Leptodactylid frogs

(Leptodactylidae)

Class Amphibia

Order Anura

Family Leptodactylidae

Thumbnail description
Small to large terrestrial and aquatic frogs with arciferal pectoral girdles, usually with teeth on the upper jaw, and no intercalary elements between the penultimate and terminal phalanges of digits

Size
0.4–10 in (10–250 mm)

Number of genera, species
45 genera; 1,124 species

Habitat
Tropical rainforest, temperate rainforest, semiarid grasslands, montane forests, and grassland above tree line

Conservation status
Critically Endangered: 5 species; Endangered: 2 species; Vulnerable: 13 species; Data Deficient: 18 species

Distribution
Leptodactylids occur throughout South America, the West Indies, Central America, and Mexico and also range into the extreme southern United States

Evolution and systematics

Fossils of the subfamily Telmatobiinae are known from Paleocene, Eocene, and Oligocene deposits in Brazil and Argentina. Ceratophryine fossils are known from the Miocene and Pliocene of Argentina, and early Tertiary fossils of Eleutherodactylus are known from the West Indies.

Formerly, Leptodactylidae included the South African Heleophryne, now placed in its own family (Heleophrynidae), and two subfamilies in Australia, now recognized as Limnodynastidae and Myobatrachidae; Limnodynastidae also included Cyclorana, now recognized as a pelodryadine hylid. Most of the features of leptodactylids are primitive for neobatrachians. There is no compelling evidence that the family is monophyletic, and it probably is paraphyletic with respect to several other neotropical families of frogs.

Classification within Leptodactylidae has not been stable; herein seven subfamilies are recognized. Of these, Cycloramphinae, Eleutherodactylinae, and Odontophryinae commonly have been recognized as tribes within Telmatobiinae.

Ceratophryinae

This group consists of medium to large frogs with broad heads, large mouths, robust bodies, and relatively short limbs. The skull is massive and casqued, and the dermal roofing bones are exostosed (with bony outgrowths). The sternum is cartilaginous. The transverse processes on the anterior vertebrae are greatly expanded, and the sacral diapophyses are rounded. The terminal phalanges are knoblike, and dermal glandular pads are absent on the dorsal surfaces of the tips of the digits. The usual karyotype consists of 13 pairs of chromosomes, but some Ceratophrys are polyploids with as many 52 pairs of chromosomes. Pigmented aquatic eggs hatch into carnivorous tadpoles. The subfamily is widely distributed in the tropical lowlands of South America from northern Argentina northward to northern Colombia. It contains 3 genera and 12 species: Ceratophrys (8 species), Chacophrys (1 species), and Lepidobatrachus (3 species).

Cycloramphinae

This group contains small to medium-sized frogs with normal heads and limbs. The skull is not casqued, and the dermal roofing bones are not exostosed. The sternum is

cartilaginous but has an osseous plate in Paratelmatobius. The transverse processes on the anterior vertebrae are short, and the sacral diapophyses are rounded or dilated. The terminal phalanges are knoblike, and dermal glandular pads are absent on the dorsal surfaces of the digital pads. Chromosomes are in 13 pairs. Eggs are deposited in moist situations and hatch as stream-inhabiting tadpoles or as nonfeeding tadpoles that complete their development in terrestrial nests. The subfamily is restricted to southeastern Brazil. It contains 8 genera and 44 species: Crossodactylodes (3 species), Cycloramphus (25 species), Paratelmatobius (6 species), Rupirana (1 species), Scythrodes (1 species), Thoropa (5 species), and Zachaenus (3 species).

Eleutherodactylinae

This group consists of small to medium-sized frogs, mostly with normal heads and limbs. The skull is not casqued, and the dermal roofing bones are not exostosed. The sternum is cartilaginous. The transverse processes on the anterior vertebrae are short, and the sacral diapophyses are rounded. The terminal phalanges are knoblike or T-shaped, and paired dermal glandular pads are absent on the dorsal surfaces of the terminal digits. Chromosomes are in 13 pairs in most genera (9 in Holoaden) but vary from 9 to 17 pairs in Eleutherodactylus. A few large, unpigmented eggs are deposited on land or in bromeliads. Eggs hatch as froglets; there is no aquatic larval stage. At least one species, Eleutherodactylus jasperi, gives birth to living young. The subfamily is widely distributed in South America from northern Argentina northward; Eleutherodactylus also occurs throughout the West Indies, Central America, Mexico, and southern Texas and Florida in the United States. It contains 12 genera and 745 species: Adelophryne (5 species), Atopophrynus (1 species), Barycholos (2 species), Dischidodactylus (2 species), Eleutherodactylus (689 species), Euparkerella (4 species), Geobatrachus (1 species), Holoaden (2 species), Ischnocnema (5 species), Phrynopus (29 species), Phyllonastes (6 species), and Phyzelaphryne (1 species).

Hylodinae

This subfamily contains small to large frogs with normal heads and limbs. The skull is not casqued, and the dermal roofing bones are not exostosed. The sternum is cartilaginous but tends to calcify in old adults. The transverse processes on the anterior vertebrae are short, and the sacral diapophyses are rounded. The terminal phalanges are T-shaped, and a pair of dermal glandular pads is present on the dorsal surfaces of the digital pads. Chromosomes are in 3 pairs. Pigmented eggs deposited in ponds or streams hatch into herbivorous tadpoles with two upper and three lower rows of labial teeth. The subfamily is restricted to southeastern Brazil and extreme northeastern Argentina. It contains 3 genera and 34 species: Crossodactylus (10 species), Hylodes (19 species), and Megaelosia (5 species).

Leptodactylinae

This subfamily consists of small to large frogs with normal heads and limbs. The skull is not casqued, and the dermal roofing bones are not exostosed. The sternum consists of a bony style. The transverse processes on the anterior vertebrae are not expanded, and the sacral diapophyses are rounded or slightly dilated. In most genera, the terminal phalanges are knoblike, but they are T-shaped in Lithodytes, and dermal glandular pads are absent on the dorsal surfaces of the terminal digits. The chromosome complement consists of 10–13 pairs, except for two tetraploid species of Pleurodema that have 22 pairs. Most genera deposit eggs in aquatic foam nests, with tadpoles hatching as free-living herbivrous tadpoles, but eggs are laid in clumps or strings in Limnomedusa, Pseudopaludicola, and some Pleurodema; eggs of Adenomera are in terrestrial foam nests and usually hatch as nonfeeding larvae. The subfamily occurs throughout South America and tropical and subtropical Mesoamerica as far north as southern Texas, United States, and also in the West Indies. It contains 9 genera and 152 species: Adenomera (7 species), Edalorhina (2 species), Hydrolaetare (1 species), Leptodactylus (66 species), Limnomedusa (1 species), Lithodytes (1 species), Physalaemus (41 species), Pleurodema (12 species), and Pseudopaludicola (11 species).

