Hyla versicolorGray Treefrog

Ge­o­graphic Range

Both gray treefrog species, Hyla ver­si­color and Hyla chrysoscelis, in­habit a wide range from south­ern On­tario and Maine, west­ward to cen­tral Texas, north­west to Man­i­toba, and north­ern Florida. An iso­lated colony is also noted in New Brunswick. The two species ap­pear phys­i­cally iden­ti­cal, and con­se­quently more stud­ies are nec­es­sary to de­lin­eate where the species over­lap. (Collins and Co­nant, 1998)

Gen­er­ally, the east­ern gray treefrog (H. ver­si­color) is mostly found to the north and north­east of the range. How­ever, the gray treefrog species are ex­tremely vari­able in their dis­tri­b­u­tion pat­tern. For in­stance, the east­ern gray treefrog is com­mon in the east­ern Great Lakes re­gion, in­clud­ing south­ern Michi­gan; how­ever, both species: H. ver­si­color and H. chrysoscelis share the same breed­ing ponds in Wis­con­sin and north­ern Michi­gan. (Hard­ing, 1997)

Habi­tat

East­ern gray Treefrogs in­habit all el­e­va­tions of wooded areas near tem­po­rary and per­ma­nent wa­ters in such di­verse sur­round­ings such as swamps, ponds, lakes, old fields, thickly wooded sub­ur­ban neigh­bor­hoods, farm wood­lots, and mixed or de­cid­u­ous forests. Dur­ing the sum­mer months, they are most often found in damp rot­ten logs or hol­low trees. In win­ter, gray treefrogs hi­ber­nate on land under woody de­bris such as logs, roots and leaf lit­ter. (Hard­ing, 1997)

  • Aquatic Biomes
  • lakes and ponds
  • temporary pools

Phys­i­cal De­scrip­tion

The east­ern gray treefrog mea­sures 1.25 to 2 inches (3-5cm) in length. The record length is 2.25 in. (6 cm). There is no sex­ual di­mor­phism. The dor­sal sur­face of the gray treefrog species is rough and lightly sprin­kled with warts,more than most frogs but less than the av­er­age toad. The large toepads pro­duce mu­cous to ad­here to smooth bark or man-made struc­tures near light sources, and are char­ac­ter­is­tic of the fam­ily Hyl­i­dae. The col­ors of a gray treefrog vary with the col­ors of its back­ground and en­vi­ron­men­tal fac­tors such as sea­son and hu­mid­ity, but shades of gray are most com­mon with black blotches on the back. Vari­a­tions of brown, green, and pearl-gray col­ors have been noted. Green col­ors are more promi­nent dur­ing the breed­ing sea­son and in year­ling frogs. Usu­ally, there is a white mark be­neath the eye. The ven­tral skin on the hind legs, in the groin re­gion, may ap­pear or­ange to golden yel­low with black speck­les and the belly is white.

If the col­oration is in ques­tion, place the treefrog in a box, allow it to sit qui­etly, and later re-ex­am­ine the spec­i­men. The year­ling frogs are about half the size of the older H. ver­si­color pop­u­la­tion, but re­tain the same char­ac­ter­is­tics. Gray treefrogs con­tinue to grow each year until they achieve the phys­i­cal limit of the species. (Collins and Co­nant, 1998; Hard­ing, 1997)

Both gray treefrog species pos­sess the same lar­val traits, but H. ver­si­color was used to ex­em­plify the tad­pole stage in Co­nant and Collins' "A Field Guide to Rep­tiles and Am­phib­ians of East Cen­tral North Amer­ica". The tad­poles are small, but col­or­ful, 1.25 in. to 1.5 in. (3.2 cm. to 3.8 cm.) long. The tip of the tail is well-de­fined with a 5 mm nar­row tip. The oral disc is com­prised of 2 upper labial tooth rows and 3 lower, ser­rated jaws, and an over­hang­ing upper jaw. The in­testi­nal coil is also vis­i­ble. The back­ground color is light green to yel­low. The tallest sec­tion of the tail fin is the mid­dle and heavy black dots are scat­tered along the mar­gin on a red or or­ange back­ground across the tail. (Collins and Co­nant, 1998; Hard­ing, 1997)

