Asian treefrogs

(Rhacophoridae)

Class Amphibia

Order Anura

Family Rhacophoridae

Thumbnail description
Small to relatively large treefrogs with the two halves of the pectoral girdle fused midventrally and expanded disks on the fingers and toes

Size
0.6–4.9 in (15–120 mm) in snout-vent length

Number of genera, species
13 genera; 341 species

Habitat
Both primary and disturbed forests, agricultural fields, ponds, streams, and savanna

Conservation status
Endangered: 1 species; Vulnerable: 2 species; Lower Risk/Near Threatened: 1 species; Data Deficient: 2 species

Distribution
Southeast Asia from eastern India, Sri Lanka, and Nepal, to Japan, Borneo, Celebes, and the Philippines; also in sub-Saharan Africa and Madagascar

Evolution and systematics

Asian treefrogs are most closely related to the true frogs (Ranidae) as evidenced by abutting epicoracoids in the pectoral girdle (firmisterny), the metasternum forming a bony style, and the presence of the cutaneous pectoris muscle. However, rhacophorids differ from ranids by having an intercalary element between the penultimate and terminal phalanges in the digits.

No fossils are known for the family. Relationships among the frogs currently assigned to Rhacophoridae are problematic. Results of morphological and molecular analyses are controversial. Some workers place rhacophorids as a subfamily of Ranidae; others recognize as many as three separate families, one of which also contains species usually placed in Ranidae. Herein, the 341 recognized species of rhacophorids are placed in 12 genera in three subfamilies, though 13 genera may be recognized.

Buergerinae

Flange on the third metacarpal bone and femoral glands absent; nuptial excrescences present; eggs deposited in water; free-living tadpoles. One genus (Buergeria) with four species in eastern Asia.

Mantellinae

Flange on third metacarpal bone absent; femoral glands present; nuptial excrescences absent; eggs deposited on ground or on vegetation; free-living tadpoles. Two genera (Mantella and Mantidactylus) in Madagascar.

Rhacophorinae

Flange on third metacarpal absent or present; femoral glands absent; nuptial excrescences present; eggs deposited in water, in arboreal cavities, or foam nests; free-living tadpoles or direct development. Nine genera: Chiromantis in sub-Saharan Africa; Aglyptodactylus and Boophis in Madagascar; Chirixalus, Nyctixalus, Philautus, Polypedates, Rhacophorus, and Theloderma in southeastern Asia.

Physical characteristics

Asian treefrogs have enlarged disks on the ends of the fingers and toes that aid in climbing on vertical surfaces and clinging to branches and leaves, and a head that is usually distinct from the body. They have varying degrees of webbing on the toes, and some have extensive webbing on the fingers. Some species of Rhacophorus with extensive webbing also have flaps of skin or fringes along the outside of the limbs and a flattened body. These characters increase their surface area, enabling them to glide or parachute from their higher perches to other trees or the ground when they jump. They have therefore been named "flying frogs." One of the most famous is Wallace's flying frog (Rhacophorus nigropalmatus) of Borneo and Southeast Asia, which can glide up to 24 ft (7.3 m) if it is dropped from a height of 17.7 ft (5.4 m).

Asian treefrogs usually have large eyes with horizontal pupils. The dorsal coloration varies from green, gray, and brown to white and black. Many have spots or irregular blotches on their backs. Some species have flash colors on

their sides and inside their thighs, and sometimes on the webbing between the fingers and toes. Several species have fringes on the forearm from the elbows to the outside of the fourth finger, and projections at the knees and vent. The skin may also vary from smooth to the very bumpy skin of the genus Theloderma. In some species the skin is co-ossified to the skull.

