Asian toadfrogs

(Megophryidae)

Class Amphibia

Order Anura

Family Megophryidae

Thumbnail description
Small to large frogs that are exceptionally well camouflaged and often secretive

Size
0.59–5.51 in (15–140 mm)

Number of genera, species
11 genera; 107 species

Habitat
Forest, mountain streams

Conservation status
Not threatened

Distribution
Eastern Asia

Evolution and systematics

Asian toadfrogs are a group of frogs with diverse morphologic features, and there is not one character that easily defines them. For this reason, using anatomy to form an understanding of the natural groupings within the megophryids or their relationship to other frog families has been difficult. There are no fossil megophryids, and thus the age of this group and its ancient distribution cannot be confirmed. Like other organisms with similar distributions and limitations to dispersal, their current ranges are the result of geologic events that took place more than 30 million years ago. In the Late Oligocene, the sea levels were much lower than they are today, creating a continuous landmass from mainland Asia to the Indo-Australian archipelago, including a portion of the Philippine islands. The tropical rainforest climate at that time was similar to the environments where megophryids are found today, and many of the megophryid genera likely were established and distributed across much of this area. Subsequent fluctuations in sea levels and shifting tectonic plates eroded these land-bridge connections, isolating representatives of Megophrys, Xenophrys, and Leptobrachium on many islands, including Borneo, Sumatra, Java, and the Philippines. The radiation of Scutiger and Oreolalax is intimately tied to the uplifting of the Tibet plateau that followed the collision of the Indian plate with Eurasia some 50 million years ago.

As of the year 2002, the family Megophryidae included 107 species divided among 11 genera and two subfamilies. The arrangement of tubercles on the hand and two distinct types of tadpoles easily distinguish the subfamilies. The subfamily Leptobrachiinae includes Leptobrachella, Leptobrachium, Leptolalax, Oreolalax, Scutiger, and Vibrissaphora. In this group a large tubercle is present at the base of the first finger. The tadpoles of all of these genera typically have a downward oriented mouth and a robust keratinized horny beak on both the upper and lower jaws. The subfamily Megophryinae includes Atympanophrys, Brachytarsophrys, Megophrys, Ophryophryne, and Xenophrys. The tubercle on the hand, at the base of the first finger, extends well onto the thumb. The tadpoles have a large umbelliform (funnel-like) mouth that is directed upward. The keratinized beak is reduced and present only on the margin of the lower jaw.

All Asian toadfrogs have eight vertebrae and intervertebral discs that are not fused to adjacent vertebrae at the time of metamorphosis. The sacral diapophyses are dilated, and the pectoral girdle is arciferal, with a long bony sternum. On the roof of the mouth the neopalatines are absent; to compensate,

a palatal process of the maxilla is elongated. Asian toadfrogs share a common ancestor with North American and European spadefoot toads (Pelobatidae) and parsley frogs (Pelodytidae). Some researchers have recognized these three families as the suborder Pelobatoidea. Asian toadfrogs can be distinguished from their sister groups by their paddle-shaped tongue and a hyoid that is simplified and elongated; the hyoid lacks any remnant of a cartilaginous connection to the back of the skull.

Physical characteristics

Asian toadfrogs come in just about every size and shape imaginable. The largest species, the broad-headed toad (Brachytarsophrys), attains a maximum length of 6.6 in (168 mm), and the smallest species, the Borneo frog (Leptobrachella), is a mere 0.7 in (17.8 mm) long. Females are typically larger than males, except among the moustache toads (Vibrissaphora) and two species of alpine toads (Scutiger). Other sexually dimorphic characters include keratinized nuptial patches on the chest and fingers of breeding male alpine toads and cateyed frogs (Oreolalax) and bizarre keratinized spines seen on the upper lip of the male moustache toad during the breeding season.