  • Sexual Dimorphism
  • sexes alike
  • Average mass
    7.175 g
    0.25 oz
    AnAge
  • Range length
    3 to 5 cm
    1.18 to 1.97 in
  • Average basal metabolic rate
    0.00433 W
    AnAge

De­vel­op­ment

Tad­poles of the east­ern gray treefrogs meta­mor­pho­size into froglets in six to eight weeks. The young frogs are ap­prox­i­mately 0.6 in. (1.5 cm) snout to vent length. The lar­val and adult rate of growth is de­pen­dent on the avail­abil­ity of food and stress of preda­tors. The sex de­ter­im­i­na­tion of am­phib­ians is ge­netic. How­ever, if lar­vae are treated with es­tro­gen, then hor­monal sex reveral is pos­si­ble after meta­mor­pho­sis. Hyla ver­si­color fol­low the XX/XY pat­tern of het­eroga­mety. (Hard­ing, 1997; Skelly, 1992; Wal­lace, et al., 1999)

Re­pro­duc­tion

The breed­ing cho­ruses of gray treefrogs begin in late April to early May after the evening air tem­per­a­ture rises above 15°C, which varies through­out the range. These frogs end their hi­ber­na­tion in the early months of spring, but do not have the en­ergy re­serves to call, yet. Warm, cloudy nights, from dusk to mid­night, pro­duce the most in­tense cho­ruses. How­ever, in­ter­ludes of cold weather may tem­porar­ily end the male gray treefrog calls. Gen­er­ally, the breed­ing cho­rus lasts for sev­eral weeks. Some­times, the breed­ing calls are con­tin­ued into late June or early July, de­pend­ing on local tem­per­a­tures and un­usual weather phe­nom­ena. (Hard­ing, 1997)

Fe­male choice dom­i­nates the mat­ing scheme of gray treefrogs, since the fe­male ap­proaches the male with the most pro­longed and fre­quent calls. If the male de­tects a nearby fe­male he will also fur­ther en­tice her with a “courtship call” that is longer and more em­phatic than the usual ad­ver­tise­ment call. Suc­cess­ful call­ing re­sults in am­plexus as the fe­male de­posits eggs which are ex­ter­nally fer­til­ized by the male. Al­most im­me­di­ately, the large egg mass breaks into small, loose egg clus­ters of 10 to 40 eggs at­tach to plants or other struc­tures within the pond. De­pend­ing on the water tem­per­a­ture, the tad­poles hatch in three to seven days. Both gray treefrog sp. do not hy­bridize due to a mat­ing bar­rier, the dif­fer­ent pulse rate and pitch be­tween the two calls. (Hard­ing, 1997; Haus­fater, et al., 1990)

East­ern gray treefrogs em­ploy their unique call from the safety of veg­e­ta­tion next to the shal­low breed­ing sites, prefer­ably in tree branches that over­hang the water. The males ag­gres­sively de­fend and use their voice to out­line their ter­ri­to­ries with ex­tended calls. Satel­lite males, often in their first breed­ing sea­son or oth­er­wise dis­ad­van­taged, do not call to save en­ergy. In­stead, they lie in wait near a call­ing male and in­ter­cept the fe­male by claim­ing the caller’s po­si­tion after he moves away. The fe­male only vis­its the breed­ing site to lay her eggs. Dur­ing the last weeks of the breed­ing sea­son, oc­ca­sional calls may still be heard as the males slowly re­treat from the shore­line and dis­ap­pear into the fo­liage. Rare calls may still be heard in the trees in late sum­mer or fall, yet they are un­re­lated to mat­ing, and occur more often dur­ing rain show­ers. Call­ing males are often at­tacked by preda­tors, and this re­sults in a fe­male-bi­ased pop­u­la­tion. (Du­ell­man and Trueb, 1986; Hard­ing, 1997; Steb­bins and Cohen, 1995)