The species of the genus Mantella do not look at all like treefrogs. They are usually not arboreal but instead spend their lives on the forest floor. As a result, most of these diminutive frogs lack one of the most conspicuous characters of this family, the expanded digital disks. Mantellas have toxic skin secretions, and bright dorsal colors similar to the poison frogs (Dendrobatidae) in South America. These colors vary from bright yellow, orange, or red on the dorsum as in the golden mantella (Mantella aurantiaca), to yellow, orange, or red on the limbs with jet black on the dorsum as in Mantella cowanii. Femoral glands are present in males of Mantella and of the highly variable genus Mantidactylus, which are more like treefrogs with expanded disks on the tips of their digits. Because of the presence of femoral glands, the absence of nuptial pads on the males, and the non-amplexing mating behavior, both of these genera are considered closely related. Frogs of the Malagasy genus Aglyptodactylus also lack enlarged disks on the tips of the fingers and toes, which indicates that these frogs are terrestrial or possibly semifossorial.

Distribution

Most Asian treefrogs occur in south, southeast, and east Asia from eastern India, Sri Lanka, and Nepal throughout Myanmar (Burma), Thailand, Laos, Kampuchea (Cambodia), and Vietnam, southeast along the Malay Peninsula onto the islands of Sumatra, Java, Borneo, and Sulawasi, and throughout the Philippines. They also occur in China and Japan, extending into temperate forests north of 40° latitude on the island of Honshu, Japan. Chiromantis, with only three species, is in sub-Saharan Africa, and four genera (Aglyptodactylus, Boophis, Mantella, and Mantidactylus) are endemic to Madagascar.

Habitat

Most Asian treefrogs occur in forests and some (Nyctixalus, Philautus, and Theloderma) usually are not near water. Many species of Chirixalus, Polypedates, and Rhacophorus also inhabit flooded rice fields and grasses or low shrubs between agricultural lands and forests. Many species of the genera Boophis and Buergeria breed in streams, others in ponds, ditches, or other sources of stagnant water. Polypedates leucomystax is abundant in and around human habitation, such as on buildings and in gardens, and even within cities throughout Southeast Asia. Species of Chiromantis inhabit dry areas of the African savanna; they can be found resting on tree limbs exposed to direct sunlight.

Behavior

Most of what is known about the behavior of rhacophorid frogs is related to mating and reproduction, since it is during the mating season that these frogs are more noticeable as males gather and call at breeding sites. In this respect, most rhacophorid frogs are active at night, when the males set up territories around a pond or stream and advertise to females. However, males can even call during the day but usually from hidden retreats. In contrast to this, males of many species of Mantella are usually active during the day, calling and fighting for territories in the open, calling from hidden positions under leaf litter, or foraging for ants, termites, and fruit flies.

Feeding ecology and diet

Adults probably feed primarily on insects, spiders, and other arthropods depending on relative size. Tadpoles of most species graze on algae on the rocks and debris. Tadpoles of Philautus carinensis and Chirixalus eiffingeri have been reported to feed on eggs of other frogs.

Reproductive biology

Most males call at night, or from a hidden retreat during the day. However, males of many species of Mantella are active and call during the day. Most species of Chirixalus, Chiromantis, Polypedates, and Rhacophorus deposit eggs in a foam mass on vegetation over ponds or swamps, and the tadpoles fall into the water below or are washed out of the foam by the next rain. In contrast to the foam-nesters, species of the genera Nyctixalus and Theloderma lay a small number of eggs on the inner walls of water-filled tree holes. The eggs hatch and the tadpoles drop into and develop within the water in the tree hole. Alternatively, the many relatively small frogs of the genus Philautus and some frogs of the genus Mantidactylus lay a small number of eggs on the ground. The embryos develop directly into froglets. All species of the genera Aglyptodactylus, Boophis, and Buergeria lay eggs in stagnant or moving water.

Most rhacophorids have prolonged breeding seasons, but Aglyptodactylus are "explosive breeders" during a few days in temporary ponds. Mantella and Mantidactylus do not engage in amplexus. Instead, there appears to be an abbreviated contact at which time the male induces the female to lay eggs by hormonal stimulation from femoral glands. For example, in Mantidactylus depressiceps, the male and female position themselves on vertical leaves so that the male is over the female and his thighs are touching her shoulder and back. He rubs his thighs against her and she almost immediately begins to deposit eggs on the upper surface of the leaf.

