Babblers (Timaliidae)

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Babblers

(Timaliidae)

Class Aves

Order Passeriformes

Suborder Passeri (Oscines)

Family Timaliidae

Thumbnail description
Small to medium-sized birds of highly variable shapes and colors

Size
3–16 in (7.6–40 cm); 0.35–5.25 oz (10–150 g)

Number of genera, species
Roughly 50 genera, around 280 species

Habitat
Generally forests, but some semi-desert, scrub, grassland, and wetlands

Conservation status
Endangered: 5 species; Vulnerable: 22 species; Near Threatened: 39 species; Data Deficient: 1 species

Distribution
Eurasia (excluding Japan), Wallacia, Africa, Madagascar, the Pacific Coast of North America (Introduced in Hawaii)

Evolution and systematics

Nearly a century ago, Ernst Hartert, Curator of the incredible Rothschild collection of preserved birds, observed: "What can't be classified is regarded as a babbling thrush." In the opinions of many of today's ornithologists, Hartert then proceeded to confuse matters further. The convoluted history of babbler classification is well treated in Sibley and Ahlquist's Phylogeny and Classification of Birds, and the reader is best referred to that book. Suffice it to say that exactly what a babbler is, and how it should be classified, has been, and remains, a controversy among ornithologists.

Jean Delacour's masterful arrangement of babblers, proposed in the 1940s and 1950s, has worked remarkably well, even as 1980s experiments in DNA-hybridization completely reorganized bird classification, and the rationale behind it. Delacour considered the babblers a vast subfamily, Timaliinae, one among many in the huge family Muscicapidae, created by Hartert in 1910, and now determined to be a hodgepodge of unrelated birds. Delacour divided his Timaliinae into a series of tribes, including one for both the wrentit (Chamaea) and bearded reedling (Panurus), both of which were traditionally placed in different families. He also gave tribe status within the babblers to the parrotbills (Paradoxornis and Conostoma) and to the genus Picathartes. Picathartes had long been especially controversial, being assigned to the starling and crow families by various authorities.

The DNA research of Charles Sibley and his associates has led them to place the babblers in the family Sylviidae, together with many of the birds traditionally called Old World warblers. Within this family, they have divided the babblers into two subfamilies. Two genera, the laughing thrushes (Garrulax) and liocichlas (Liocichla), are given their own subfamily, Garrulacinae. The other babblers, including the parrotbills, which have traditionally been placed in a subfamily, Paradox-ornithinae (or Panurinae), are included by Sibley in the sub-family Sylviinae, divided into three tribes. All but one of these are in the tribe Timaliini, the exception being the wrentit, comprising its own tribe, Chamaeini. The third tribe, Sylviini, consists of the warblers of the genus Sylvia, including such familiar European birds as the blackcap, whitethroats, and the garden warbler. One controversial inclusion in the tribe Timaliini, as Sibley defines it, is the genus Rhabdornis, the Philippine creepers, traditionally placed near the nuthatches and creepers. On the other hand, the genera Pomatostomus and Garritornis, considered by Delacour and others to be close relatives of the scimitar babblers, now constitute their own family, Pomatostomidae, which Sibley and colleagues placed near the crow family, Corvidae. The rail babblers (Eupetes and Ptilorrhoa), which Delacour initially assigned to a tribe within the Timaliinae, then later considered a separate subfamily, are classified by Sibley as a subfamily within the Corvidae. It is not surprising that Sibley removed Picathartes, long a source of disagreement, from the Timaliinae, nor is his refusal to place their family in close proximity to any other. What is truly startling is his inclusion, in the family Picathartidae, of the rock jumpers (Chaetops), traditionally classified as thrushes (Turdidae).

Paleontology has, thus far, not served to clarify the origins and relationships of babblers, the only fossils being a Middle Pleistocene example of the modern species, the Arabian babbler (Turdoides squamiceps). DNA research seems to support Asian origins in the Oligocene, about 40 million years ago.

Physical characteristics

Almost any book published over the last 50 years is likely to present a different definition of exactly what constitutes "the babblers." Over 40 years ago, Oliver Austin, in his now classic Birds of the World, summed up the situation by describing this group as "poorly defined and loosely delimited." All of these birds have 10 primary feathers, the outermost shorter than the rest. The tail is composed of 12 retrices. The plumage is soft and loose. Juvenal plumage is never speckled. The legs and feet are proportionately large and powerful, while flight potential is comparatively small (leading, in part, to the great number of species with small ranges). In appearance babblers range from the utterly nondescript to the unforgettable.

Despite the great disagreements among most books, one usually finds the babblers arranged thusly. To begin with are several genera of "small brown birds," otherwise referred to as "somber," "dull" or "nondescript." Often their most recognizable feature, at least when they're alive, are their voices. The number of genera is in dispute, but a few more than 30 species compose these "jungle-babblers" (Pellorneum, Trichastoma, Illadopsis, Kakamega, Malacopteron, etc.), all of them tropical, most of them forest-dwellers, occurring through large areas of Africa and Asia. There follow around a dozen scimitar babblers (Pomatorhinus, Xiphirhynchus, and Jaboulleia), named for their long curved bills, some quite spectacular. They appear to be closely related to the around 20 very different-looking wren-babblers (Rimator, Ptilocichla, Napothera, Pnoepyga, Spelaeornis, etc.). These are short-tailed, mostly plump-bodied birds, some tiny, none with brilliant colors, but some with vivid patterns. Some strongly resemble New World antbirds. Scimitar babblers and wren-babblers are Asian, with one species extending down the Lesser Sundas to Timor. What comes next depends on the book one is reading, and here things may differ wildly. Suffice to say that there follows the majority of the babblers, in many genera great and small. Towards the end, one usually finds the parrotbills (Paradoxornis and Conostoma) of Palearctic and Oriental Asia, and the primarily Palearctic bearded reedling (Panurus biarmicus), mostly compact-bodied and long-tailed, many with distinctive bills, one, (Paradoxornis paradoxus), uniquely with three toes instead of four. Finally, one may or may not find the unique and magnificent Picathartes. What they are remains a question.

Distribution

Asia is the center of evolution for babblers. With the exception of the unique picathartes, which DNA research suggests may not be babblers at all, but an ancient African lineage, most of the 33 African babblers appear to be derivative of large Asian genera. Only recently have several of them been removed from Asian genera, and placed in ones exclusive to Africa, such as Illadopsis and Pseudoalcippe, while 15 are clearly members of Turdoides, widespread in the Indian subcontinent. Typical of the island, the six species in Madagascar, all endemic, have no connection to the African mainland.