The group takes its family name from the genus Megophrys, derived from Greek words (meg + ophrys) that mean "large eyebrow." This refers to the species Megophrys montana, which has long, fleshy appendages above the eyes. This trait is present in most species of Megophryinae, though it is not as pronounced. Leptobrachiinae includes two genera of warty toadlike species, the cateyed frogs and alpine toads. The remainder of the Leptobrachiinae, the slender frogs (Leptolalax), leaf litter frogs (Leptobrachium), and moustache toads, are not as toadlike, but they have fairly large eyes in proportion to their heads. The Borneo frog has unique toe disks that are swollen and have a pointed tip; all other megophryids have simple rounded toes with no visible disks. The pupils are vertical in all genera except the Borneo frog and the mountain toad (Ophryophryne), in which they are either horizontal or diamond-shaped. The color pattern is generally cryptic, but in some species the males may exhibit breeding coloration in the form of bright highlights on the digits or vocal sac, and even fewer have brilliant spots (red, yellow, or orange) along the flanks or thighs.

Distribution

Asian toadfrogs occur throughout Southeast Asia, as far north as Shanxi, China; south to the island of Java (Indonesia); and east from Bengal, India, to Mindanao in the Philippines. The Asian horned frog (Megophrys montana) has been collected at sea level on the beaches of Sarawak, whereas the Nyingchi lazy toad (Scutiger nyingchiensis) has adapted to the harsh climate at 16,732 ft (5,100 m) on the southern slopes of the Himalayas. Of the 11 genera of megophryids, only one genus, Leptobrachella, occurs exclusively on the island of Borneo (Indonesia and Malaysia) or a nearby oceanic island. The most widely distributed genus and the largest, in terms of number of species, is Xenophrys. It occurs throughout much of the range of the family at moderate elevations, between 1,968 and 6,889 ft (600–2,100 m). Leptobrachium and Leptolalax have a more restricted distribution that covers southern China, Indochina, and the island of Borneo; Ophryophryne and Brachytarsophrys occur in Vietnam and southern China. The five species of Vibrissaphora live on only two widely disjunct mountain ranges in southern China and northern Vietnam. Because of their preference for montane habitats, certain species in each genus are unique to a specific mountain range. Of the 107 species of Asian toadfrogs, 26 are known from only a single location.

Habitat

Asian toadfrogs have an extensive north-south distribution that encompasses temperate to tropical rainforest climates. In their northern distribution (approximately 8–35° north latitude), they are found in seasonal monsoon forests where the dry season may last 2–5 months and where the annual rainfall is 106 in (270 cm) a year. In their southern equatorial limit, they occur in aseasonal tropical rainforests in which the annual rainfall may be as much as 263 in (668 cm) a year. Asian toadfrogs require primary or old-growth secondary montane forests. In either case, the forest canopy is generally dense, and the ground is covered with accumulated leaf litter. The tadpoles need clear mountain streams of varying depths. The slender-bodied tadpoles of Leptolalax and Leptobrachella live in swift torrents among the small stones that line the streambed. Larger-bodied tadpoles, such as those of Leptobrachium, Vibrissaphora, Scutiger, and Oreolalax occupy the deeper splash pools, where they are able to avoid the current. The funnel-mouth tadpoles opt for calmer edges of larger streams or are found in the clear, shallow seeps that are scarcely deeper than the tadpoles' bodies.

Behavior

Asian toadfrogs are nocturnal, coming out at dusk to forage and breed. In seasonal climates, their activity is dictated by the wet and dry seasons, but in the tropics they may be active all year long. In the north, breeding takes place during the wet season; Asian toadfrogs may be very abundant at this time. During the dry season it is difficult to find Asian toad-frogs, and it is thought that they return to the forest, taking refuge under rocks and logs. There is one report of five male moustache toads "hibernating" in a tree hole. Asian toadfrogs are poor jumpers; in fact, the leaf litter frog is more inclined to walk slowly away from a disturbance than it is to hop. The cryptic appearance of most species is their only defense against predation, but if they are threatened, the broad-headed toads will open their large mouths and lunge as if to bite.