In most species there is no parental care of eggs and tadpoles. However, in some Mantidactylus, males sit on egg masses, apparently guarding them against possible desiccation or predation. In some foam-nesting frogs, females return to the foam nest to add more foam or urinate on the nest probably to prevent desiccation. Female Chirixalus eiffingeri return to tree holes to feed unfertilized eggs to their own tadpoles.

Conservation status

Due to a drastic reduction in populations and habitat, the IUCN lists Philautus schmackeri as Endangered and Nyctixalus spinosus and Mantella aurantiaca as Vulnerable; P. alticola and P. poecilus are listed as Data Deficient. In addition, two species of Philautus are listed in CITES as Near Threatened and Endangered, and M. aurantiaca and N. spinosus are listed as Vulnerable. Several species are endemic to small regions or islands. For example, the genera Aglyptodactylus, Boophis, Mantella, and Mantidactylus are all endemic to the island of Madagascar and are probably highly impacted by deforestation, as are many other animals on that island.

Significance to humans

Most rhacophorid frogs receive little attention from the people of the regions where they live. Most of the frogs are too small to eat, but the legs of some larger Mantidactylus appear in food markets. Because of their striking colors, frogs of the genus Mantella have been captured and sold in the pet trade in the same fashion as poison frogs of South America.

Species accounts

List of Species

Buerger's frog

Buergeria buergeri

subfamily

Buergerinae

taxonomy

Hyla bürgeri Temminck and Schlegel, 1838, Japan.

other common names

English: Kajika frog; Japanese: Kajika-Gaeru.

physical characteristics

A medium-sized treefrog; males are 1.5–1.7 in (37–44 mm), and females are 1.9–2.7 in (49–69 mm) in snout-vent length. The body is slender and dorsoventrally depressed. This frog has a ground color of ash gray to brown with an irregular darker pattern on the back that blends in with the rocks on which it sits. The skin on the back has scattered irregular granules with blunt tips. The legs have a banding pattern but the abdomen is cream to white. The tips of the fingers and toes are expanded into large truncated disks.

distribution

This species is endemic to the mountainous regions of the islands of Honshu, Kyushu, and Shikoku, Japan.

habitat

Usually this species breeds in mid-sized streams with numerous boulders. Outside of the breeding season it has been seen

along forest roads and in trees, and overwintering on river banks under stones and among sand.

behavior

Male territoriality observed during the breeding season.

feeding ecology and diet

Known to feed on rather small insects and spiders.

reproductive biology

This is a prolonged breeder, with males territorial on rocks within riffles of streams and calling both day and night from April into August. Females enter the stream from upland areas throughout the breeding season. Females are quickly amplexed by males when they enter a breeding site. Amplectant pairs may travel up to 130 ft (40 m) to a spawning site. Spawning occurs under rocks in the stream. The egg masses contain 200–600 eggs. The tadpoles with ventrally directed mouths feed on algae among the pebbles and rocks.

conservation status

This species is relatively common throughout its range and is not considered threatened.

significance to humans

The call of this species is a high trill of 10 or more clear notes that is sometimes mistaken for that of a bird. Many hot-spring resorts in Japan take advantage of this beautiful call as a tourist attraction during the spring and summer.

Betsileo golden frog

Mantella betsileo

subfamily

Mantellinae

taxonomy

Dendrobates betsileo Grandidier, 1872, Pays des Betsileos, Madagascar.

other common names

English: Betsileo poison frog.

physical characteristics

This is a small frog of 0.8–1.1 in (20–28 mm) in snout-vent length. The tips of the fingers and toes are not expanded into disks. It has a yellow to orange middorsal color that abruptly turns black dorsolaterally. There is a pale line along the upper lip. The venter is black with irregular blue spotting. The chin is blue. The legs are gray or brown with black bands. The upper half of the iris is golden.

distribution

This species is found from sea level to about 1,640 ft (500 m) in northeastern, western, and southern regions of Madagascar.

habitat

Lowland coastal areas, usually outside of forest.

behavior

Active during the day; males call from exposed areas and fight with other males.

feeding ecology and diet

The diet consists of ants, fruit flies, and small beetles. These frogs actively search for insects during the day.

reproductive biology

The call of the male consists of two short clicks with a short interval between them. Females lay clutches of eggs near streams where the tadpoles wash into pools.

conservation status

Although this species is not threatened, it is exploited in the pet trade.

significance to humans

This frog may be desired as a pet, as are other mantella frogs.