China, where the Palearctic and Oriental zoogeographical realms meet, is a great center of distribution, and at least 143 (including four Taiwanese endemics) occur there, many nowhere else. Chinese babblers include 35 of the 49 species of laughing thrushes (Garrulax), 14 of the 16 fulvettas (Alcippe), and every one of the 19 parrotbills (Paradoxornithinae). Through Southeast Asia, there continues a great diversity of babblers, especially in Indochina, where several were only recently discovered. Fifty-five species have been recorded for Java, Sumatra, and Borneo, many of them endemic, or otherwise found only on the Malay Peninsula. While the "babblers" inhabiting Australia and New Guinea are no longer considered Timaliids, a few species occur in Wallacea, including the enigmatic malia (Malia grata), found only on Sulawesi, and remarkably, the fragile-looking pygmy wren-babbler (Pnoepyga pusilla), found all the way from Nepal to Timor. It is remarkable that Japan and Okinawa have no native babblers. On the other hand, 19 species occur in the Philippines, including 10 of the 24 species of Stachyris, and all but one are endemic. The Philippines and Timor are the western-most of the Pacific islands where babblers occur naturally. (Four Asian species have been successfully introduced in Hawaii.)

The Indian subcontinent, including Sri Lanka, is another babbler stronghold, and 131 species are recorded there. The majority are birds of the Himalayas, shared with China, but a number are endemic to Peninsular India or Sri Lanka. In the Middle East, babblers are represented by three species of Turdoides. Europe, including the United Kingdom, share the bearded reedling (Panurus biarmicus) with temperate Asia. Finally, on the Pacific Coast of North America is the wrentit (Chamaea fasciata).

Habitat

Forests are the stronghold of babblers, and most are dependent on one type or another. However, natural selection being what it is, there are babblers adapted to scrublands, near-desert, grassy savannas, and marshes, and some species have become partial to orchards, fields, and gardens.

Behavior

For so diverse a group of birds, generalizations are difficult. As did Bertram Smythies in the first edition of this encyclopedia, one cannot do better than to quote Jean Delacour, from his monumental 1946 monograph of the babblers: "They move restlessly among twigs and on the ground; they hop about and dig among fallen leaves. Usually they live in the undergrowth, sometimes on the ground among dense plant growth, fallen branches, climbers and evergreen trees, where they can be observed searching for berries and insects. While doing so, they move busily, flutter the wings a great deal, wag their tails and utter noisy calls. As rule, they are loud and varied vocally, hence the name babbler, for they are virtually never quiet. Some sing very well and their full melodies ring out far. Outside the breeding season, they move about in small troops. Often they join with other birds into the mixed flocks characteristic of tropical forest, all seeking food together."

Delacour's description fits most of the forest-dwelling babblers fairly well. It is in other sorts of habitat that particularly interesting variations have evolved. The Arabian babbler (Turdoides squamiceps), of the semi-desert, with its highly developed "tribal" social system, and the semi-aquatic bearded reedling (Panurus biarmicus) of the marshes, are two striking examples.

Feeding ecology and diet

Insects are the core of babbler diets. Some species appear to feed exclusively on them, while many also eat fruit, other invertebrates, and small frogs and reptiles. As with any huge family, there have evolved some peculiar specialists. While the bearded reedling feeds vigorously on insects for most of the year, and rears its young entirely on them, it subsists on seeds during the winter, its digestive tract making remarkable adjustments to this change in diet. As one might expect, the desert-adapted Arabian babbler will eat practically anything. On the other hand, the fire-tailed myzornis (Myzornis pyyrhoura) has come to resemble a hummingbird or sunbird, consuming nectar with its insects, and becoming a pollinator in the process.

Reproductive biology

In general, babblers form pairs in the breeding season, establish territories, raise one or two broods, then reassemble as flocks. Again, this seems to be the case for forest species, with most exceptions being birds of other habitat. The bearded reedling has evolved to produce up to four broods in quick succession each breeding season, with the first potentially able to breed themselves by season's end. The Arabian babbler's reproductive system, however, is remarkably different. Its highly regimented social units, with birds waiting as long as six years for their first opportunity to breed, is only the most extreme situation in its large and widespread genus Turdoides. Researchers have confirmed that at least 14, and possibly 26, of the 29 species in this genus practice some sort of cooperative breeding, with groups defending territories where only a few members will breed at any given time. The Turdoides are primarily open-country birds. In India, the tendency to always remain in their social unit has earned some species the name "Seven Sisters," and India's revered ornithologist Salim Ali refers to their groups as "sisterhoods." Another departure from the norm are the colonial picathartes, who use their unique, mud pottery-like nests, built on rock faces, as residences, and raise chicks in them year after year. Such nests are unique among babblers, who, in general, construct cup or bowl-shaped, or spherical, nests of plant materials, usually not far from the ground. Eggs run a spectrum from white and patternless, to various beautiful colors and intricate patterns of spots and streaks.

Conservation status

As of 2002, the IUCN and BirdLife International designated five babblers as Endangered, 22 as Vulnerable, and 39 as Near Threatened. In addition, it appears that subspecies of two otherwise globally non-threatened species may be extinct: The southern Turkish bearded reedling (Panurus biarmicus kosswigi) appears to have been a victim of wetlands destruction, while the mysterious "Astley's leiothrix" (Leiothrix lutea astleyi), known only from the bird trade more than 80 years ago, may have been exterminated through trapping.

All five Endangered babblers are primarily threatened by habitat loss. All are forest birds. Two are Philippine endemics: the flame-templed babbler (Stachyris speciosa) occurs only on Negros and Panay, while the Negros striped-babbler (S. negrorum) is entirely restricted to that severely deforested island. The remaining three depend on high-elevation forests: the Nilgiri laughing thrush (Garrulax cachinnnans), one of many imperiled inhabitants of India's Nilgiri Hills, the white-throated mountain babbler (Kupeornis gilberti), known to science only since 1949, restricted to several places in Nigeria and Cameroon, and the gray-crowned crocias (Crocias lang-bianis), rediscovered in 1994 after 56 years of no records, from a few locations in Vietnam.

Three Vulnerable species, the Omei Shan liocichla (Liocichla omeiensis), found only around Mt. Emei, in southwestern China, and the white-necked and gray-necked picathartes (Picathartes gymnocephalus and P. oreas), of the Guinea forests of West Africa, were vigorously exploited by the cage-bird trade, resulting in their listing by the Convention on International Trade in Endangered Species (CITES). Like the 19 other Vulnerable babblers, they are threatened by habitat destruction as well.