Feeding ecology and diet

The broad-headed toads are sit-and-wait predators, consuming fairly large prey that may be moving along the forest floor. Little else is known about the foraging activities of the remaining species. Random examination of stomach contents has found that moths, spiders, crickets, cockroaches, beetles, scorpions, centipedes, and snails are all potential prey of Asian toadfrogs.

Funnel-mouth tadpoles feed on minute particles on the surface of the water. While feeding, the larvae position their upturned lips at the level of the water. Taking advantage of the gentle currents that they prefer, they simply allow water and any small particles on the surface film to flow over the edge of the funnel and into the mouth. Papillae (small fleshy fingerlike projections) around the lips direct the food particles into the mouth. When the papillae come in contact with a particle that is too large, the tadpole quickly dives to avoid the obstruction and resurfaces to resume the feeding process. The non–funnel-mouth tadpoles of Leptobrachiinae all forage on the detritus or algae that accumulate in streams. One study has shown that in the same microhabitat, leptobrachine larvae consume food particles that are on average three times the size of what the funnel-mouth tadpoles eat.

Reproductive biology

In seasonal climates breeding activity occurs during the wet season. In Vietnam, this is typically late fall to early spring, and it may last one to two months. In these climates, it is not uncommon to find peak breeding activity when evening temperatures are 41–44°F (5–7°C). Males also may be heard calling during the day, but these efforts are never made with the enthusiasm that is heard at night. Female leaf-litter frogs from Borneo are full of eggs in January, June, July, and August; these equatorial megophryids may breed all year round.

The males of most species situate themselves along the stream bank, either in the vegetation or under the boulders that are at the sides of the stream. The semi-arboreal Ophryophryne calls from vegetation up to 3 ft (1 m) above the ground. Larger species (Megophrys and Brachytarsophrys) space themselves 162–324 ft (50–100 m) along the stream bank. Smaller species, such as Leptolalax, Ophryophryne, and Scutiger, may form aggregations of five to 10 males along a 75-ft (23 m) stretch of a stream. The calls of Xenophrys, Brachytarsophrys, and Leptobrachium

sound like a guttural bark, "grrrack," whereas Ophryophryne produces a series of eight to 10 quick whistles. Leptolalax and Leptobrachella have calls made up of series of quick pulses that sound like a finger dragging across a comb. Amplexus is either axillary, where males clasp the females at the level of the shoulder, or inguinal, where males clasp females around the waist. Alpine toads (Scutiger) engage in inguinal amplexus and are said to use their keratinized chest patches to stimulate the female to lay eggs. The eggs always are deposited on the underside of large boulders at the edges of streams. Parental care has been recorded in the moustache toad—males remain with the clutches until they develop into tadpoles.

Conservation status

Asian toadfrogs vary from being extremely common to exceedingly rare. The slender mud frog (Leptolalax pelodytoides) has a vast distribution and during the breeding season can be the most abundant species in a stream habitat. On the other hand, Sung's slender frog (Leptolalax sungi) is known only from a 50-yd (50 m) stretch of one mountain stream. Efforts are being made to protect the rapidly disappearing habitat in which Asian toadfrogs live. As of the year 2002, no Asian toad-frogs were listed as endangered or threatened by the IUCN.

Significance to humans

During the winter breeding season, when Asian toadfrogs are most abundant, the larger species, such as the Annam broad-headed toad (Brachytarsophrys intermedia), are eaten by local people. The immediate relevance of the smaller species to humans has not been discovered. Given their preference for mature, undisturbed forests and the larval requirement for clean streams, however, they may prove to be a valuable indicator for evaluating the health of Southeast Asian montane forests.