Odontophryinae

This group contains medium-sized frogs with robust bodies and relatively large heads. The skull is not casqued, and the dermal bones are not exostosed except in Proceratophrys. The sternum is cartilaginous. The transverse processes on the anterior vertebrae are not widely expanded, and the sacral diapophyses are rounded or slightly dilated. The terminal phalanges are knoblike, and dermal glandular pads are absent on the dorsal surfaces of the terminal digits. The known chromosome complement is 11 pairs. Eggs are deposited in ponds and hatch into herbivorous tadpoles. The subfamily ranges from eastern Brazil to central Argentina. It contains 3 genera and 27 species: Macrogenioglottus (1 species), Odontophrynus (9 species), and Proceratophrys (17 species).

Telmatobiinae

This subfamily contains the basal leptodactylids that have normal heads and bodies. The skull is not casqued and dermal roofing bones are not exostosed, except in Caudiverbera. The sternum is cartilaginous but tends to calcify in old adults. The transverse processes on the anterior vertebrae are short, and the sacral diapophyses are rounded. The terminal phalanges are knoblike (T-shaped in Batrachyla), and dermal glandular pads are absent on the dorsal surfaces of the terminal digits. Chromosomes are in 13 pairs. Pigmented eggs deposited in water hatch into herbivorous tadpoles. The subfamily is restricted to the temperate forests and Patagonian Region of southern Chile and Argentina but extends northward in the Andes to Ecuador. It contains 11 genera and 92 species: Alsodes (14 species), Atelognathus (8 species), Batrachophrynus (2 species), Batrachyla (5 species), Eupsophus (8 species), Hylorina (1 species), Insuetophrynus (1 species), Somuncuria (1 species), Telmatobius (47 species), and Telmatobufo (3 species).

Physical characteristics

Leptodactylids range in size from minute species of Eleutherodactylus with a snout-vent length of only 0.4 in (10 mm) to large terrestrial species (Ceratophrys aurita) and aquatic species (Telmatobius culeus) with snout-vent lengths of 10 in (250 mm). Body shape varies from robust toadlike species (e.g., Odontophrynus) with extremely large heads (ceratophryines) to dorsoventrally flattened aquatic species (e.g., Atelognathus and some Telmatobius) with loose flaps of skin. Some long-legged, terrestrial species (e.g., some Eleutherodactylus and some Leptodactylus) resemble ranids but lack webbing between the toes. Some other arboreal Eleutherodactylus have expanded digits.

All members of the family have eight separated presacral vertebrae, except that the first and second are fused in Telmatobufo. The two halves of the pectoral girdle overlap midventrally to produce the arciferal condition. Usually the pectoral girdle contains two cartilaginous elements, the sternum and omosternum; in leptodactylines, the sternum has a bony style and in Paratelmatoibius, a bony plate. Maxillary and premaxillary bones usually bear teeth. The terminal phalanges of the digits are knoblike or T-shaped. The skin on the dorsum varies from smooth (with or without longitudinal ridges) to pustular or tubercular. Species of Ceratophrys and Proceratophrys have fleshy eyelid "horns," and Edalorhina and many species of Eleutherodactylus have elongate tubercles on the snout, eyelids, and/or heels. The constricted pupil on the eye is horizontally elliptical in most leptodactylids, but it is vertically elliptical in some telmatobiines (Caudiverbera, Hylorina, and Telmatobufo), leptodactylines (Hydrolaetare and Limnomedusa), and one ceratophryine (Lepidobatrachus).

Dorsally most leptodactylids are varying shades of gray, brown, or dull green, and the venter usually is dull white or cream. However, many species of Eleutherodactylus have pale longitudinal stripes and/or bright flash colors on the flanks or limbs that are not visible when the frog is in a resting position. The striped pattern is most evident in the black Lithodytes lineatus, which also has red spots in the groin and on the thighs.

Most leptodactylid tadpoles have a globular body with a single sinistral spiracle and well-developed caudal fins; the oral disc usually has keratinized jaw sheaths and two anterior and three posterior rows of labial teeth. Tadpoles of Lepidobatrachus have paired spiracles and lack keratinized mouth-parts. Some stream-inhabiting tadpoles (e.g., Cycloramphus and Thoropa) have long, muscular tails with extremely low fins.

Distribution

With the exception of the Atacama Desert, leptodactylids occur throughout South America from the Straits of Magellan northward; they range from sea level to 16,200 ft (5,000m) in the Andes. In so doing, the family contains the southernmost frog in the world (Pleurodema bufonina) and the species reaching the highest elevation in the New World (Pleurodema marmorata). Leptodactylus ranges northward to southern Texas, United States, and on Hispaniola, Puerto Rico, and the Lesser Antilles in the West Indies. Eleutherodactylus occurs throughout the West Indies and Mesoamerica to southwestern United States. The only other genera not confined to South America

are Pleurodema, which extends into Panama, and Physalaemus, which ranges northward into Mexico.

Habitat

Leptodactylids occur wherever moisture is present sometime during the year. Ceratophryines mostly inhabit dry regions, but three species inhabit humid forests. Cycloramphines, hylodines, and eleutherodactylines mostly inhabit humid forests, but some eleutherodactylines occur above the tree line in the Andes; many Eleutherodactylus are arboreal in tropical forests. Leptodactylines occur in semiarid regions as well as humid forests, and three species of Pleurodema exist above tree line in the Andes; Hydrolaetare is aquatic. Odontophryines inhabit humid forests, grasslands, and semiarid regions. Telmatobiines are most diverse in humid temperate forests but also range into semiarid regions; Telmatobius and Batrachophrynus inhabit lakes and streams in the high Andes, and Somuncuria inhabits streams originating from hot springs.

Behavior

Most leptodactylids are nocturnal; daytime retreats are under logs or leaf litter, in burrows, or in bromeliads or other epiphytes. However, hylodines are diurnal in mesic montane environments. Two genera of leptodactylines, Edalorhina and Pseudopaludicola, also are diurnal. Even at high elevations when nighttime temperatures are only slightly above freezing, many species of Eleutherodactylus and Phrynopus and three species of Pleurodema are active at night.

The large, carnivorous Ceratophrys secret themselves in shallow excavations amidst leaf litter with only the tops of their heads visible. During the dry season, Lepidobatrachus burrow into the mud in the bottoms of drying ponds; once underground, they shed successive layers of skin that harden into a cocoon that protects them from desiccation.

Escape behavior in most leptodactylids consists of leaping away from potential predators, but some (e.g., Ceratophrys and Edalorhina) sit still and rely on their cryptic coloration and disruptive outlines to avoid predators; this is accompanied by stretching out the limbs in the cryptically colored Proceratophrys appendiculata. When disturbed, Caudiverbera and some species of Leptodactylus inflate their lungs and thereby increase their size to a potential predator. Many species of Physalaemus and Pleurodema have a pair of large, elevated, and brightly colored glands on the posterior part of the body. These frogs assume a defensive posture by lowering their heads and elevating the posterior part of the body, thereby presenting the glands to the potential predator. These glands have been interpreted as "eyespots" and can be construed by the predator as representing a much larger organism.