Fourteen island endemics, seven restricted to the Philippines, are included among the 39 Near Threatened species, and of the remainder, nine occur only in the Malay Peninsula, Sumatra, and Borneo. Again, loss of habitat, in often restricted ranges, is the cause for their designation. Finally, there is one categorized as Data Deficient, the miniature titbabbler (Micromacronus leytensis), at 3 in (7.6 cm), the smallest babbler. Restricted to the Philippine islands of Leyte, Samar, and Mindanao, it is a forest-dependent species in a land of increasing deforestation, and has remained rare, and little known since its discovery in 1961.

Significance to humans

Babblers have long been admired for their appearance, songs, and behavior. One aspect of this admiration has been the compulsion to keep them in captivity. The enormous volume of commercial trade in living babblers has caused increasing concern. At the same time, over 30 species have been hatched in captivity.

Some Chinese babblers, especially laughing thrushes (Garrulax sp.) and parrotbills (Paradoxornis sp.) cause some damage to crops, but this appears to be minor, and offset by insect control by these same birds. On the other hand, babblers are playing an increasingly important role in the developing economy of ecotourism. Such species as the fire-tailed myzornis are specifically featured in advertisements enticing trekkers to Nepal and Bhutan, already attracted simply by the potential of seeing great numbers of species in the foraging "bird waves" that sweep across the Himalayas. As ecotourism continues to grow, increasing numbers of people from around the world will enjoy the magnificent mixed choruses of otherwise unobtrusive brown babblers in Borneo or Myanmar, search for Madagascar's peculiar endemics, observe the colonial nesting of fantastic-looking picathartes in Ghana or Gabon, and search for laughing thrushes only recently unknown to science in the highlands of Vietnam. At the same time, vigorous efforts are being made by environmentalists in these countries to instill pride in a precious natural heritage.

Species accounts

List of Species

Rufous-winged akalat
Chestnut-backed scimitar-babbler
Pygmy wren-babbler
Flame-templed babbler
Wrentit
Arabian babbler
White-crested laughing thrush
Yellow-throated laughing thrush
Hwamei
Omei Shan liocichla
Red-billed leiothrix
Black-crowned barwing
Golden-breasted fulvetta
Whiskered yuhina
Fire-tailed myzornis
Crossley's babbler
Bearded reedling
Vinous-throated parrotbill
White-necked picathartes
Gray-necked picathartes

Rufous-winged akalat

Trichastoma rufescens

subfamily

Timaliinae

taxonomy

Turdirostris rufescens Reichenow, 1878, Liberia. Monotypic. Endemic African genus Illadopsis has been merged by various authors into Asian genera Malacocincla or Trichastoma.

other common names

English: Rufous-winged illadopsis; French: Akalat à ailes rousses; German: Rostschwingen-Buchdrossling.

physical characteristics

7.2 in (18 cm); 1.2 oz (35 g). Profoundly nondescript thrush-like bird, as are most of the other six African members of the genus. Sexes alike. Rufous-brown dorsally, whitish ventrally, with indistinct brown wash on chest and brownish gray sides. Eyes dark; legs and feet pale.

distribution

Endemic to Upper Guinea forest, from southern Senegal to Togo.

habitat

Undergrowth in primary and disturbed forest.

behavior

Ground dweller. Found in pairs or small flocks, sometimes associates with other species of insectivorous passerines. Territorial.

feeding ecology and diet

Actively searches the ground for insects and tiny frogs, scratching at substrate with feet.

reproductive biology

Undescribed.

conservation status

Near Threatened due to former paucity of observations, and continuous destruction of forests throughout narrow range. Recent surveys have found species common in some localities, and occurring in areas where previously unrecorded. Population densities in logged forest are a fraction of those in primary habitat, but has been found in plantations and open thickets.

significance to humans

None known.

Chestnut-backed scimitar-babbler

Pomatorhinus montanus

subfamily

Timialiinae

taxonomy

Pomatorhinus montanus Horsfield, 1821, Java. Four subspecies.

other common names

English: Yellow-billed scimitar babbler; French: Moineau friquet; German: Rotrückensäbler.

physical characteristics

8 in (20 cm). Striking bird reminiscent of New World thrashers (Mimidae). Bright chestnut mantle, flanks, and vent. Brownish primaries and tail. White chest and throat. Remainder of head black, except for well-defined white eyebrow. Long curving bill bright yellow, with black at rear of upper mandible, extending along part of culmen. Legs gray. Melodious voice.

distribution

Malay Peninsula, Sumatra, Java, Bali, and Borneo.

habitat

Lower and middle stories of forests, at all altitudes.

behavior

Often associated with laughing thrushes (Garrulax sp.). Single birds may be observed, but usually in small flocks. Vocal, but usually hard to see.

feeding ecology and diet

Continually hunting for snails, spiders, and insects, as well as small seeds and fruits.

reproductive biology

Monogamous. Presumed to be similar to that of other members of genus, with cone or bowl-shaped nest of plant material constructed by both sexes, and containing four white eggs.

conservation status

Not threatened but endemic in areas subject to increasing habitat loss due to logging.

significance to humans

For many years, fairly small numbers have been exported through the cage-bird trade.

Pygmy wren-babbler

Pnoepyga pusilla

subfamily

Timaliinae

taxonomy

Pnoepyga pusillus Hodgson, 1845, Nepal. Eight subspecies; some may constitute separate species.

other common names

English: Lesser scaly-breasted wren-babbler, brown wrenbabbler; French: Turdinule mailée; German: Moostimalie.

physical characteristics

3.5 in (9 cm). Tiny, almost tail-less brown bird with speckled underparts, and dark eyes. Light and dark-breasted phases are present. Short, thin pointed bill.

distribution

Himalayas, east from Nepal, southern China, Taiwan, Indochina, Malay Peninsula, Sumatra, Java, Flores, and Timor.

habitat

Highland forest floor thickets and moist ravines.

behavior

Highly furtive and mouse-like, staying concealed in ferns and other plants, but surprisingly loud, piercing whistle, followed by softer note, is heard often, apparently a contact call. Distinctive voice.

feeding ecology and diet

Presumably small invertebrates.

reproductive biology

Undescribed.

conservation status

Not threatened, but would be vulnerable to any logging of montane forests.

significance to humans

Target species for ecotourists.