Species accounts

List of Species

Bana leaf litter frog

Leptobrachium banae

subfamily

Leptobrachiinae

taxonomy

Leptobrachium banae Lathrop Murphy, Orlov, and Cuc, 1998, Gia Lai Province, Vietnam.

other common names

None known.

physical characteristics

Males are 2.3–2.9 in (57.2–73.0 mm) long, and females are 3.1–3.3 in (79.9–84.2 mm) long. This heavy-bodied frog has a head that is broad and flat. The limbs are slender and short and seem disproportionately small for the body. The protruding eyes are dark, except for the upper third, which is white. The pupil is vertical. A narrow white membrane is visible around the margin of the eye. From above, the Bana leaf litter frog is uniformly dark brown with red spots on the flanks and hind limbs; the belly is gray with minute white spots.

distribution

This species inhabits the central highlands of Vietnam.

habitat

This species is known only from primary forests at elevations between 2,620 and 3,280 ft (800–1,000 m).

behavior

The Bana leaf litter frog is nocturnal and terrestrial; it spends most of its time taking refuge in the leaf litter deep in the forest.

feeding ecology and diet

The feeding habits are not known, but the diet likely includes a variety of large to medium-size insects.

reproductive biology

Solitary males have been heard calling from burrows or under logs some distance away from the nearest stream.

conservation status

The species is not listed as threatened, but prevailing habitat destruction for coffee plantations is jeopardizing the only known population of this species.

significance to humans

None known.

Slender mud frog

Leptolalax pelodytoides

subfamily

Leptobrachiinae

taxonomy

Leptobrachium pelodytoides Boulenger, 1893, Karin Hills, Burma (Myanmar). This species has the most extensive distribution of any Asian toadfrog and probably represents a complex of species.

other common names

English: Mountain short-legged toad; German: Schlamm-Schlankfrosh; Vietnamese: Cóc mày buèn.

physical characteristics

Males grow up to 1.31 in (33.4 mm) and females to 1.62 in (41.2 mm). An elongate frog, it is orange to light brown, with irregular dark brown mottling on the back and head. The chin and belly are creamy white, and the slender limbs have black transverse bars. The upper lip includes several vertical black bars and one cream-colored vertical bar at the apex of the snout. Small tubercles may be scattered along the back. The tadpole is long and slender and has a subterminal mouth. The body and tail are light brown, and the edges of the tail fin are translucent.

distribution

The slender mud frog ranges across Hong Kong, southern China, Myanmar, Thailand, Vietnam, and Malaysia.

habitat

This species occurs in the vicinity of montane streams but is tolerant of disturbed habitats.

behavior

The frogs are nocturnal and terrestrial.

feeding ecology and diet

The diet is unknown, but it presumably consists of small insects.

reproductive biology

Males call from rocks in and along the edges of streams. Once approached by a female, the male places his chin on the female's shoulder and guides her to a site to lay eggs. Eggs are deposited on the underside of partially submerged rocks.

conservation status

This species is common in montane stream habitats and is not considered threatened.

significance to humans

None known.

Schmidt's lazy toad

Oreolalax schmidti

subfamily

Leptobrachiinae

taxonomy

Scutiger schmidti Liu, 1947, Mount O-mei, Szechwan, China.

other common names

English: Webless toothed toad.

physical characteristics

Males are 1.7–2.0 in (44–52 mm) in length, and females are 1.8–2.1 in (45–54 mm) long. This species has an overall toad-like appearance, including numerous scattered warts on the back and limbs and a thick layer of skin that hides the tympanum. Males have two large nuptial patches on the chest and many keratinized spines on the first finger. The color of the back is grayish brown; the belly and chin are flesh-colored and partially transparent. The short limbs are covered with dark brown transverse bars. The pupil is vertical, and the iris is golden. The thick-bodied tadpole has a subterminal mouth, and the body and tail are ashy-brown with gold and green flecks.

distribution

This species is distributed throughout the Hengduanshan Mountains in southern Szechwan and Yunnan, China.

habitat

The species lives around the headwaters of high mountain streams in desolate high-altitude valleys at elevations of 5,700–7,800 ft (1,740–2,380 m).

behavior

Schmidt's lazy toad is nocturnal, terrestrial, and inactive.

feeding ecology and diet

The diet is unknown.

reproductive biology

The males are extremely persistent callers and will not be deterred even if they are covered with leeches or if someone overturns the stone under which they are calling. Females lay about 120 eggs in balls affixed to the underside of rocks in small mountain streams at a time when the tadpoles from the previous year are on the verge of metamorphosing. Males seem to seek out and court females; individual females may be surrounded by several calling males.

conservation status

Not threatened.

significance to humans

None known.