Feeding ecology and diet

Most leptodactylids are sit-and-wait predators on small arthropods. But Caudiverbera, Ceratophrys, and large species of Leptodactylus also feed on other vertebrates, including frogs, lizards, and small snakes, birds, and mammals. Some species of Eleutherodactylus feed only on ants, and Physalaemus feeds almost exclusively on termites.

Reproductive biology

Leptodactylids living in seasonal environments and at least some living in continuously humid forests have defined breeding seasons usually associated with the beginning of the rainy season. Those species living in continuously humid environments may breed several times a year.

Males of most species of leptodactylids vocalize to attract females. Calls vary from a single "peep" or series of short notes in various species of Eleutherodactylus to a loud "baaa" in Ceratophrys and a loud "whoorup" in some Leptodactylus. At least some species of Physalaemus and Eleutherodactylus have more complex calls consisting of notes that are territorial and others that are courtship calls. Selection of a mate seems to be mostly by female choice. Once a female approaches a male, he grasps her from above with his hands in her armpits (axillary amplexus), except in the telmatobiine Batrachyla, in which amplexus is around the waist.

All ceratophryines, cycloramphines, odontophryines, odontophryines, and telmatobiines deposit their eggs in water or at the edge of water and have aquatic tadpoles. Clutches vary from a few dozen to hundreds of eggs, mostly depending on the size of the frog. Most leptodactylines construct a foam nest by the pair or with only the male kicking the mucous secreted with the eggs and trapping air bubbles within it. The nests float on water or are constructed in depressions that become inundated shortly after the eggs hatch. The small leptodactyline Adenomera has terrestrial foam nests, and the eggs hatch as nonfeeding larvae that complete their development in the nest. The foam nests develop a sticky exterior and contain moisture within, thereby protecting the eggs from desiccation. These frogs commonly deposit their eggs earlier than sympatric pond-breeders, and therefore in temporary ponds the tadpoles get an early start before potential competitors and predators.

Insofar as known, all eleutherodactylines deposit their eggs in moist situations on the ground or in epiphytic plants. Clutches usually contain fewer than 50 relatively large eggs that undergo direct development, thereby eliminating the aquatic larval stage. One species, Eleutherodactylus jasperi, is known to give birth to one to six living young; fertilization is internal and the eggs are retained in the oviducts.

Developmental time is highly variable. In most species with aquatic eggs, hatching occurs three to five days after deposition and the larval period lasts for four to nine weeks, but in Lepidobatrachus and Odontophrynus eggs hatch within two days and the larval period is only about three weeks. In contrast, in some telmatobiines, which deposit eggs in cold water, ovarian development may require as long as 20 days and the larval period lasts up to two years. In eleutherodactylines, development from time of fertilization to hatching of froglets usually is only three to four weeks, and the development period in Eleutherodactylus jasperi is only 30 days.

Parental care in the form of male attendance of terrestrial or arboreal clutches of eggs is common among Eleutherodactylus in the West Indies, but only a few instances of parental care (by females) are known among Eleutherodactylus on the mainland. In some species of Leptodactylus, the female remains with the foam nest during embryonic development; after hatching the tadpoles remain closely associated with the female. Female Leptodactylus bolivianus have been observed to modify the depth of the pond or to guide the school of tadpoles to deeper water, thereby protecting their tadpoles from possible desiccation. Females of Leptodactylus fallax remain with the foam nest; when the larvae hatch, the females deposit unfertilized eggs in the foam nest and the larvae feed on the eggs.

Conservation status

Many leptodactylids are threatened by habitat destruction. Possibly several species are extinct, including the large Eleutherodactylus karlschmidti and the live-bearing Eleutherodactylus jasperi in Puerto Rico. The 2002 IUCN Red List includes 36 species: 5 are categorized as Critically Endangered; 2 as Endangered; 13 as Vulnerable; and 18 as Data Deficient.

Significance to humans

Several large leptodactylids (Caudiverbera, Batrachophrynus, Leptodactylus, and Telmatobius) are consumed by humans. In the Andes of Peru and Bolivia, Telmatobius culeus are captured by "raneros" in Lago Titicaca, and Batrachophrynus macrostomus are likewise taken from Lago Junín. Restaurants in villages near these lakes commonly advertise that frogs are on the menu. Ceratophrys have become popular in the pet trade and are bred in captivity for this purpose. The Puerto Rican Eleutherodactylus johnstonei has been introduced intentionally into Colombia and Venezuela because people who vacationed in Puerto Rico were enamored by the call. In contrast, the unintentional introduction of Eleutherodactylus coqui in Hawaii has caused distress among inhabitants—they are unused to nocturnal vocalization, because there are no frogs native to Hawaii.

Species accounts

List of Species

Surinam horned frog

Ceratophrys cornuta

subfamily

Ceratophryinae

taxonomy

Rana cornuta Linnaeus, 1758, "Virginia" (in error).

other common names

English: Horned frog, packman frog.

physical characteristics

This large, robust frog has an immense head, the width of which is about one-half of the snout-vent length, which is as great as 3.1 in (80 mm) in males and 4.7 in (120 mm) in females. The skin on the dorsum and flanks is finely rugose with conical tubercles, and the venter is nearly smooth. A distinguishing feature is the presence of a large, triangular, dermal process ("eyelid horn") extending upward on each eyelid. The fingers are unwebbed, and the hind limbs are moderately short with toes that are about one-half webbed. The dorsum is green or brown with brown markings, and the venter is dull cream except for a dark brown or black throat. The iris is creamy tan with brown flecks. Breeding males have tan nuptial excrescences on the thumbs.

distribution

This species is widely distributed in the Amazon Basin and Guianan Region in South America.

habitat

This frog is a denizen of lowland tropical rainforest.

behavior

Using its cryptic color pattern as camouflage, this frogs wriggles into the leaf litter on the forest floor, so that only the head is exposed. Individuals may remain in the same place for several days and nights before moving to another site on rainy nights.

feeding ecology and diet

A classic sit-and-wait predator, Ceratophrys cornuta apparently will eat anything that moves by it and is not too large to swallow. It makes a short lunge at its prey, which consists of ants, spiders, and other small arthropods, but the bulk of its prey are large grasshoppers, frogs, and even snakes, lizards, and mice.

reproductive biology

This is an explosive breeder at the time of the first heavy rains of the rainy season. Males call from the edges of ponds or while sitting in shallow water; the call is a low-pitched "baaa." Amplexus is axillary, and clutches of up to 2,000 small, pigmented eggs are deposited in water. Tadpoles attain a total length of about 2.5 in (65 mm). The body is broadly ovoid with a bluntly rounded snout and small eyes directed dorsolaterally. The oral disc is large and directed anteriorly. The jaw sheaths are massive; a long, pointed median process on the lower sheath inserts into a notch on the upper sheath; there are 13 rows of labial teeth on the upper lip and eight rows on the lower lip. The tadpoles are voracious carnivores and feed on other tadpoles in the pond and even are cannibalistic. Feeding is a gape-and-suck process, during which the prey is punctured by the process on the lower jaw sheath and quickly ingested; attack and swallowing takes only about five seconds.

conservation status

Although Ceratophrys cornuta is locally abundant throughout its range, clearing of forest is restricting its habitat. It is not listed by the IUCN.

significance to humans

This species is found in the pet trade.

