog tadpoles. Forages in grass, mud, and shallow water.

reproductive biology

In captivity is sequentially polyandrous, female mating with two or more males. Breeds mainly during rains. Nest a bowl of grass or sedges in vegetation over water. Eggs: four to five. Incubation: 17–18 days, by male only; young cared for by male; fledge at 28 days; one to three broods per season.

conservation status

Possibly uncommon throughout range; sometimes locally common after good rains.

significance to humans

None known.

Spotted rail

Pardirallus maculatus

subfamily

Rallinae

taxonomy

Rallus maculatus Boddaert, 1783, Cayenne, French Guiana. Two subspecies recognized.

other common names

French: Ràle tacheté; German: Fleckenralle, Spanish: Rascón Overo.

physical characteristics

10–12.6 in (25–32 cm); 4.6–7.7 oz (130–219 g). Long-billed; blackish brown, heavily streaked white on foreparts and barred on flanks; undertail-coverts white; bill yellow-green with red base. Juvenile duller; three morphs: dark morph (almost plain), pale morph (pale underparts), and barred morph (barred underparts).

distribution

P. m. insolitus: Mexico to Costa Rica; P. m. maculatus: Cuba, West Indies, and from Colombia and Ecuador to eastern Brazil and northern Argentina.

habitat

Marshes, swamps, and wet grassland.

behavior

Generally secretive. Gives a distinctive rasping screech "g'reech" and gruff pumping notes. Territorial when breeding.

feeding ecology and diet

Invertebrates; also small fish and some plant material. Forages in mud or shallow water.

reproductive biology

Monogamous. Breeds mainly June through September. Nest a cup of grass or rushes, low in marsh vegetation, often over water. Eggs two to seven.

conservation status

Not threatened, though poorly known and overlooked; may be locally common.

significance to humans

None known.

Chestnut rail

Eulabeornis castaneoventris

subfamily

Rallinae

taxonomy

Eulabeornis castaneoventris Gould, 1844, Flinders R., Gulf of Carpentaria. Two subspecies recognized.

other common names

French: Râle à ventre roux; German: Mangroveralle; Spanish: Cotara Australiana.

physical characteristics

17.3–20.5 in (44–52 cm); 1.2–2.0 lb (550–910 g). Thickset, with gray head and pinkish brown underparts. Three color morphs, with upperparts olive, chestnut, or olive-brown. Juvenile has duller bare parts.

distribution

E. c. castaneoventris: northern coast of Australia; E. c. sharpei: Aru Island.

habitat

Dense mangroves.

behavior

Diurnal and nocturnal, according to tidal cycle. Shy, secretive, and alert. Has a strutting walk; runs very swiftly. Gives a characteristic harsh screech. Territorial when breeding.

feeding ecology and diet

Eats mainly crabs; also other invertebrates. Uses stones as anvils to break shells of hermit crabs.

reproductive biology

Monogamous. Breeds September through February. Nest a bulky platform of sticks, grass, leaves, bark and seaweed, in mangroves. Eggs four to five young fledged at nine weeks.

conservation status

Not threatened. Patchily recorded; possibly overlooked. Probably not uncommon locally.

significance to humans

None known.

Takahe

Porphyrio mantelli

subfamily

Rallinae

taxonomy

Notornis mantelli Owen, 1848, Waingongoro, North Island, New Zealand. Nominate race of North Island recently extinct, may merit species status; one extant race P. m. hochstetteri.

other common names

French: Talève takahé; German: Takahe; Spanish: Calamón Takahe.

physical characteristics

24.8 in (63 cm); 4–9 lb (1.8-4.2 kg). Flightless; thickset, with massive red bill, and purple and green plumage; undertailcoverts white. Juvenile brownish-gray. Chick has black fluffy down and black bill that turns red in the adult stage.

distribution

Fiordland, South Island, New Zealand; also introduced to four nearshore islands.

habitat

Alpine tussock grassland, scrub and beech forest (Fiordland); pastures (islands).

behavior

Strongly territorial; permanent territories 5–148 acres (2–60 ha), much smaller on islands. Shy; normally has slow, deliberate walk but runs quickly. During winter snow, descends from grassland to forest or scrub up to 6.2 miles (10 km) away.

feeding ecology and diet

Eats leaf bases of alpine grasses; in winter also seeds and fern rhizomes. On islands eats introduced grasses.

reproductive biology

Breeds mainly October through December. Nest a bowl of grass on ground. Eggs one to three (usually two); pale buff color with blotches of mauve and brown; incubation 29–31 days. Young dependent on adults for food for four months. Age of first breeding two years. Survival to one year 27–71%; survival on islands 89%.

conservation status

"Rediscovered" in 1948; some translocated to islands in 1984–1991. Captive-bred birds are released into all populations. Endangered, with a very small population (100–160 birds from 1980 to 2000 in Fiordland; 55 adults on islands in 1998). Island populations breed successfully.

significance to humans

Possibly hunted widely in the past.