Flame-templed babbler

Stachyris speciosa

taxonomy

Dasycrotapha speciosa Tweeddale, 1878, Negros. Listed by some authorities in the monotypic genus Dasycrotapha.

other common names

English: Rough-templed tree-babbler; French: Lobélia spéciosa; German: Goldstirn-Buschtimalie.

physical characteristics

5.25 in (13.2 cm). Unique, rather bizarre appearance: black head with golden yellow forehead, lores, eye ring, and chin; bright reddish orange tufts on side of head. Streaked olive mantle with brown wings and tail. Breast yellow with black spots.

distribution

Restricted to central Philippine islands of Negros and Panay.

habitat

Lowland forest understory.

behavior

Forages in small flocks, individually, or with other species. Very responsive to contact calls.

feeding ecology and diet

Presumed to be insects and seeds.

reproductive biology

Nest and eggs undiscovered as of 2000.

conservation status

Endangered due to severe reduction and fragmentation of range resulting from deforestation.

significance to humans

Negros and Panay "flagship" potential.

Wrentit

Chamaea fasciata

subfamily

Timaliinae

taxonomy

Parus fasciatus Gambel, 1845, Monterey, California. Six subspecies. While Delacour long ago considered this species a babbler, many authorities have persisted in assigning it its own family or subfamily. However, the DNA hybridization research

of Sibley and Ahlquist suggests the wrentit is the only New World babbler, whose ancestors crossed the Bering land bridge in the mid-Miocene (15–20 million years ago). Sibley and Ahlquist do place it in its own tribe, Chamaeini.

other common names

French: Cama brune; German: Chaparraltimalie.

physical characteristics

6.3 in (16 cm); 0.5 oz (14 g). Gray (southern) or brown (northern) compact bird with long tail (usually held up), pale eyes, and small, sharp bill. Song is a very distinctive accelerating series of high notes.

distribution

Pacific Coast of North America, from northern Oregon to northern Baja California.

habitat

Chaparral and coniferous scrub.

behavior

Generally live in pairs, communicating continuously with unique calls. Frequents heavy low vegetation.

feeding ecology and diet

Forages for insects and berries in undergrowth.

reproductive biology

Monogamous, paired for life. Sexes share in all aspects of nesting and chick rearing. Nest is located deep in bushes, a tightly woven cup of twigs, bark, and feathers, lined with spider webs. Three to five greenish blue, unpatterned eggs compose clutch.

conservation status

Not threatened, though significant portions of range have been developed.

significance to humans

Popular with birdwatchers.

Arabian babbler

Turdoides squamiceps

subfamily

taxonomy

Malurus squamiceps Cretzschmar, 1827, Transjordan. Three subspecies.

other common names

French: Cratérope écaillé; German: Graudrossling; Spanish: Tordalino Arábigo.

physical characteristics

11 in (28 cm). Slender grayish brown, long-tailed bird, which may remind Americans of thrashers (Mimidae). Plumage identical in both sexes, but male's eyes pale yellow, female's brown.

distribution

Israel (Negev, Dead Sea Depression, Arava Valley, Sinai), Jordan, coastal Arabian Peninsula (except for Red Sea Coast).

habitat

Dependent on vegetation and water sources, but otherwise a dry country bird, favoring acacias, tamarisk, saltbush, and datepalm groves and gardens.

behavior

Complex social system, reminiscent of wolves. Lives in groups of six to a dozen, but sometimes more than 20, composed of an alpha breeding pair, usually the oldest birds, which maintain a strict dominance hierarchy over rest of unit. Groups compete with others for fixed territories, and may displace them, sometimes violently. Most unit members related to breeders, but some recruitment of unrelated birds from nearby colonies. When not actively foraging, generally secreted in bushes.

feeding ecology and diet

Opportunistic feeders, energetically foraging in groups for anything edible. Main items include very wide variety of

arthropods, small reptiles, berries, flowers, nectar, seeds, and garbage.

reproductive biology

In some units, only the alpha male mates with any of the females. In others, subordinate males may breed with subordinate females, but the alpha female is always the one most fiercely defended by the alpha male. Courtship behavior appears to be initiated by females soliciting subordinate males in the group, causing the alpha male to aggressively prevent copulations. No copulations take place if other members of the group are present. (As a rule, they actively interfere with impending matings, only dominant males being able to prevent other group members from following potential breeding pair). Successful copulation takes place in seclusion of bushes. Nest is large, untidy, cup-shaped structure, composed of coarse plant materials, with no clearly defined lining. Generally, only one is constructed in a group's territory. Up to three females may lay eggs in group's nest. Eggs are glossy turquoise. Clutch size per female varies from three to five. At night, only the alpha female incubates, but any female in the group may do so in daytime. Eggs take 13–14 days to hatch, and hatching may occur continuously through breeding season, so that nest may be occupied by large number of chicks of disparate ages. All members of group feed chicks, often competing to do so. Unusually long reproductive longevity for a passerine bird. Sexual maturity not attained until two years, some may remain non-breeders in group as long as six years. Individuals may still be group's alpha breeders at 13 years of age.

conservation status

Globally non-threatened. Range has expanded, especially in Israel, due to irrigation and abundance of food (including garbage) around residences.

significance to humans

This species may pose some harm to certain types of agriculture, eating such products as sorghum. On the other hand, they consume vast numbers of potentially harmful insects, such as termites and weevils, and are especially avid tick eaters. Subject of continuous research for decades.

White-crested laughing thrush

Garrulax leucolophus

subfamily

Timaliinae

taxonomy

Corvus leucolophus Hardwicke, 1815, Uttar Pradesh. Five subspecies. DNA hybridization study suggests Garrulax and Liocichla compose a separate subfamily (Garrulacinae) from the other babblers, which belong in another subfamily, together with some warblers (Sylviinae).

other common names

English: White-crested jay thrush; French: Garrulaxe à huppe blanche; German: Weisshaubenhäherling.

physical characteristics

12 in (30 cm). Instantly recognizable dark-bodied, whiteheaded bird with bushy crest and black mask and beak. Sexes monomorphic. Subspecific variation fairly pronounced: Himalayan nominate subspecies has brown chest and underparts. These areas are white in southeastern Yunnan and Indo-Chinese

G. l. diardi, which also has a much grayer nape. G. l. patkaicus of Assam, northern Myanmar, and western Yunnan, has brown underparts, but a pure white nape. Sumatran G. l. bicolor has underparts, back, and tail blackish brown, a very different shade from that of other subspecies.