Budgett's frog

Lepidobatrachus laevis

subfamily

Ceratophryinae

taxonomy

Lepidobatrachus laevis Budgett, 1899, Paraguayan Chaco, South America.

other common names

Guaraní: Kururú chiní.

physical characteristics

Adults of this broad-headed frog with a flattened body attain snout-vent lengths of 4.5–5.1 in (110–130 mm). The eyes are small and close together on the top of the head. The snout is broad and sloping. The fingers are unwebbed, and the toes are nearly fully webbed. A large, spade-like, black inner metatarsal tubercle is present on the base of each hind foot. The dorsal skin is glandular, and the skin on the venter is granular. The dorsum is dull brown or gray with faintly darker blotches or paler streaks; the belly is white. The iris is pale cream, and the pupil is round.

distribution

Budgett's frog occurs only in the dry Chaco Region in northern Argentina and southern Paraguay.

habitat

This species inhabits dry scrub forest.

behavior

This frog is active only during the short rainy season, November through January, when individuals swim in temporary ponds. As the ponds dry up toward the end of the rainy season, the frogs burrow backwards, using the spade-like tubercles on the hind feet, deep in the mud in the bottoms of ponds. Once below the surface, they shed the outer layers of skin several times; this skin forms an impermeable cocoon that protects the frog from desiccation during the long dry season. With the advent of following rainy season, moisture softens the cocoon, and the frogs emerge into the water, eat the shed skin, and begin a new season of activity. Budgett's frog is aggressive and opens its large mouth as a defensive posture.

feeding ecology and diet

Apparently most feeding takes place in the water. These frogs eat snails and smaller frogs; in captivity they also will eat fish.

reproductive biology

Males call while floating on the water; the call is a loud "eeee." Amplexus is axillary. As many as 1,200 small pigmented eggs are laid in water, and these sink to the bottom, where they hatch in about 18 hours. The tadpoles, which are carnivores, metamorphose about 20 days after hatching. The tadpoles, which reach a total length of about 2 in (50 mm), have broad, depressed bodies, paired spiracles, and large mouths with weak labial teeth and no horny jaw sheaths. The tadpoles feed on smaller tadpoles, which they swallow whole.

conservation status

Populations seem to be stable, and the species is not listed by the IUCN.

significance to humans

None known.

Rock river frog

Thoropa miliaris

subfamily

Cyclorampinae

taxonomy

Rana miliaris Spix, 1824, "Amazon River" (in error).

other common names

None known.

physical characteristics

Males attain a maximum snout-vent length of 2.8 in (71 mm) and females, 3.2 in (81 mm). The head is broad with a rounded snout and large, distinct tympanum. The skin of the dorsum is smooth to weakly granular with scattered tubercles; the venter is smooth. The fingers and toes lack webbing and have slightly swollen tips. Breeding males lack vocal slits but have small nuptial spines on the thumb and first and second fingers. The dorsum is tan or brown, and the groin is dull yellow; the throat and belly are gray and the anterior and posterior surfaces of the thighs are dull yellow with dark brown bars. The iris is reddish copper with black reticulations.

distribution

Thoropa miliaris ranges in the Atlantic Coast Forest from Espírito Santa to São Paulo in southeastern Brazil.

habitat

This species inhabits humid tropical and subtropical forests.

behavior

This species is nocturnal and terrestrial and is most common along streams.

feeding ecology and diet

Presumably the diet includes small arthropods.

reproductive biology

Males call from rock faces along streams; the call is a short, pulsed, low-pitched note. Eggs are deposited in streams. Tadpoles wriggle onto wet rocks faces. They have depressed bodies, long and muscular tails without noticeable fins, and ventrally directed oral discs with slender jaw sheaths and two anterior and three posterior rows of labial teeth.

conservation status

Although not listed by the IUCN, this species is threatened by habitat destruction.

significance to humans

None known.

Puerto Rican coqui

Eleutherodactylus coqui

subfamily

Eleutherodactylinae

taxonomy

Eleutherodactylus coqui Thomas, 1966, 7.3 mi (11.8 km) south of Palmer, Puerto Rico.

other common names

Spanish: Coquí.

physical characteristics

Males attain a snout-vent length of 2 in (50 mm) and females, 2.5 in (63 mm). The dorsum is shagreen with scattered small tubercles, and the venter is areolate. The snout is subacuninate, and the tympanum is distinct. The fingers and toes are long, unwebbed, and bear terminal, expanded, truncate discs. The dorsum is various shades of brown, commonly with a middorsal or pair of dorsolateral creamy tan stripes. A distinct dark brown bar extends from the nostrils through the reddish bronze eye to a point above the tympanum. The venter is grayish white.

distribution

This frog occurs throughout Puerto Rico to elevations of 3,900 ft (1,200 m). It has been introduced on St. Thomas and St. Croix in the U.S. Virgin Islands, and into southern Florida, Louisiana, and Hawaii, United States.

habitat

The Puerto Rican coqui lives in nearly all regions of Puerto Rico; it inhabits humid montane forest, dry forest, gardens, and houses.

behavior

This strictly nocturnal species takes refuge under objects, in axils of palms, and especially in bromeliads. At night it is active on the ground but usually on vegetation to heights of more than 50 ft (15 m). Individuals seldom move more than 20 ft (6.5 m) from their diurnal retreats, and when feeding at night they move no more than about 2 in (50 mm). Males establish territories by vocalization and are aggressive toward other males that enter their territories.

feeding ecology and diet

Feeding occurs on vegetation at night; the frogs consume vast quantities of insects, principally ants, crickets, and roaches, as well as spiders, snails, and even small frogs.

reproductive biology

Breeding occurs throughout the year but is reduced in the driest times of the year (January through March). Males call at night. The call is a multiple note, "co-qui." The "co" solicits response from females, whereas the "qui" is a territorial call, which is repeated rapidly upon the intrusion of another male. Amplexus involves the male sitting on the body of the female with his arms around her body; fertilization is internal via cloacal apposition. Clutches of about 26 eggs are deposited on leaves of bromeliads or other plants. The female abandons the eggs, which are attended by the male, who commonly places his body over the eggs. Development is direct into a froglet within the egg capsule and requires 17–26 days. Late embryos develop a tubercle on the tip of the snout ("egg-tooth") that is used to rip open the capsule. Hatchings are about 0.23 in (6 mm) long. The frogs reach sexual maturity in less than one year and have a life span of four to five years. Females can breed as often as every 58 days.

conservation status

This ubiquitous species is common throughout its range.

significance to humans

Exportation of plants, especially bromeliads, from Puerto Rico has resulted in the accidental introduction of the Puerto Rican coqui on the U.S. Virgin Islands, into Florida and Louisiana, and Hawaii, where there are no native frogs. People in Hawaii complain about the nocturnal "noise" made by the coqui. Because of its abundance and ease for study in Puerto Rico, this species has been investigated more thoroughly than any other tropical anuran.