Ailao moustache toad

Vibrissaphora ailaonica

subfamily

Leptobrachiinae

taxonomy

Vibrissaphora ailaonica Yang Chen, and Ma, 1983, Jingdong County, Yunnan, China. Some authorities include Vibrissaphora in the genus Leptobrachium.

other common names

English: Ailao spiny toad.

physical characteristics

Males are slightly larger than females. Males can grow to 3.2 in (81.6 mm) and females to 3.1 in (78.5 mm). Both sexes have a fine network of ridges over the body and limbs. Keratinized spines develop on the upper lip of the males; females have white spots that correspond to the spines found on the males. The adults are reddish brown with indistinct dark spots; juveniles are light brown with more conspicuous spotting. The pupil is vertical, and the iris is nearly black except for the top third, which is a striking lime green. Tadpoles have a thick body and a subterminal mouth; the body and tail are brown with a light brown line above that bifurcates into a Y where the body joins the tail.

distribution

This species is distributed across the Ailao Shan and Wuliang Shan mountain ranges; a southern extension of the Tibet plateau, in Yunnan province, China; and probably northern Vietnam.

habitat

The Ailao moustache toad prefers closed canopy montane forests at elevations of 7,220–8,200 ft (2,200–2,500 m).

behavior

This species is primarily terrestrial, though males become aquatic during the breeding season.

feeding ecology and diet

Not known.

reproductive biology

The breeding season occurs in late winter and lasts 2–6 weeks. There appears to be a considerable paternal investment; males construct nests underneath large boulders and undergo substantial physical transformations. At the onset of the breeding season, 20–60 keratinized spines develop on the upper lip of the males; the forearms become very thick, and the skin begins to loosen, forming numerous folds on the back and sides of the body. After the female deposits eggs in the nest, the male stays to guard the eggs while attempting to attract additional females. Nesting sites may contain several males and several egg masses. Fertilized eggs take more than a month to develop into tadpoles, and metamorphosis does not take place for two years.

conservation status

Not threatened.

significance to humans

None known.

Annam broad-headed toad

Brachytarsophrys intermedia

subfamily

Megophryinae

taxonomy

Megalophrys intermedius Smith, 1921, Annam, Vietnam.

other common names

English: Annam spadefoot toad; Vietnamese: Cóc mắt trung gian.

physical characteristics

Males grow up to 4.6 in (118.3 mm) and females to 5.5 in (139.5 mm) in length. One of the largest Asian toad frogs, it has a stout body and a broad, flat head. The limbs are short and thick, and webbing is absent on the feet. A fleshy appendage is present above the eyes, and there are irregular folds and ridges on the flanks and back. The color of the back is light brown to reddish brown. The pupil is vertical, and the iris is nearly black. Tadpoles have a funnel-shaped mouth and are brown on the back, with contrasting black and white bars that extend from the belly to the tail.

distribution

The Annam broad-headed toad occupies the central highlands of Vietnam.

habitat

The species prefers montane forest and streams at elevations of 2,460–3,940 ft (750–1,200 m).

behavior

This toad is solitary and inactive and relies on its cryptic appearance to avoid predation. If disturbed, it will not attempt to escape but rather gape its mouth and threaten to bite.

feeding ecology and diet

The Annam broad-headed toad is a sit-and-wait predator that will eat nearly anything that approaches. The diet is known to

include smaller frogs, beetles, crickets, spiders, and moths. Small rodents are a likely prey as well.

reproductive biology

This species is known to breed in late fall and early spring. Males migrate to streams and call from underneath large boulders, where the eggs are deposited.

conservation status

Not threatened.

significance to humans

The Annam broad-headed toad is occasionally used as a food source by local people.