distribution

The length of the Himalayas, with some gaps, Assam, Myanmar, southern China, Indochina, and an isolated subspecies in Sumatra.

habitat

Evergreen forests, especially secondary, where heavy under-growth and bamboo stands abound.

behavior

Frequents lower storeys of forest, roaming in flocks of varying sizes (up to 40), often with other bird species. Distinctive ringing, antiphonal vocalizations are frequent.

feeding ecology and diet

Groups continuously forage through plants and leaf litter for insects, lizards, fruit, nectar, and seeds.

reproductive biology

Species observed in flocks during the breeding season. From captive observations, however, it may prove that only one pair actually breeds. Clutch size is usually three to five. Eggs are white. The wide, shallow nest is usually only about 6 ft (1.8 m) above ground. Bamboo leaves are preferred material for nest. In China, two broods are usually raised each year, any time from March through August. In zoos, breedings have been recorded all year.

conservation status

Not threatened. Preference for secondary forest decreases vulnerability to habitat loss. Traditionally exploited for cage-bird trade; as of 2002, only Indonesia exporting.

significance to humans

Very popular in zoos; more than 400 hatched in United States alone since 1968.

Yellow-throated laughing thrush

Garrulax galbanus

subfamily

Timaliinae

taxonomy

Garrulax galbanus Godwin-Austin, 1874, Manipur. Three subspecies. Chinese populations may constitute separate species.

other common names

English: Yellow-bellied laughing thrush; French: Garrulaxe à gorge jaune; German: Gelbbauchhäherling.

physical characteristics

9.1 in (23 cm); 1.75 oz (50 g). Elegant thrush-shaped bird with brown mantle, black mask, and yellow throat and underparts. Sexes monomorphic. Nominate subspecies, from the western part of the range, has an olive green nape and crown, while the two isolated eastern subspecies have brilliant dark blue napes and crowns instead. Jiangxi population has a clear, brilliant yellow chest, while birds from Yunan have yellowish gray chests.

distribution

Western population in eastern India (the Manipur Valley, Nagaland, Assam, and Mizoram), the Chin Hills of Mynamar, and possibly Bangladesh. Isolated Chinese populations: one in Jiangxi Province, another in Yunnan, and perhaps another yet to be ascertained, from which captive specimens have arrived.

habitat

Dense scrub.

behavior

Lives in flocks, stays near ground.

feeding ecology and diet

Hunts through leaf-litter and other substrate for insects. Also consumes fruit.

reproductive biology

Captive birds produce clutches of two or three eggs, in smallish cup-shaped nests of twigs and plant fibers. Incubation, lasting 13 days, is performed by both parents. As with other Garrulax, while a flock may be present during nesting, only one pair actually breeds.

conservation status

Though previously listed as Near Threatened, IUCN designated it as globally non-threatened as of 2002. However, isolated Chinese populations, possibly constituting a separate species, appear to be very small and occupy limited ranges. Not listed by CITES. Commercial live bird exports banned by People's Republic of China in 2001.

significance to humans

International captive breeding projects hatched over 100 since 1990.

Hwamei

Garrulax canorus

subfamily

Timaliinae

taxonomy

Turdus canorus Linnaeus, 1758, Amoy. Three subspecies.

other common names

English: Melodious jay thrush, spectacled thrush, Chinese thrush; French: Garrulaxe hoamy; German: Augenbrauenhäherling.

physical characteristics

9 in (22 cm). Thrush-shaped. Uniformly brown with fine darker streaking. Distinctive white eye ring, with long rearwards extension lending an "Egyptian" look. Bill pale. Taiwanese subspecies more heavily streaked with much reduced eye ring.

distribution

Central and southeast China, Taiwan, Hainan, and northern Indochina. Feral populations established on the Hawaiian Islands of Oahu, Maui, Hawaii, Molokai, and Kaua'i.

habitat

Scrub, secondary forest, and nearby farmland.

behavior

A notably shy bird, yet famous for male's robust song. Stays near the ground, a weak flyer. Usually in pairs or small groups.

feeding ecology and diet

Forages in leaf litter for insects, fruit, seeds, and insects.

reproductive biology

Monogamous. Large bowl or oval-shaped nest situated in thickets, made of leaves, pine needles, grasses, fine twigs, vines, and other plant material. May or may not be lined. Three to five bluish eggs, sometimes spotted. Female alone incubates for 15 days. Chicks fledge after 13 days of being fed by both parents. Two breeds raised between February and September.

conservation status

Not threatened, though Taiwanese G. canorus taewanus may be threatened due to introduction of mainland nominate subspecies. As disturbed and agricultural habitat is favored, still quite common on Chinese mainland, despite massive exploitation for the cage-bird trade. Added to Appendix II of CITES in 2001.

significance to humans

Most revered songbird in China. Apparently inflicts limited damage to peanut and pea crops in China, but negative presence offset by insect control.

Omei Shan liocichla

Liocichla omeiensis

subfamily

Timaliinae

taxonomy

Liocichla omeiensis Riley, 1926, Mt. Emei, Sichuan. Only recently recognized as full species, distinct from Taiwanese endemic Steere's babbler (L. steerii).

other common names

English: Mount Omei babbler; French: Garrulaxe de l'Omei; German: Omei-Haeherling.

physical characteristics

7 in (17 cm). Gray with bright reddish orange pattern on wings, squared-off, red-tipped tail, red-and-black vent, yellowish ear coverts, dark eyes and bill.

distribution

The vicinity of Mt. Emei, southern Sichuan and northeastern Yunnan.

habitat

Mountain forests with bamboo stands.

behavior

Noted for being not as shy as other babblers, but rather curious.

feeding ecology and diet

Small flocks forage for insects and fruits both in trees and on the ground.

reproductive biology

Male displays to female by exposing bright colors on wings while vocalizing. Nest and eggs undescribed.

conservation status

Vulnerable. Still occurs in some numbers, but threatened both by bamboo shoot harvesting and other forms of habitat destruction, and by the cage-bird trade. Accorded CITES Appendix II status in 1997. Export from China prohibited in 2001.

significance to humans

Target species for ecotourists.