Golden coqui

Eleutherodactylus jasperi

subfamily

Eleutherodactylinae

taxonomy

Eleutherodactylus jasperi Drewry and Jones, 1976, 3.7 mi (6 km) southeast of Cayey, Puerto Rico.

other common names

Spanish: Coquí dorado.

physical characteristics

This small frog attains a maximum snout-vent length of 8.5 in (21.5 mm). The dorsum is shagreen, and the venter is areolate. The snout is bluntly rounded and nearly truncate in dorsal view; the tympanum is about one-half of the diameter of the eye. The fingers and toes are moderately long, unwebbed, and have rounded terminal discs. The dorsum is golden yellow to orange yellow, and the venter is pale yellow, except that the skin covering the abdomen is transparent. The iris is pale gray with black flecks.

distribution

This species has been known only from elevations of 2,100–2,750 ft (650–850 m) in the Sierra de Cayey, Puerto Rico.

habitat

This strictly nocturnal frog inhabits arboreal bromeliads in subhumid forest.

behavior

This small nocturnal species seeks shelter in bromeliads by day.

feeding ecology and diet

Presumably the diet includes small arthropods.

reproductive biology

Males call from bromeliads at night; the call consists of a series of six to eight notes, "tuit-tuit-tuit-tuit." Eleutherodactylus jasperi is the only member of the family that is known to have internal fertilization and give birth to living young. The species is ovoviparous, in that the eggs are retained in the oviduct and the yolk within the egg capsule supplies all nutrition. The eggs are up to 0.2 in (5 mm) in diameter and require about 30 days to develop into froglets. The number of young is three to five; upon birth they are 0.3 in (7 mm) long and contain a large amount of yolk in the abdomen.

conservation status

Although listed as Data Deficient by the IUCN, this species is presumed to be extinct; it was last observed in 1981.

significance to humans

Because of its nearly unique reproductive mode, this small frog was of immense interest to biologists, but only limited data were obtained before it disappeared.

Warty tree toad

Hylodes asper

subfamily

Hylodinae

taxonomy

Elosia aspera Müller, 1924, Barreria, Rio de Janeiro, Brazil.

other common names

None known.

physical characteristics

Males attain a maximum snout-vent length of 1.7 in (43 mm) and females, 2 in (50 mm). The snout is rounded, and a tympanum is present but not always distinct. The fingers are unwebbed but fringed, and the toes are unwebbed; terminal

segments of all digits are expanded, truncate, and have a pair of scutes on the dorsal surfaces. Males have vocal sacs that are expanded laterally but lack nuptial excrescences. The dorsum is dull brown with irregular darker brown to black markings, but the upper surfaces of the truncate digits are white. The upper lip is white with narrow brown bars. The venter is pale tan with darker mottling or spots, and the iris is pale bronze.

distribution

This species is distributed in the coastal mountain ranges from Rio de Janeiro to Santa Catarina in southeastern Brazil.

habitat

Hylodes asper inhabits humid montane forest.

behavior

This diurnal terrestrial species is most commonly seen on rocks and low vegetation along mountain streams. Calling males also display by waving their hind feet one at a time.

feeding ecology and diet

Presumably the diet consists of small arthropods.

reproductive biology

Males call by day from rocks at the edges of streams. The call is a long high-pitched whistling trill. Amplexus is axillary, and eggs are deposited in water, where they hatch into herbivorous tadpoles. The tadpoles have rather slender bodies and long tails with moderately low fins; the oral disc is directed ventrally and has heavy, coarsely serrate jaw sheaths and two anterior and three posterior rows of labial teeth.

conservation status

Although not listed by the IUCN, this species is threatened by the great reduction in habitat that also affects other inhabitants of the Atlantic Coastal Forest in southeastern Brazil.

significance to humans

None known.

Perez's snouted frog

Edalorhina perezi

subfamily

Leptodactylinae

taxonomy

Edalorhina perezi Jiménez de la Espada, 1870, Napo, Ecuador.

other common names

None known.

physical characteristics

Males of this small frog attain a snout-vent length of 1.2 in (30 mm) and females, 1.4 in (35 mm). The snout is short and truncate; prominent, pointed tubercles are present on the upper eyelid, and a distinct dorsolateral fold extends from the orbit to the groin. The dorsum may be tuberculate, smooth with a few scat-tered tubercles, or having several longitudinal ridges between the dorsolateral folds; the venter is smooth. The dorsum is gray or brown with or without reddish brown streaks, the flanks are black, and the venter is white with extensive black markings. The iris is grayish tan with a reddish copper ring around the pupil.

distribution

This species is distributed in the upper Amazon Basin from southern Colombia to northern Bolivia.

habitat

Edalorhina perezi inhabits lowland tropical rainforest.

behavior

This diurnal species is active on the forest floor, where its cryptic coloration blends well with the leaf litter.

feeding ecology and diet

A great variety of small arthropods, including spiders, flies, crickets, and roaches are eaten while the frogs forage in the leaf litter.

reproductive biology

Males call solitarily from the leaf litter by day; the call consists of three to five low whistles with two pulses per note. Amplexus is axillary, and the pair moves to a small body of water, usually temporary ponds, where 78–98 eggs are deposited in a foam nest constructed by the pair kicking the eggs, secretions, and water into a small, spherical mound that floats on the surface of the water. The eggs hatch in four to six days, and the tadpoles develop in water. Tadpoles attain a maximum total length of about 0.8 in (20 mm). The body is ovoid with a bluntly rounded snout and dorsally positioned eyes. The oral disc is directed anteroventrally; the jaw sheaths are finely serrate, and there are two rows of labial teeth on the anterior lip and three on the posterior lip. The body and caudal musculature are tan, and the belly is greenish yellow.

conservation status

Not listed by the IUCN. However, as in the case of all inhabitants of the Amazonian rainforest, the continuous range of this species is being fragmented by clearing of the forest.

significance to humans

None known.