Free Madagascar frog

Mantidactylus liber

subfamily

Mantellinae

taxonomy

Rhacophorus liber Peracca, 1893, Andrangoloaka, Madagascar.

other common names

None known.

physical characteristics

This species is a relatively small treefrog with a snout-vent length of 0.8–1.1 in (21–29 mm) in males and 1.06–1.1 in (27–28 mm) in females. The fingers and toes have expanded disks. There is no webbing between the fingers and only moderate webbing between the toes. The males have either diffuse or large distinct femoral glands. They also possess a large white subgular vocal sac. The smooth dorsum varies between red, gray, or green, with a dark bar between the eyes, a pale median band, and white or yellow spots on the side under the hind legs. The legs and ventrum may be black.

distribution

Eastern and central Madagascar from sea level to 3,900 ft (1,200 m).

habitat

This species is usually found in or around water holding plants within or out of primary forests.

behavior

Males call from vegetation near swamps, pools, and slow-moving water during the rainy season.

feeding ecology and diet

Probably feeds on small insects, spiders, and other arthropods.

reproductive biology

The male's call sounds like two pebbles hitting each other. The female is attracted to a male by his call. She may nudge him from behind, and he places the ventral side of his thighs on her head and shoulders and pulsates laterally. She then begins depositing between 30 and 90 eggs on a leaf overhanging water into which the tadpoles drop after five to seven days of development.

conservation status

Not threatened.

significance to humans

None known.

Forest bright-eyed frog

Boophis erythrodactylus

subfamily

Rhacophorinae

taxonomy

Hyperolius erythrodactylus Guibé, 1953, Forêt de Mahajeby, près de Morafenobe, Ouest de Madagascar.

other common names

None known.

physical characteristics

This is a small, bright green treefrog (snout-vent length of 0.8–1.3 in [20–32 mm]), with many red spots, each surrounded by a yellow ring. The upper eyelids are yellow with brown spots, and there is a yellow line from the snout to the eye. The venter is transparent, the bones are green, and the disks of the fingers and toes are red.

distribution

Eastern Madagascar.

habitat

Usually found on leaves of trees and shrubs near rapids of large streams.

behavior

Not known.

feeding ecology and diet

Probably feeds on insects and other small arthropods.

reproductive biology

Breeds in streams, where the tadpoles live in the rapids.

conservation status

Not threatened, but a limited range in southeastern Madagascar.

significance to humans

None known.

Eiffinger's Asian treefrog

Chirixalus eiffingeri

subfamily

Rhacophorinae

taxonomy

Rana eiffingeri Boettger, 1895, Liukiu [Ryukyu] Islands, either and probably from Okinawa, the middle group, or from Ohoshima, the northern group, Japan.

other common names

English: Big-thumbed treefrog.

physical characteristics

This is a relatively small but stout treefrog, with the males 1.2–1.4 in (31–35 mm) and the females 1.4–1.6 in (36–40 mm) in snout-vent length. The rough skin on the back has scattered small round tubercles and short ridges, is pale brown to dark brown with some black-brown spotting, sometimes with a dark triangle between the eyes, an X-shaped mark on the back, and cross-bands on the legs. The tips of the fingers and toes have expanded round disks.

distribution

Yaeyama Island Group, Japan; Taiwan.

habitat

This species inhabits mountain forests not necessarily near water and occurs in groves of bamboo.

behavior

Males usually call near tree holes or from within cut bamboo that have filled with water.

feeding ecology and diet

Presumably feeds on small insects.

reproductive biology

Males call near tree holes or within cut water-filled bamboo. This species breeds throughout the year in tree holes or cut water-filled bamboo 1.6–4.9 ft (50–150 cm) above the ground. Usually the bottom of the holes is covered with rotting leaves. The eggs are laid separately or in a small mass of about 20–70 eggs above water on the inner walls of the hole. Males often stay in the breeding hole, probably to moisten and, thus, keep the eggs from desiccating. Females spawn continually over several months and periodically return to the breeding site and lay unfertilized eggs directly into the water as food for the tadpoles.

conservation status

This species is not considered threatened or endangered, but it has limited distribution in Taiwan and two small islands out-side of Taiwan.

significance to humans

The practice of cutting bamboo may actually create a breeding habitat for these frogs in areas where tree holes are uncommon.