Asian horned frog

Megophrys montana

subfamily

Megophryinae

taxonomy

Megophrys montana Kuhl and Van Hasselt, 1822, Java, Indonesia.

other common names

English: Asian spadefoot toad; Dialect (unspecified): Katak bertanduk, Takang.

physical characteristics

Males grow to 1.7–3.6 in (44–92 mm) in length and females to 2.6–4.4 in (67–111 mm). This is a stocky, large-bodied frog with a bizarre, elongated "horn" on the upper eyelid and, in some forms, a fleshy appendage projecting off the nose. This skin is smooth, except for one or two pairs of fleshy ridges that extend from behind the head to the groin. The color of the back is light brown to reddish brown, occasionally with a few black tubercles. The flanks bear numerous fleshy tubercles and are slightly darker in color than those on the back. The color and overall shape of this species is a perfect imitation of a dried leaf. The pupil is vertical, and the iris is dark brown. Tadpoles have a funnel mouth, and the body and tail are brown.

distribution

The species inhabits Thailand, Malaysia, Sumatra, Java, Natuna, Borneo (Indonesia), and the Philippines.

habitat

This species prefers dense tropical forests from elevations between sea level and 7,220 ft (2,200 m). On rare occasions it is discovered in agricultural areas.

behavior

The Asian horned frog is nocturnal. Its impeccable camouflage makes it extremely difficult to see on the forest floor. If it is discovered, either during the day or at night, it will crouch down further into the leaf litter and wait for the disturbance to go away.

feeding ecology and diet

This species consumes relatively large prey, including cockroaches, scorpions up to 3.9 in (10 cm) in length, and snails with diameters greater than 1.6 in (4 cm).

reproductive biology

During the breeding season these frogs migrate to streams to breed. Males call individually and do not form choruses. The call sounds like a resonant honk.

conservation status

Not threatened.

significance to humans

None known.

Asian mountain toad

Ophryophryne microstoma

subfamily

Megophyrinae

taxonomy

Ophryophryne microstoma Boulenger, 1903, Tonkin, Vietnam.

other common names

English: Narrow-mouthed horned toad.

physical characteristics

Males grow up to 1.43 in (36.2 mm) in length and females up to 1.79 in (45.4 mm). This bizarre toothless frog has a narrow mouth and an extremely truncated snout. Small, pointy tubercles are present above the eye and leaflike venations are on the back. The color of the back ranges from light to dark brown, with some irregular mottling on the head and back. The pupil is diamond-shaped, and the iris is golden brown.

distribution

The Asian mountain toad lives in Vietnam and southwestern China.

habitat

The species prefers montane forests and streams at elevations above 1,300 ft (400 m).

behavior

This nocturnal frog is terrestrial to semi-arboreal. It relies on its camouflage to avoid predation.

feeding ecology and diet

The species feeds on small insects.

reproductive biology

Males typically call from an elevated position above a stream, either in the vegetation or on rocks. The call includes a series of five to 10 quick whistles. Males do not form choruses, but they often are compelled to respond with calls to nearby calling males.

conservation status

This species is not threatened, but its habitat is disappearing quickly. The habitats in Vietnam (populations in the north and in the central highlands) are becoming urbanized or altered for agricultural purposes.

significance to humans

None known.