South American bullfrog

Leptodactylus pentadactylus

subfamily

Leptodactylinae

taxonomy

Rana pentadactyla Laurenti, 1768, "Indiis" (Surinam).

other common names

None known.

physical characteristics

Males of this large, robust frog are slightly larger than females; they attain a maximum snout-vent length of 7.3 in (180 mm), whereas the maximum length in females is 6.9 in (176 mm). The body is robust; the head is large with an acutely rounded snout and prominent tympanum. The skin on the dorsum and venter is smooth, and a prominent dorsolateral dermal fold extends from the orbit to the groin. The fingers and toes are long with slender tips and lack webbing. Breeding males have greatly swollen forelimbs and one large, pointed, black spine on the inner surface of the thumb and two black spines on each side of the chest. The dorsum is tan to reddish brown with broad, reddish brown marks on the body between the yellowish tan dorsolateral folds. The dorsal surfaces of the limbs are tan to reddish brown with narrow transverse brown bars. The upper lip is tan with a brown margin and dark brown triangular spots. The venter is cream with bold dark brown to black mottling, especially on the belly and hind limbs. The iris is bronze.

distribution

This frog ranges in lowlands (below 3,800 ft or 1,200 m) from northern Honduras to the Pacific lowlands of Ecuador and throughout the Guianas and northern two-thirds of the Amazon Basin in South America.

habitat

Principally a denizen of tropical rainforest, this species also invades dry forest and lower montane forests.

behavior

This nocturnal species spends its days in burrows, under logs, or hidden in leaf litter. Defensive mechanisms include noxious skin secretions and posturing by inflating the lungs and elevating the body on all four limbs. When grasped, these frogs usually emit a high-pitched scream.

feeding ecology and diet

Juveniles feed on small arthropods, but large adults feed on large arthropods, frogs, lizards, snakes, and small birds and mammals. Tadpoles are omnivorous, feeding on vegetation, tadpoles, and eggs, even of their own species.

reproductive biology

Males call solitarily from margins of ponds and backwaters of streams; the call is a loud "whoorup" repeated at intervals of five to 10 seconds. An attracted female is grasped by the male by axillary amplexus and held firmly by the muscular forearms and nuptial spines on the thumbs and chest. About 1,000 eggs are deposited in a large foam nest by backward and forward motions of the male's hind limbs that mix air, water, eggs, and secretions into the nest, which usually is deposited in a depression adjacent to water. The eggs hatch in two to three days; subsequent rains flood the nest site, and the tadpoles move into the pond or slow-moving stream. Development is rapid, and metamorphosis occurs about four weeks after hatching. Tadpoles attain a maximum total length of about 3.3 in (83 mm). The body is ovoid with a rounded snout with large eyes directed dorsolaterally. The oral disc is nearly terminal and bears finely serrate jaw sheaths and two anterior and three posterior rows of labial teeth. The body and caudal musculature are brown.

conservation status

Populations of this species seem to be stable. It is not listed by the IUCN.

significance to humans

Some indigenous people eat these frogs, after they have been washed thoroughly.

Gold-striped frog

Lithodytes lineatus

subfamily

Leptodactylinae

taxonomy

Rana lineata Schneider, 1799, Guyana.

other common names

None known.

physical characteristics

Males attain a maximum snout-vent length of 1.8 in (45 mm) and females, 2.2 in (56 mm). The body is slender; the snout is rounded, and a distinct tympanum is present. The fingers and toes are unwebbed and slender with slightly dilated tips. The skin on the dorsum is finely spiculate, and the venter is smooth. The dorsum and flanks are black; a pair of broad yellow stripes extends from the tip of the snout to the groin. A large red spot is present in the groin, and a smaller red spot is present on the posterior surface of each thigh. The throat and chest are grayish brown, and the undersurfaces of the hind limbs are gray. The iris is coppery bronze. Males lack nuptial excrescences.

distribution

The species is widely distributed in the upper and middle Amazon Basin and in the Guianan region in northeastern South America.

habitat

This species is restricted to humid tropical lowland rainforest.

behavior

Juveniles are active on the ground by day and night, whereas adults are primarily nocturnal. Adults have been found in association with the large earthen nest of leaf-cutting ants (Atta), and males are known to call from subterranean tunnels in these nests.

feeding ecology and diet

This species feeds on a variety of small arthropods and also earthworms on the forest floor.

reproductive biology

Males call from mouths of burrows or other partially concealed sites; the call is a series of melodious notes. About 200 unpigmented eggs are deposited in a foam nest constructed at the edge of water. The tadpoles remain in the foam nest for seven to 15 days after hatching and then disperse into the water. Tadpoles metamorphose about nine weeks after hatching. Tadpoles attain a maximum total length of about 2 in (50 mm). The body is elongately ovoid with a truncate snout and dor-sally situated eyes. The oral disc is directed anteroventrally and bears slender, finely serrate jaw sheaths and up to two rows of labial denticles on the anterior lip and up to three rows on the posterior lip. The tadpoles are bright pink with a short middorsal white stripe.

conservation status

Clearing of rainforest threatens to limit the distribution of this species. It is not listed by the IUCN.

significance to humans

Lithodytes is not known to have toxic skin secretions like the poison frogs of the family Dendrobatidae; however, the color pattern of Lithodytes closely resembles that of the sympatric poison frog Epipedobates femoralis and thus may be a case of mimicry.

Túngara frog

Physalaemus pustulosus

subfamily

Leptodactylinae

taxonomy

Paludicola pustulosa Cope, 1864, New Grenada and Truando River (Colombia).

other common names

None known.

physical characteristics

This small, toadlike anuran attains a maximum snout-vent length of 1.4 in (35 mm). The head is relatively small with a subacumi-nate snout and no distinct tympanum. The dorsum is tuberculate, and the venter is smooth. A well-defined elongate gland is present on the flank. The first finger is longer than the second, and the fingers and toes lack webbing. The dorsum is dull brown with or without irregular darker brown spots; the venter is grayish white with black spots, and the iris is tan with black flecks. Breeding males have brown nuptial excrescences on the thumbs.

distribution

The species is widely distributed in the lowlands of southern Mexico, Central America, northern Colombia, the coastal region and llanos of Venezuela eastward to Guyana, and the islands of Trinidad and Tobago.

habitat

Physalaemus inhabits grasslands, pastures, and open forest formations.

behavior

This small species is nocturnal and active only in the rainy season. During the day they are hidden in leaf litter or under objects; in the dry season they burrow in the ground and may remain inactive for many months.

feeding ecology and diet

The diet consists of a variety of small arthropods.

reproductive biology

Breeding takes place primarily at the beginning of the rainy season when males congregate in small bodies of water, even puddles in roads, and commence calling while floating on the surface of the water. The call consists of a whine followed or not by one or more short notes, "chuck." Females swim to males and preferentially select males with low-pitched "chucks." Amplexus is axillary; as the eggs are extruded, they are kicked into a foam nest by the feet of both individuals. Foam nests float on the water and contain 80–450 small eggs that hatch in two to three days into tiny larvae, which may remain in or under the foam nest for up to five days if the water level has dropped. Tadpoles grow to a length of 0.8 in (20 mm) and metamorphose in five to nine weeks. The larval body is ovoid; the eyes are directed dorsolaterally, and the caudal fins are shallow. The oral disc is directed anteroventrally; the jaw sheaths are moderately massive, and there are two anterior and three posterior rows of labial teeth. In the laboratory, the frogs reach breeding condition two to three months after metamorphosis.

conservation status

Inasmuch as this small frog is not an inhabitant of dense forest, clearing of forests probably has enhanced its abundance and distribution. It is not listed by the IUCN.

significance to humans

Physalaemus pustulosus has contributed to knowledge of amphibian biology by being the object of studies on vocal communication, sexual selection, and avoidance of predation.