Gray treefrog

Chiromantis xerampelina

subfamily

Rhacophorinae

taxonomy

Chiromantis xerampelina Peters, 1854, Tette and Sena, Mozambique.

other common names

English: Foam nest frog, southern foam nest treefrog, great African gray treefrog, African gray treefrog; German: Ruderfrosch.

physical characteristics

This species is a relatively large tree frog with a snout-vent length of 2.8 in (72 mm) in males and 3.3 in (85 mm) in females. It is robust with long limbs. The fingers and toes have expanded disks, and the two outer fingers are opposable to the inner fingers, thereby enabling better grasp of the limbs on which it perches. The dorsum is shades of gray and brown with variable darker markings on a roughly textured skin, effectively concealing the frog against different backgrounds, especially the bark of trees. However, the color can change to almost white as temperature rises.

distribution

Savannas of coastal Kenya and northeastern Namibia south to Natal, Republic of South Africa.

habitat

This species usually occurs in warm regions at low elevations and is common in dry savanna.

behavior

This frog and one of the two other species of this genus are unique to rhacophorids in their ability to conserve water so that they can live in the dry African savanna. Individuals orient themselves when resting on tree limbs so that their bodies do not receive full exposure to the sunlight. They tuck the arms and legs under their body, thereby decreasing the amount of surface area exposed to the air and thus reducing evaporative water loss. Also, the frogs turn almost white during the hottest times of the day to reduce heat absorption. While estivating during the dry season, this frog secretes a fluid that turns into a waterproof cocoon. In addition, physiologically this species has been shown to be more tolerant of higher temperatures, and its skin is resistant to water loss of up to 35 times that of other frogs. However, if the temperature becomes too high, this frog will produce drops of water on the skin to cool it down by evaporative cooling. Also, instead of producing urine, as in most frogs, or ammonia, as in highly aquatic frogs, this species, like reptiles, produces the semi-solid uric acid to conserve water.

feeding ecology and diet

Probably feeds on insects and other arthropods.

reproductive biology

This species deposits fertilized eggs as a foam nest. The female may begin building the foam nest, stop, climb down from her perch, and rehydrate in the pool below. At this point the male usually releases her. She then climbs back up and continues building the foam nest. She may repeat this two to four times. The male that originally was on her back may not be the one that is on her back when she releases 500–1,200 eggs into the foam nest. The female may return to the nest the following night and add more foam but not more eggs to the nest to keep it from dehydrating. Communal nests result from up to 20 females and twice as many males building nests close enough together so that they coalesce. Sperm competition is thought to occur in this species (as in many foam-nesting treefrogs with multiple males spawning with one female). This is further confirmed by the presence of large testes in the males that allows them to produce and shed sperm multiple times throughout the breeding season. After four to six days the developing larvae drop from the nest into the water below.

conservation status

Not threatened.

significance to humans

None known.

Painted Indonesian treefrog

Nyctixalus pictus

subfamily

Rhacophorinae

taxonomy

Ixalus pictus Peters, 1871, Sarawak, Malaysia (Borneo).

other common names

English: Cinnamon treefrog, Peter's treefrog.

physical characteristics

This small- to medium-sized treefrog, males 1.12–1.5 in (30–37 mm) and females 1.46–1.54 in (37–39 mm) in snout-vent length, has a relatively long pointed snout and slender limbs. The skin on the back is rough with numerous spiny tubercles, and the skin on the head is co-ossified to the skull. The dorsum is cinnamon to chocolate brown with small white spots scattered throughout, but which also form a broken line

from the edge of the snout, along the edge of the upper eyelid, and continuing partway down the side of the back. Some individuals are red or orange. The upper half of the iris is white, and the lower half is brown. The webbing on the hand is absent or only basal, and the webbing on the foot is moderate.