Red-billed leiothrix

Leiothrix lutea

subfamily

Timaliinae

taxonomy

Sylvia lutea Scopoli, 1786, Anhui. Five subspecies generally recognized.

other common names

English: Pekin robin, red-billed hill tit; French: Léiothrix jaune; German: Sonnenvogel.

physical characteristics

6 in (15.5 cm); 0.8 oz (22 g). Unmistakable combination of red bill, dark eyes with white ring, orange chest, yellow

throat, olive back, orange and yellow wings, and uniquely forked tail.

distribution

The length of the Himalayas, Assam, western and northern Myanmar, southern China, northern Vietnam. Introduced in Hawaiian islands of Oahu, Hawaii, Maui, Molokai, and Kaua'i, though now possibly extinct on last island.

habitat

Undergrowth of secondary forests and bamboo stands, gardens.

behavior

Found in groups, which, when perched, typically maintain close bodily contact with frequent mutual preening. In winter, moves to lower elevations.

feeding ecology and diet

Groups mostly forage near ground for insects, fruits, seeds, and flowers.

reproductive biology

Apparent group courtship, followed by pairs separating off and setting up territories. Distinctive male territorial song. Both sexes build cup-shaped nest from a variety of plant materials and cocoon silk. Usual clutch three to four greenish white, variously speckled eggs.

conservation status

Not threatened. Massive exploitation for cage bird trade led to listing as CITES Appendix II in 1997. China banned commercial export of songbirds in 2001. Mysterious, brightly colored L. lutea astleyi may represent a population trapped to extinction.

significance to humans

Enormously popular cage bird. Implicated in spreading malaria to native Hawaiian birds.

Black-crowned barwing

Actinodura sodangorum

subfamily

Timaliinae

taxonomy

Actinodura sodangorum Eames, Le, Nguyen, and Eve, 1999, Ngoc Linh, Vietnam. Monotypic.

other common names

French: Actinodure de Vietnam; German: Schwarzkronensibia.

physical characteristics

9.6 in (24 cm). Like the other six members of the genus, a compact-bodied, brownish bird, with a short bushy crest, a long tail, finely barred with black, and edged with white, and primary feathers barred like tail. Unique among genus in possessing a black crown, bold streaking on the throat, and very dark wings. Sexes monomorphic.

distribution

Discovered in 1996 on Mount Ngoc Linh, Kon Tum province, in the western highlands of Vietnam. Since found in six further Kon Tum sites, and on the Laotian Dakchung plateau.

habitat

Montane evergreen and pine forest, as well as grasslands and cultivated land.

behavior

Only observed as individuals or pairs, rarely with other species.

feeding ecology and diet

Gleans insects from foliage, primarily high in trees, sometimes in scrub.

reproductive biology

Monogamous. Pairs often duet antiphonally. Nest undescribed.

conservation status

Vulnerable. Habitat increasingly encroached upon by agriculture, though able to tolerate at least some cultivation.

significance to humans

Target for ecotourists.

Golden-breasted fulvetta

Alcippe chrysotis

subfamily

Timaliinae

taxonomy

Proparus chrysotis Blyth, 1845, Nepal. Five subspecies.

other common names

English: Golden-breasted tit babbler; French: Alcippe à poitrine dorée; German: Gold-Alcippe.

physical characteristics

4.3 in (11 cm).The most brilliantly colored member of a large genus of mostly brown birds. Chest orange or yellow; head black, with gray throat and whitish cheeks and crown. Wings black with brilliant orange and yellow highlights. Tail black, edged with orange or yellow. Mantle grayish and rump yellow. Sexes similar. Shape typical of genus: rounded, like a titmouse or kinglet, with a short, sharp bill, and tail of moderate length.

distribution

Himalayas, east from Nepal, Assam, northeastern Myanmar, southwestern China, and northwestern Vietnam.

habitat

Highland evergreen forest and scrub.

behavior

Member of perpetually moving "bird waves," mixed-species flocks sweeping from one feeding site to another, continuously vocalizing.

feeding ecology and diet

Typical mixed-species flock foliage gleaner, moving from plant to plant on a perpetual search for insects and spiders on bark, twigs, and under leaves. May also eat seeds and fruit.

reproductive biology

Not known.

conservation status

Not threatened, though parts of range subject to deforestation.

significance to humans

A popular target for ecotourists, many of whom come to the Himalayas to momentarily spot as many participants in "bird waves" as possible.

Whiskered yuhina

Yuhina flavicollis

subfamily

Timaliinae

taxonomy

Yuhina flavicollis Hodgson, 1836, central Nepal. Seven subspecies.

other common names

English: Yellow-naped yuhina, yellow-collared ixulus; French: Yuhina à cou roux; German: Gelbnackenyuhina.

physical characteristics

5 in (13 cm). Like other nine members of the genus, a compact, quietly colored bird with small, pointed bill and short, but well-defined, crest. Distinguished by combination of black malar stripe ("moustache") and white streaking on brown flanks. Wings, mantle, and tail dark brown. Chest and throat white. Back of head gray, bordered by orange-brown, then white collars.

distribution

The length of the Himalayas, western China, Myanmar, and northern Indochina.

habitat

Montane evergreen and deciduous forests.

behavior

Usually in large, noisy mixed-species flocks, with birds of similar size. Altitudinal migrant, moving downhill in winter.

feeding ecology and diet

Continuously foraging in interspecies groups for insects, berries, and nectar.

reproductive biology

Monogamous, pairs become territorial during breeding season. Bowl-shaped nest of fine plant materials tied on to branches of shrubs and climbing plants. Two to three speckled eggs, incubated by both male and female.

conservation status

Not threatened.

significance to humans

While considered "uncommon" in China, it is one of the core species in "bird wave" multi-species flocks which attract birders from around the world to the Himalayas.

Fire-tailed myzornis

Myzornis pyrrhoura

subfamily

Timaliinae

taxonomy

Myzornis pyrrhoura Blyth, 1843, Nepal. Monotypic and only member of genus. Position within family remains unclear.

other common names

French: Myzorne queue-de-feu; German: Feuerschwänzchen.

physical characteristics

5 in (12.5 cm). Startling bright green bird with brilliant green, red, and black tail, reddish undertail coverts, black and orange pattern on wings, red wash on chest, scaly crown, red eyes, and slender, black, slightly down-turned bill.

distribution

Himalayas, east from Nepal, and south, along the Salween Divide, into western Sichuan, Yunan, and northeastern Myanmar.

habitat

Rhododendron, bamboo, juniper, and other montane forests, as well as tree farms.

behavior

May be seen singly, in groups of three to four, and in parties of sunbirds, warblers, and other small babblers. Altitudinal migrant; from 9,800–12,000 ft (3,000–3,660 m) in summer, to 6,600 ft (2,000 m) in winter.

feeding ecology and diet

Remarkably similar to unrelated sunbirds (Nectariniidae), with whom it often associates. Continuously probes flowers, especially rhododendrons, becoming covered with pollen, while extracting nectar and insects. Observed by Salim Ali to hover at

flowers like a sunbird, also to run up moss-covered trunks, creeper-like, in search of insects and spiders. Also eats berries.

reproductive biology

Eggs undescribed. Observed feeding chicks at nest in May.

conservation status

Not threatened.

significance to humans

The "Living Emerald" of the Himalayas, a major draw for ecotourists trekking in Nepal and Bhutan.