Gray four-eyed frog

Pleurodema bufonina

subfamily

Leptodactylinae

taxonomy

Pleurodema bufonina Bell, 1843, Puerto Deseado and Río Santa Cruz, Patagonia, Argentina.

other common names

None known.

physical characteristics

Males of this toadlike species attain a snout-vent length of 1.8 in (45 mm), and females reach 2.2 in (56 mm). The skin on the dorsum is shagreen and glandular; the belly is smooth. The snout is bluntly rounded, and a distinct but small tympanum is present. The fingers are unwebbed, and the toes are basally webbed. A distinct feature is the pair of large, ovoid, lumbar glands that are about one-third of the length of the body. The dorsum is dull brown with or without darker brown spots and/or a tan middorsal stripe; the venter is creamy tan. The iris is pale bronze with black flecks.

distribution

This is the southernmost frog in the world. Its distribution extends from the Straits of Magellan northward to 36° south latitude in Patagonian Argentina and adjacent Chile; the elevational range is from sea level to 7,500 ft (2,300 m).

habitat

This small frog inhabits the harsh semiarid Patagonian scrub and steppe, where it is most common in arroyos and margins of lakes.

behavior

Pleurodema bufonina is active by day and night, especially after rains. It seeks shelter under stones and in crevices.

feeding ecology and diet

Nothing is known; presumably it feeds on small arthropods.

reproductive biology

Breeding takes place in shallow water in the austral spring. Males do not call; amplexus is inguinal. Eggs are laid in irregular strings in shallow water. Tadpoles attain a maximum length of about 1.4 in (35 mm); the body is ovoid, and the eyes are small and directed dorsolaterally. The caudal musculature is moderately robust, and the dorsal fin does not extend onto the body. The oral disc is small and directed anteroventrally; the jaw sheaths are broadly arched and finely serrate, and there are two anterior and three posterior rows of labial teeth. The body and caudal musculature are grayish brown, and the belly is gray.

conservation status

Not threatened.

significance to humans

None known.

Cururu lesser escuerzo

Odontophrynus occidentalis

subfamily

Odontophryinae

taxonomy

Ceratophrys occidentalis Berg, 1896, Arroyo Agrio, Neuquén, Argentina.

other common names

None known.

physical characteristics

Males attain a maximum snout-vent length of 2.4 in (60 mm) and females, 2.6 in (65 mm). The body is robust and toadlike. The head is broad with a rounded snout and small tympanum. The skin on the dorsum is pustular with enlarged glands on the eyelids, posterior to the eyes, arms, and legs; the venter is coarsely areolate. The fingers and toes have narrow lateral fringes, and the toes are about one-third webbed. A large, shovel-shaped tubercle is present at the base of the foot. The dorsum is various shades of brown with a middorsal tan stripe; the venter is cream. The iris is dull bronze with black flecks. Breeding males have dark brown nuptial excrescenses on the inner surface of the thumb and dorsal surface of the first finger.

distribution

This species is widely distributed at elevations from near sea level to 6,500 ft (2,000 m) in central and western Argentina.

habitat

This frog inhabits arid and semiarid sandy areas; usually individuals are near streams.

behavior

This nocturnal species is active above ground only during the rainy season. Using their hind feet for digging, individuals spend the dry season underground. In exceptionally dry years, the frogs do not emerge and are known to spend two years in their underground retreats.

feeding ecology and diet

For its size, this frog eats relatively large prey—large arthropods and small mice.

reproductive biology

Males call day and night while floating in water; the call is a long series of low-pitched notes. Heavily pigmented eggs are deposited in deep, natural pools at the edges of streams. Tadpoles grow to a maximum length of 4.6 in (117 mm); they have globular bodies with dorsally directed eyes, high caudal fins, and oral discs that have two anterior and three posterior rows of labial teeth.

conservation status

Not threatened.

significance to humans

None known.

Patagonia frog

Atelognathus patagonicus

subfamily

Telmatobiinae

taxonomy

Batrachophrynus patagonicus Gallardo, 1962, Laguna Blanca, Neuquén Province, Argentina.

other common names

None known.

physical characteristics

Adults attain a maximum snout-vent length of 2 in (50 mm). The snout is acutely rounded in dorsal and lateral views. The eyes are small and directed anterolaterally; the tympanum is obscured by a dermal fold. The skin on the dorsum and venter is smooth; in aquatic adults, loose flaps of skin are present on the sides of the body and on the thighs, but these are absent in terrestrial subadults. The fingers are unwebbed, and the toes are fully webbed. The dorsum is dull brown to live brown with faint darker spots or flecks, and the venter is pale orange; the iris is pale bronze-brown. Breeding males have smooth gray nuptial excrescences.

distribution

This species is known only from the basaltic Laguna Blanca and nearby small lakes in northern Patagonia, Argentina.

habitat

Adults inhabit cold lakes with rocky bottoms; subadults are terrestrial in grassy pampas, where they take refuge under stones.

behavior

Adults are aquatic and swim among submerged rocks on the bottoms of shallow lakes. Upon metamorphosis, young move onto land and subsequently enter lakes, where they develop loose, baggy skin, which provides additional surface area for integumentary respiration in the cold water.

feeding ecology and diet

The diet consists of aquatic arthropods, especially amphipods.

reproductive biology

Small eggs are randomly attached to aquatic plants. Tadpoles are bottom-dwellers in shallow water. They reach a total length of about 2 in (50 mm) and have a golden brown dorsum with small brown spots and translucent fins. The body is depressed, and the eyes and nostrils are dorsal; the oral disc is directed anteroventrally and bears two anterior and three posterior rows of labial teeth.

conservation status

This species is not listed by the IUCN. However, introduction of trout into Laguna Blanca has resulted in a decline in abundance of this species, which might be near extinction.

significance to humans

None known.