distribution

Borneo, Malaya, Sumatra, and Palawan Island, Philippines.

habitat

This species is found in both lowland and montane forests from near sea level to 5,400 ft (1,650 m). Adults have been found on leaves of shrubs and small trees one to three meters above the ground but probably are also higher in trees.

behavior

Not known.

feeding ecology and diet

Presumably feeds on small invertebrates.

reproductive biology

This species deposits approximately 10 eggs in a gelatinous mass on the inner walls of water-filled tree holes. The hatch-ling tadpoles drop into the water and feed on the detritus within the tree holes.

conservation status

Not threatened.

significance to humans

None known.

Luzon bubble-nest frog

Philautus surdus

subfamily

Rhacophorinae

taxonomy

Polypedates surdus Peters, 1863, Luzon, Philippines.

other common names

English: Common forest treefrog.

physical characteristics

This small treefrog is 0.9–1.1 in (22–28 mm) in snout-vent length. The tips of the fingers and toes are expanded into disks. There is a pair of tubercles within dark spots at the shoulder and tubercles on the upper eyelids. Webbing is absent between the fingers but moderate between the toes.

distribution

Bohol, Mindanao, and Polillo Islands, Philippines.

habitat

Primary forests from 1,640 to 6,560 ft (500–2,000 m) in elevation, not necessarily near water.

behavior

Not known.

feeding ecology and diet

Probably feeds on small insects and spiders.

reproductive biology

Breeding occurs throughout the year. This species lays five to 19 large, unpigmented eggs in the leaf axils of ferns. The embryos develop directly into froglets. The tail is absorbed just before hatching.

conservation status

Not threatened.

significance to humans

None known.

Kinugasa flying frog

Rhacophorus arboreus

subfamily

Rhacophorinae

taxonomy

Polypedates arboreus Okada and Kawano, 1924, Kinugasa, Kyoto, Honshu, Japan.

other common names

English: Forest green treefrog; Japanese: Mori-ao-gaeru.

physical characteristics

This relatively large treefrog has a large head. Males are 1.7–2.4 in (42–60 mm), and the females are 2.3–3.2 in (59–82 mm) snout-vent length. The dorsum is bright green with or without black or brown spots with black edges. The abdomen is white or cream with pale brown spots. The iris is orange to brownish red. The backs of the thighs are white with black mottling reticulation. The webbing on the hand is well developed, but the webbing on the toes is moderate. The tips of the toes and fingers are expanded into large, truncated disks. The skin on the back is rough with tubercles on the upper eyelid, elbow, and shanks.

distribution

The species is endemic to Honshu, Japan, and the small island of Sado off northeastern Honshu; it occurs from sea level to over 6,560 ft (2,000 m) and is most common in mountainous regions.

habitat

Outside of the breeding season it is found perched in trees or under leaf litter, usually in forested areas but also in urban gardens. During the breeding season it is often seen in trees, grass, and on the ground near ponds and rice fields. During the winter it hibernates under moss or shallow soil.

behavior

This frog is a prolonged breeder, beginning in April and continuing through July. Males set up territories around a breeding site, usually a pond or rice field from where they call.

feeding ecology and diet

This frog feeds on insects.

reproductive biology

The males' call is a series of two to six clicks usually followed by a lower series of clucking sounds. This species deposits eggs in a foam nest on vegetation or the ground over standing water. A female exudes an albumen-based fluid from her cloaca, which she beats with her hind feet into an elliptical foam mass approximately 3.5 × 4.7 in (88 × 120 mm). The amplexing male may also participate by beating the foam with his hind feet. After the foam nest is completed, she deposits 300–800 eggs into the nest, and the male sheds his sperm over the eggs as they leave her cloaca. In some cases the female is surrounded by several males in addition to the male on her back, and all will participate in beating the fluid into a foam mass. These males then shed sperm into the foam mass along with the amplexing male. The foam mass hardens on the outside, protecting the developing embryos from desiccation and predation. After the tadpoles have hatched, the bottom of the foam nest softens by weathering or possibly enzymes released from the hatched eggs, and the tadpoles fall from the foam nest into the standing water below where they develop to metamorphosis.