Crossley's babbler

Mystacornis crossleyi

subfamily

Timaliinae

taxonomy

Bernieria crossleyi Grandidier, 1870, Madagascar. Monotypic. Position within family unclear.

other common names

French: Mystacornis; German: Mystacornis.

physical characteristics

6.3 in (16 cm). Compact-bodied, with relatively long bill. Sexes strongly dimorphic. Male distinguished by black throat and gray crown, while female's white throat and brown crown gives it a markedly different appearance. Both sexes have a broad white malar stripe ("moustache"), far more noticeable in male, as well as black band through eye, and small white spot above eye. Mantle, wings, and tail brown, belly gray, off-white under-tail coverts. Eyes dark.

distribution

Eastern Madagascar.

habitat

Understory of primary evergreen humid forests.

behavior

Found either individually or in family units. Adult males do not tolerate the presence of others, and respond aggressively to others' vocalizations. (Unlike many other babblers, fledged juveniles do not resemble adults, but are instead uniform rufous.)

feeding ecology and diet

Walks around on forest floor, in fairly open places, looking for insects.

reproductive biology

Monogamous. Cup-shaped nest built from twigs, close to the ground.

conservation status

Not threatened, but habitat dependent. Range includes several reserves.

significance to humans

A target species for ecotourists.

Bearded reedling

Panurus biarmicus

subfamily

Paradoxornithinae

taxonomy

Parus biarmicus Linnaeus, 1758, Holstein. Three subspecies. Some classification systems have placed this species and other

parrotbills in separate family (Panuridae or Paradoxornithidae), but Delacour considered them subfamily within babblers. DNA hybridization research suggests they may belong in the same subfamily (Timaliinae) as most babblers.

other common names

English: Bearded tit, marsh pheasant (East Anglia); French: Mesange a moustaches; German: Bartmeise; Spanish: Bigotudo.

physical characteristics

5 in (12.7 cm). Highly sexually dimorphic. Male unmistakable: unique elongated black feathers in front of eye droop down to chest, forming a "moustache" that contrasts strikingly with orange eyes and bill; gray head, white throat, warm brown mantle, rump, and tail, black-white-and-brown-patterned wings, and black vent. Female: long-tailed, compact-bodied bird like male, with similarly patterned wings, and orange eyes and bill, but otherwise generally buffy-brown. Juvenile male very distinct: gray eyes set in black mask, black back, black on tail, and overall golden color. Legs black and unusually long.

distribution

Scattered range in British Isles and western and central Europe, then broadly across central and northeastern Asia to the Pacific (Bokai Sea). Accidental to Japan.

habitat

Wetlands, especially Phragmites reed beds.

behavior

European populations essentially nonmigratory, but inclined to wander in winter, and severe weather may cause wholesale relocations. East Asian populations appear to regularly migrate from interior to coast in winter. Highly gregarious and social outside of breeding season. Complex series of communication calls and displays. Usual flock size 10–20. Noted for skillful negotiation through reed beds, typically grasping a different stalk in each foot. Also capable of what Otto Koenig termed "flutterswimming," should bird fall in water.

feeding ecology and diet

Remarkable seasonal changes in digestive tract. Winter diet largely Phragmites, Typha and rush (Juncus) seeds. Stomach then muscular and hard-walled, and bird ingests numerous small stones. During remainder of year, when diet is mostly insects, spiders, and snails, stomach is smaller, flaccid, and contains no, or fewer, stones. Forages both on the ground and in vegetation.

reproductive biology

Adapted to produce large numbers of offspring quickly, in response to relatively unstable environment. Regularly hatches three broods a year, sometimes four, and young from earliest clutch may themselves breed before end of season, in fall. Pairs mate for life, but nest in loose colonies without territorial boundaries. Nest is a deep cup of reed leaves and other plants, lined with reed flower-heads and feathers, located deep amidst plant stalks, above water, or on land, constructed by both members of pair, who continue to add material throughout 10–14 day incubation period. Four to eight pale, streaked, and speckled eggs incubated by both parents, who also raise chick together.

conservation status

Not threatened, but distinctive, pink-bellied southern Turkish P. biarmicus kosswigi appears to be extinct. In early nineteenth century common along the Thames, from the estuary to Oxford, but British Isles population reduced to two to four pairs in East Anglia by 1948 due to marsh and fen draining, hard winters, and private and commercial egg collectors. Since then British population has increased to more than 500 pairs, at least partially recolonized from continental Europe during hard winter dispersals. German range has expanded since the 1950s. All European populations are closely monitored.

significance to humans

Since the 1990s large numbers were available cheaply from People's Republic of China for the captive-bird trade. Export prohibited by China in 2001, but may continue from Russian Federation.

Vinous-throated parrotbill

Paradoxornis webbianus

subfamily

Paradoxornithinae

taxonomy

Suthora webbiana Gould, 1852, Shanghai. Seven subspecies.

other common names

English: Rufous-headed crowtit; French: Paradoxornis de Webb; German: Papageischnabel.

physical characteristics

5 in (12 cm). Tiny, long-tailed, brownish bird with rufous accents, finch-like yellow bill, and dark eyes. Sexes similar.

distribution

Greater portion of eastern China, north to southeastern Siberia, south to northeastern Myanmar and northwestern Vietnam, as well as Korean Peninsula and Taiwan.

habitat

Summer: undergrowth of mixed conifer and broad-leafed secondary forest. Winter: grassy hillsides, reed beds, thickets, farmlands.

behavior

Some populations migrate south, most only change habitat in winter. Flocks usually contain about 10 birds, may include up to 80. Continuous contact calls. Noted for great agility among dense vegetation.

feeding ecology and diet

Continuously forages through vegetation in flocks for seeds, insects, spiders, and occasionally cultivated grain.

reproductive biology

Monogamous. Male constructs outer framework of deep bowl-shaped nest of grass and various fibers, in small trees, bamboo, dense grass, or thickets. Female assists in lining with moss, cobwebs, hair, etc. Thirteen-day incubation of 3–5 blue-green or white eggs shared by parents.

conservation status

Not threatened. Habitat may have expanded through recent human activity. Recent massive commercial exportation from China ended by ban in 2001, though shipments may arrive from Siberia.

significance to humans

May pose minor threat to millet, sorghum, wheat, and rice. Continuous acrobatic activity has made it a popular traditional cagebird in China and Japan. Before the 1949 Chinese Revolution, males popular for wagered bird fights.