Burmese spadefoot toad

Xenophrys parva

subfamily

Megophryinae

taxonomy

Xenophrys monticola Günther, 1864, Khasi Hills, India. A study of karyotypes found X. parva to have six large and seven small chromosomes, a pattern shared with other Xenophrys species. Preliminary DNA analyses indicate that the Vietnamese populations are distinct from those in Nepal. Further population sampling across the broad distribution of this species undoubtedly will confirm that this is a complex of species.

other common names

English: Concave-crowned horned toad.

physical characteristics

Males grow to 1.8 in (44.6 mm) and females to 2.0 in (51.0 mm) in length. The width of the body is slightly less than that of the head, and the snout is flat and shieldlike. A distinct ridge extends from the tip of the nose over the tympanum to the shoulder. There are one or two fleshy ridges on the back and a small cone-shaped tubercle is often present above the eye. The top of the head has a dark brown triangle, and there is a similar brown X on the back. The color of the back ranges from reddish brown to golden brown, and the throat and chest are mottled with dark brown. The pupil is vertical, and the iris is golden brown. Tadpoles have a funnel-shaped mouth, and the body is almost uniformly light brown, except for the translucent extremities of the tail fin.

distribution

The species inhabits eastern and central Nepal, northeastern and eastern India, Bangladesh, Myanmar, northern Thailand, northern Vietnam, and southern China.

habitat

This montane frog prefers forests and streams at elevations above 4,950 ft (1,400 m).

behavior

The species is terrestrial and relatively active. It hides in leaf litter by day and relies on its cryptic coloration to avoid predators.

feeding ecology and diet

The Burmese spadefoot toad actively forages at night, feeding on small insects, including crickets, spiders, and moths.

reproductive biology

The species is known to breed in early spring.

conservation status

Not threatened.

significance to humans

None known.

Resources

Books

Bourret, René. Les Batraciens de l'Indochine. Vol. 6. Hanoi: Institut Océanographique de l'Indochine, 1942.

Duellman, William. E., and Linda Trueb. Biology of Amphibians. New York: McGraw-Hill Book Co., 1986.

Inger, Robert, F. "Distribution of Amphibians in Southern Asia and Adjacent Islands." In Patterns of Distribution of Amphibians: A Global Perspective, edited by William Duellman. Baltimore: Johns Hopkins University Press, 1999.

Zhao, Er-Mi. "Distribution Patterns of Amphibians in Temperate Eastern Asia." In Patterns of Distribution of Amphibians: A Global Perspective, edited by William Duellman. Baltimore: Johns Hopkins University Press, 1999.

Zug, George R., Laurie J. Vitt, and Janalee P. Caldwell. Herpetology: An Introductory Biology of Amphibians and Reptiles. 2nd ed. San Diego: Academic Press, 2001.

Periodicals

Dring, Julian. "Frogs of the Genus Leptobrachella (Pelobatidae)." Amphibia-Reptilia 4, no. 2–4 (1983): 89–102.

Dubois, Alain, and Annemarie Ohler. "A New Species of Leptobrachium (Vibrissaphora) from Northern Vietnam, with a Review of the Taxonomy of the Genus Leptobrachium (Pelobatidae, Megophryinae)." Dumerilia 4, no. 1 (1998): 1–32.

Inger, Robert F. "Diets of Tadpoles Living in a Bornean Rain Forest." Alytes 5, no. 4 (1986): 153–164.

Lathrop, Amy. "Taxonomic Review of the Megophryid Frogs (Anura: Pelobatoidea)." Asiatic Herpetological Research 7 (1997): 68–79.

Xie, Feng, and Zhuwang Wang. "Review of the Systematics of Pelobatids (Anura: Pelobatidae)." Cultum Herpetologica Sinica 8 (June 2000): 356–370.

Other

Frost, Darrel R. Amphibian Species of the World: An Online Reference. Vers. 2.20. 1 Sept. 2000 (8 May 2002) <http://research.amnh.org/herpetology/amphibia/index.html>.

Orlov, Nikolai, Roman Khalikov, Robert W. Murphy, and Amy Lathrop. Atlas of Megophryids (Megophryidae: Anura: Amphibia) of Vietnam. Compact Disc. Saint Petersburg: Zoological Institute of Saint Petersburg, 2000.

Amy Lathrop, MA