conservation status

This frog has some protection because of its rarity in a few prefectures in Japan (i.e., Nagano Prefecture).

significance to humans

This species is a predominant part of rural life in Japan, as the chorus of frogs signifies the long summer nights. There are a few ponds where literally hundreds of adult frogs are seen breeding day and night and are therefore set aside as tourist attractions.

Resources

Books

Alcala, A. C., and W. C. Brown. Philippine Amphibians: An Illustrated Field Guide. Makati City, Philippines: Bookmark, Inc., 1998.

Channing, A. Amphibians of Central and Southern Africa. Ithaca, NY: Comstock Publishing Associates, 2001.

Glaw, F., and M. Vences. A Fieldguide to the Amphibians and Reptiles of Madagascar. Frankfurt, Germany: Edition Chimaira, 1999.

Inger, R. F., and R. B. Stuebing. A Field Guide to the Frogs of Borneo. Kota, Indonesia: Natural History Publications, 1997.

Maeda, N., and M. Matsui. Frogs and Toads of Japan. Tokyo, Japan: Bun-Ichi Sogo Shuppan Co., 1990.

Passmore, N. I., and V. C. Carruthers. South African Frogs: A Complete Guide. Johannesburg, South Africa: Witwatersrand University Press, 1995.

Schiøtz, A. Treefrogs of Africa. Frankfurt, Germany: Edition Chimaira, 1999.

Zug, G. R., L. J. Vitt, and J. P. Caldwell. Herpetology: An Introductory Biology of Amphibians and Reptiles. San Diego: Academic Press, 2001.

Periodicals

Bossuyt, F., and M. C. Milinkovitch. "Convergent Adaptive Radiations in Madagascan and Asian Ranid Frogs Reveal Covariation Between Larval and Adult Traits." Proceedings of the National Academy of Science 97 (2000): 6585–6590.

Brown, W. C., and A. C. Alcala. "Philippine Frogs of the Family Rhacophoridae." Proceedings of the California Academy of Sciences 48 (1994): 185–220.

Channing, A. "A Re-evaluation of the Phylogeny of Old World Treefrogs." South African Journal of Zoology 24 (1989): 116–131.

Emerson, S. B., C. Richards, R. C. Drewes, and K. M. Kjer. "On the Relationships Among Ranoid Frogs: A Review of the Evidence." Herpetologica 56 (2000): 209–230.

Glaw, F., M. Vences, and W. Böhme. "Systematic Revision of the Genus Aglyptodactylus Boulenger, 1919 (Amphibia: Ranidae), and Analysis of Its Phylogenetic Relationships to Other Madagascan Ranid Genera (Tomopterna, Boophis, Mantidactylus, and Mantella)." Journal of Zoology, Systematics, and Evolutionary Research 36 (1998): 17–37.

Kaul, R., and V. H. Shoemaker. "Control of Thermoregulatory Evaporation in the Waterproof Treefrog Chiromantis xerampelina." Journal of Comparative Physiology and Biochemistry 158 (1989): 643–649.

Richards, C. M., and W. S. Moore. "A Molecular Phylogenetic Study of the Old World Treefrog Family Rhacophoridae." Herpetological Journal 8 (1998): 41–46.

Richards, C. M., R. A. Nussbaum, and C. J. Raxworthy. "Phylogenetic Relationships Within the Madagascan Boophids and Mantellids As Elucidated by Mitochondrial Ribosomal Genes." African Journal of Herpetology 49 (2000): 23–32.

Wassersug, R. J., K. J. Frogner, and R. F. Inger. "Adaptations for Life in Tree Holes by Rhacophorid Tadpoles from Thailand." Journal of Herpetology 15 (1981): 41–52.

Jeffery Wilkinson, PhD