Helmeted water toad

Caudiverbera caudiverbera

subfamily

Telmatobiinae

taxonomy

Lacerta caudiverbera Linnaeus, 1758, Peru (in error).

other common names

None known.

physical characteristics

Males of this large frog attain a maximum snout-vent length of 4.8 in (120 mm); females as large as 12.8 in (320 mm) have been reported. The body is robust, and the head is large with short, rounded snout. The eyes are small with a vertical pupil and are directed anterolaterally; the tympanum is large and distinct. The skin is smooth with elongate pustules on the dorsum. The fingers are moderately short and unwebbed, and the toes are about one-half webbed. The dorsum is dull brown with faint, paler, irregular markings, and the venter is grayish white. The iris is dull bronze; breeding males have black nuptial excrescences on the inner surfaces of the thumbs.

distribution

This species ranges throughout the lowlands of Chile between 30° and 42° south latitude.

habitat

This species is primarily aquatic in ponds, lakes, and rivers.

behavior

These large frogs are active by day and night. They are aggressive toward potential predators. The frogs inflate the lungs, elevate the body, open the mouth, lunge, and bite.

feeding ecology and diet

Caudiverbera is a voracious carnivore. Adults eat aquatic insect larvae, fishes, frogs, and even small birds and mammals.

reproductive biology

Breeding occurs in September and October when males call from shallow water. The call is a loud "oouü." Amplexus is axillary. Eggs are laid in clumps in shallow water; clutches consist of 1,000–10,000 eggs about 0.10–0.12 in (2.7–3.1 mm) in diameter. The eggs hatch about 20 days after deposition, and the larval duration is about two years. The tadpoles reach a maximum length of about 6 in (150 mm). The body is ovoid, slightly wider than high, with an angular snout; the eyes are directed dorsolaterally. The caudal musculature is moderately robust, and the dorsal fin originates on the posterior part of the body. The oral disc is directed anteroventrally; there are three anterior and three posterior rows of labial teeth. The body and anterior two-thirds of the tail are grayish brown, and the posterior part of the tail is dark brown to black.

conservation status

This species is negatively affected by habitat degradation and hunting pressure. It is listed as Data Deficient by the IUCN.

significance to humans

Caudiverbera is subjected to human consumption.

Emerald forest frog

Hylorina sylvatica

subfamily

Telmatobiinae

taxonomy

Hylorina sylvatica Bell, 1843, Chonos Island, Chile.

other common names

None known.

physical characteristics

This frog attains a maximum snout-vent length of 2.5 in (62 mm). The skin on the dorsum is slightly tubercular, and the belly is smooth. The snout is bluntly rounded; the eyes are large and prominent with vertical pupils, and the tympanum is distinct and about one-half of the diameter of the eye. The fingers and toes are long, slender, and unwebbed. The dorsum is pale green with coppery brown markings; the venter is pale cream. The iris is brown; breeding males have smooth, gray nuptial excrescences on the thumbs.

distribution

This species is restricted to the austral humid forests of southern Chile and adjacent Argentina.

habitat

Hylorina inhabits humid forests.

behavior

This nocturnal species spends its days under logs; at night it perches in bushes.

feeding ecology and diet

Presumably this species feeds on small arthropods.

reproductive biology

Most reproductive activity occurs in January, when males call from the edges of ponds. The call is a series of low-pitched notes. Amplexus is axillary. Clumps of 400–500 eggs (ova about 0.08 in or 2 mm in diameter) are deposited at the bases of plants in shallow water. In about 10 days, tadpoles hatch in developmental Stage 21. Tadpoles attain a maximum size of about 2.4 in (60 mm) and require about one year to develop to metamorphosis. Tadpoles have a broad, slightly depressed body with dorsolateral eyes. The oral disc is directed anteroventrally and has two anterior and two posterior rows of labial teeth.

conservation status

Although this frog is not listed by the IUCN, extensive deforestation is restricting its habitat.

significance to humans

None known.

Titicaca water frog

Telmatobius culeus

subfamily

Telmatobiinae

taxonomy

Cycloramphus culeus Garman, 1875, Lake Titicaca, Peru and Bolivia.

other common names

None known.

physical characteristics

This large, aquatic frog attains a snout-vent length of about 6 in (150 mm). The snout is acutely rounded, and the eyes are relatively small and protuberant dorsally; a tympanum is not evident. The skin is nearly smooth and tends to be loose and somewhat baggy. The digits are long with narrowly rounded tips; fingers are unwebbed, and the toes are about one-half webbed. The dorsum is dull olive green or dark brown, with or without paler or darker spots. The venter is creamy gray, and the iris is dull bronze.

distribution

This species is known only from Lake Titicaca and nearby lakes in the Titicaca Basin at elevations of about 12,300 ft (3,800 m) in the Andes in southern Peru and adjacent Bolivia.

habitat

This strictly aquatic frog inhabits shallower parts of lakes where the water temperature is about 50°F (10°C).

behavior

Living in cold, well-oxygenated water, Telmatobius culeus has a low metabolic rate; the lungs are relatively small, and apparently all respiration occurs through the skin.

feeding ecology and diet

Nothing is known.

reproductive biology

Eggs are laid in water and hatch into feeding tadpoles, which attain maximum total lengths of about 3.1 in (80 mm). The tadpoles have a large, globular body with a round snout. The oral disc is directed anteroventrally and bears keratinized jaw sheaths and two anterior and three posterior rows of labial teeth. The body is dark gray with white flecks, and the caudal fins are tan.

conservation status

This frog is not listed by the IUCN. However, in Lake Titicaca, Telmatobius culeus is threatened by pollution and hunting.

significance to humans

This frog is a staple for residents in the vicinity of Lake Titicaca; the frogs are collected with seines (nets) and sold in local markets.

Resources

Books

Cei, José M. Batracios de Chile. Santiago: Ediciones se la Universidad de Chile, 1962.

Joglar, Rafael L. Los Coquíes de Puerto Rico. San Juan: Editorial de la Universidad de Puerto Rico, 1998.

Ryan, Michael J. The Túngara Frog. Chicago: University of Chicago Press, 1985.

Periodicals

Cei, José M. "The Amphibians of Argentina." Monitore Zoologico Italiano Monografia 2 (1980): 1–609.

Heyer, W. Ronald. "A Preliminary Analysis of the Intergeneric Relationships of the Frog Family Leptodactylidae." Smithsonian Contributions from Zoolology 31 (1975): 1–55.

Heyer, W. Ronald, A. Stanley Rand, Carlos A. G. da Cruz, Oswaldo L. Peixoto, and Craig E. Nelson. "Frogs of Boracéia." Arquivos de Zoología Univ. São Paulo 31 (1990): 231–410.

Lynch, John D. "Evolutionary Relationships, Osteology, and Zoogeography of the Frog Family Leptodactylidae." Miscellaneous Publications of the Museum of Natural History, University of Kansas 53 (1971): 1–238.

Lynch, John D. "A Re-assessment of the Telmatobiine Leptodactylid Frogs of Patagonia." Occasional Papers of the Museum of Natural History, University of Kansas, no. 72 (1978): 1–57.

Lynch, John D., and William E. Duellman. "Frogs of the Genus Eleutherodactylus (Leptodactylidae) in Western Ecuador: Systematics, Ecology, and Biogeography." Natural History Museum, Universoity of Kansas Special Publication 23 (1997): 1–236.

William E. Duellman, PhD