White-necked picathartes

Picathartes gymnocephalus

subfamily

Picathartinae

taxonomy

Corvus gymnocephalus Temminck, 1825, Coast of Guinea. Monotypic.

Classification long controversial. Prior to Delacour's placement as a subfamily within babblers, often considered in crow family (Corvidae). DNA melting curve experiments show no close relation to babblers, but instead suggest Picathartes belongs in the same family with Southern African rock jumpers (Chaetops sp.), with possible relationship to corvids.

other common names

English: White-necked bald crow, yellow-headed rockfowl, Guinea picathartes; French: Picatharte de Guinée; German: Weisshals Stelzenkraehe; Spanish: Picatartes Cuelliblanco.

physical characteristics

16 in (40 cm), 7 oz (200 g). Elegantly proportioned bird of unmistakable appearance. Head unfeathered, with unique black and orange-yellow skin pattern. Eyes and powerful bill dark. Mantle, wings, and tail black, or nearly so. Neck and under-parts creamy white.

distribution

Guinea, south to Ghana.

habitat

Primary and mature secondary forest with boulders or rock formations.

behavior

Keeps to understory, mostly stays in vicinity of nest sites throughout the year. May be found singly, in pairs, or in small groups. Moves about as much by leaping and hopping as by flying. Noted to be inquisitive of humans entering habitat.

feeding ecology and diet

Forages on or near ground for wide variety of insects, as well as other arthropods, snails, earthworms, frogs, and lizards. Follows army ant columns in company of other insectivorous birds, snatching fleeing insects.

reproductive biology

Monogamous. Usually nests in colonies of several pairs. Thick-walled, cup-shaped nests are composed largely of mud, with incorporated plant material, constructed on rock faces (cave mouths, boulders, cliffs), or sometimes stream banks or large fallen trees. Separate roosting and breeding nests thought to be routine. Nests are reused over periods of at least several years. Only one or two blotched, variously colored eggs laid, with rather long incubation period of 23–28 days. Usually two clutches a year.

conservation status

Vulnerable. One of Africa's most well-known threatened birds. As of 2002, main threats are from forest clearance. Prior to 1973 CITES Appendix I listing, Liberian animal dealers appeared to have destroyed entire colonies. Recent fieldwork has located previously unknown populations, such as one in Mont Peko National Park, Ivory Coast, discovered in 1998. A number of protected areas include breeding colonies.

significance to humans

Chicks and adults sometimes taken for food. Once popular zoo bird, as of 2002, none outside Africa since 1998. Future "flagship" and ecotourism potential.

Gray-necked picathartes

Picathartes oreas

subfamily

Picathartinae

taxonomy

Picathartes oreas Reichenow, 1899, Cameroon. Monotypic.

other common names

English: Red-headed rockfowl, blue-headed picathartes, gray-necked bald crow, Cameroon picathartes; French: Picatharte du Cameroun; German: Blaustirn Stelzenkrahe; Spanish: Picatartes Cuelligris.

physical characteristics

14 in (35 cm), 7.7 oz (220 g). Appears obviously related to preceding species, but markedly distinct. Uniquely beautiful blue, black, and red pattern of bare skin on head, blue extending to base of bill. Neck, mantle, back, and tail gray. A patch of black bristles on crown, and short ruff at base of bald head, can be erected when bird is excited. Primaries black; underparts pale yellow.

distribution

Primarily Cameroon, and contiguous Nigeria and Gabon. Recently discovered to occur on the island of Bioko, Gulf of Guinea.

habitat

Closed canopy, undisturbed rainforest, often hilly, always in vicinity of rock formations.

behavior

Found on or near ground, singly, in pairs, or in groups of up to 10. Flocks have been observed bounding along, almost in unison. Shares previous species' curiosity towards humans. Groups gather to roost in communal nesting sites at night.

feeding ecology and diet

Actively hunts, poking through leaf litter, pouncing on prey, primarily arthropods (including crabs), snails, worms, frogs, and lizards. Often follows army ants, and may poke through bat guano.

reproductive biology

Monogamous. Mud and fiber nests similar to that of white-necked picathartes, similarly plastered to rocks, built by both sexes. Construction may take months, sometime a year. Clutch of usually two variously colored speckled eggs incubated by both parents for 24 days.

conservation status

Vulnerable due to dependence on primary forest. Still abundant in various locations, and new populations have been recently discovered. Limited commercial exploitation as zoo birds from 1968 to 1970. CITES Appendix I status, awarded in 1973, prevents further international trade.

significance to humans

Future "flagship" and ecotourism potential.

Resources

Books

Ali, S. The Book of Indian Birds. New York: Oxford University Press, 1996.

Collar, N. J., A. V. Andreev, S. Chan, M. J. Crosby, S. Subramanya, and J. A. Tobias. Threatened Birds of Asia. Barcelona and Cambridge: Lynx Edicions and BirdLife International, 2001.

Cramp, S., C. M. Perrins, and D. J. Brooks, eds. Handbook of the Birds of Europe, the Middle East, and North Africa, Vol. 7. New York: Oxford University Press, 1993.

Fry, C. H., S. Keith, and E. K. Urban, eds. The Birds of Africa, Vol. 6. London: Academic Press, 2000.

Kennedy, R. S., P. C. Gonzales, E. C., Dickinson, H. C. Miranda, and T. H. Fisher. A Guide to the Birds of the Philippines. New York: Oxford University Press, 2000.

MacKinnon, J. R., K. Phillipps, and Fen-qi He. A Field Guide to the Birds of China. New York: Oxford University Press, 2000.

Morris, P., and F. Hawkins. Birds of Madagascar. New Haven and London: Yale University Press, 1998.

Sibley, C. G., and B. L. Monroe. Distribution and Taxonomy of Birds of the World. New Haven and London: Yale University Press, 1990.

Sibley, C. G., and J. E. Ahlquist. Phylogeny and Classification of Birds: A Study in Molecular Evolution. New Haven and London: Yale University Press, 1990.