Giant coot

Fulica gigantea

subfamily

Rallinae

taxonomy

Fulcia [sic] gigantea Eydoux & Souleyet, 1841, Peru. Monotypic.

other common names

French: Foulque géante; German: Riesenbläßhuhn; Spanish: Focha gigante.

physical characteristics

19–23 in (48–59 cm); 4.5–5.3 lb (2.02–2.4 kg). Heavy-bodied, with small head and knobs above eyes. Dark slate-gray with white on undertail-coverts. Bill and shield white, yellow and red; legs red. Juvenile dark dull gray, with paler underparts and dark bare parts. Adults normally too heavy to fly.

distribution

Andes of central Peru, Bolivia, north Chile, and northwestern Argentina.

habitat

Ponds and lakes in highlands of puna zone.

behavior

Permanently territorial. Quite confident unless persecuted.

feeding ecology and diet

Vegetarian, taking mostly aquatic vegetation; also grazes on shore. Feeds from water surface, up-ends, occasionally dives.

reproductive biology

Monogamous. Breeds all year, peaking in winter. Nests in water c. 3 ft (1 m) deep; nest permanent; of aquatic vegetation, often resting on bottom of lake and 3 ft (1 m) wide and up to 10 ft (3 m) long at waterline, projecting up to 20 in (50 cm) above water. Eggs three to seven; young fed until two months old; fledge at four months.

conservation status

Locally scarce to common. In 1998 considered Vulnerable in Chile.

significance to humans

Many eggs are taken by people at some sites.

Resources

Books

Bird, D.M. The Bird Almanac: The Ultimate Guide to Essential Facts and Figures of the World's Birds. Buffalo: Firefly Books, 1999.

Craig, J.L., and I.G. Jamieson. "Pukeko: different approaches and some different answers." In Co-operative Breeding in Birds: Long-term Studies of Ecology and Behavior, edited by P.B. Stacey and W.D. Koenig. Cambridge: Cambridge University Press, 1990.

Cramp, S., and K.E.L. Simmons, eds. Hawks to Bustards. Vol. 2, The Birds of the Western Palearctic. Oxford: Oxford University Press, 1980.

Fjeldså, J., and N. Krabbe. Birds of the High Andes. (Copenhagen and) Svendborg: Zoological Museum, University of Copenhagen and Apollo Books, 1990.

Glutz von Blotzheim, U.N., K.M. Bauer, and E. Bezzel. Handbuch der Vögel Mitteleuropas. Vol. 5. Frankfurt am Main: Akad. Verlag, 1973.

Marchant, S., and P.J. Higgins, eds. Raptors to Lapwings. Vol. 2, Handbook of Australian, New Zealand and Antarctic Birds. Melbourne: Oxford University Press, 1993.

Olson, S.L. "A synopsis of the fossil Rallidae." In Rails of the World: A Monograph of the Family Rallidae, by S.D. Ripley. Boston: Godine. 1977.

Potapov, R.I., and V.E. Flint, eds. Galliformes, Gruiformes. Vol. 4, Handbuch der Vögel der Sowetjunion, edited by V.D. Illicev and V.E. Flint. Wittenberg Lutherstadt: A. Ziemsen Verlag, 1987.

Stattersfield, A.J., and D.R. Capper, eds. Threatened Birds of the World: The Official Source for Birds on the IUCN Red List. Cambridge, United Kingdom: BirdLife International, 2000.

Tacha, T.C., and C.E. Braun, eds. Management of Migratory Shore and Upland Game Birds in North America. Washington, DC: International Assoc. Fish and Wildlife Agencies, 1994.

Taylor, B., and B. van Perlo. Rails: A Guide to the Rails, Crakes, Gallinules and Coots of the World. Sussex: Pica Press, 1998.

Urban, E.K., C.H. Fry, and S. Keith, eds. The Birds of Africa. Vol. 2. London: Academic Press, 1986.

Wanless, R.M. The Reintroduction of the Aldabra Rail Dryolimnas cuvieri aldabranus to Picard Island, Aldabra Atoll. MSc thesis: University of Cape Town, 2002.

Periodicals

Eddleman, W.R., F.L. Knopfe, B. Meanley, F.A. Reid, and R. Zembal. "Conservation of North American Rallids." Wilson Bulletin 100 (1988): 458–475.

Halse, S.A., and R.P. Jaensch. "Breeding Seasons of Water-birds in South-western Australia—The Importance of Rainfall." Emu 89 (1989): 232–249.

Livezey, B.C. "A Phylogenetic Analysis of the Gruiformes (Aves) Based on Morphological Characters, with an Emphasis on the Rails (Rallidae)." Philosophical Transactions of The Royal Society of London B 353 (1998): 2077–2151.

McRae, S.B., and T. Burke. "Intraspecific Brood Parasitism in the Moorhen: Parentage and Parasite-host Relationships Determined by DNA Fingerprinting." Behaviorial Ecolology and Sociobiolology 38 (1996): 115–129.

Olson, S.L. "Evolution of the Rails of the South Atlantic Islands (Aves: Rallidae)." Smithsonian Contribtributions to Zoolology 152 (1973): 1–53.

Olson, S.L. "A Classification of the Rallidae." Wilson Bulletin 85 (1973): 381–446.

Other

"Black Rail." Illinois Birds. 30 June 1998. Illinois Natural Resources Information Network. 06 Dec. 2001 <http://www.inhs.uiuc.edu/chf/pub/ifwis/birds/black-rail.html>

Tan, Ria. "White-breasted Waterhen." Apr 2001. Sungei Buloh Nature Park. 06 Dec. 2001 <http://www.naturia.per.sg/buloh/birds/Amaurornis_phoenicurus.html>

Barry Taylor, PhD