  • Breeding interval
    Treefrogs breed once yearly.
  • Breeding season
    Late April to May or until the temperature rises above 15°C
  • Range number of offspring
    1,000 to 2,000
  • Range time to hatching
    3 to 7 days
  • Average age at sexual or reproductive maturity (female)
    2 years
  • Average age at sexual or reproductive maturity (female)
    Sex: female
    912 days
    AnAge
  • Range age at sexual or reproductive maturity (male)
    1 to 2 years

Fe­male gray treefrogs in­vest in their off­spring by pro­vid­ing yolk to the eggs, and choos­ing ponds that are rel­a­tively free of preda­tors (they es­pe­cially try to avoid fish). Males do not in­vest in the off­spring, and fe­male in­vest­ment ends when she lays her eggs. (Hard­ing, 1997; Skelly, 1992)

  • Parental Investment
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth

Lifes­pan/Longevity

One cap­tive gray treefrog lived for over seven years in cap­tiv­ity. Un­fo­tu­nately, it was not dis­tin­guished as H. chrysoscelis or H. ver­si­color. The po­ten­tial lifes­pan in cap­tiv­ity and the wild is un­known. It is likely that few gray treefrogs die of old age, preda­tors, dis­ease and cli­mac­tic ex­tremes are more likely causes of death. (Hard­ing, 1997)

  • Range lifespan
    Status: captivity
    7 (high) years
  • Average lifespan
    Status: captivity
    7 years
    AnAge

Be­hav­ior

Male east­ern gray treefrogs are ter­ri­to­r­ial dur­ing the breed­ing sea­son. Dur­ing the rest of the year, males and fe­males are tol­er­ant of the pres­ence of con­specifics of sim­i­lar size. How­ever, they are op­por­tunis­tic can­ni­bals, and may eat other gray treefrogs if they are small enough to catch and swal­low. (Du­ell­man and Trueb, 1986)

As a mem­ber of the genus Hyla, east­ern gray treefrogs pos­sess ad­vanced toe pads to ad­here more strongly to ver­ti­cal sur­faces of glass, metal, and pri­mar­ily tree bark. A very low angle be­tween the toe pads and sub­strate with a com­bi­na­tion of mu­cous glands and sur­face mois­ture cre­ate sur­face ten­sion to sup­port the body mass. The toe tips are able to be flex­i­ble and grip more firmly due to small bone or car­ti­lage be­tween two ter­mi­nal toe bones that sup­port the toe pad. These spe­cial­ized ad­he­sion abil­i­ties would not exist if a de­ter­gent were ap­plied to the pad; how­ever, the ad­he­sion would re­turn with the re­moval of the de­ter­gent. Climb­ing abil­ity is an im­por­tant adap­tive trait of this species and it is cen­tral to their sur­vival. (Steb­bins and Cohen, 1995)

When gray treefrogs hi­ber­nate, they ap­pear rigid, and have a high freez­ing tol­er­ance due to glyc­erol in their blood. Dur­ing hi­ber­na­tion, 80% of the body freezes and the eye be­comes opaque as breath­ing and heart­beat are tem­porar­ily sus­pended. Their high tol­er­ance for freez­ing tem­per­a­tures en­abled the gray treefrogs to ex­pand their ter­ri­tory north­ward and to­wards higher el­e­va­tions. (Du­ell­man and Trueb, 1986; Du­ell­man and Trueb, 1986)

Home Range

The home range of the east­ern gray treefrog varies widely de­pend­ing on the suit­abil­ity of the habi­tat and the avail­able food sup­ply. Due to their small size, sev­eral frogs may in­habit one tree if there is a suf­fi­cient food sup­ply. (Hard­ing, 1997)

Com­mu­ni­ca­tion and Per­cep­tion

The male's ad­ver­tise­ment call is the main trait to dis­tin­guish the east­ern gray treefrog (H. ver­si­color) from Cope's gray treefrog (H. chrysoscelis). In gen­eral, the sound is com­prised of a res­o­nant mu­si­cal trill. The east­ern gray treefrog has a slower trill than Cope’s, which is faster and higher pitched. An in­crease in air tem­per­a­ture raises the rate of the trill and tape record­ings may be nec­es­sary for pos­i­tive iden­ti­fi­ca­tion, es­pe­cially if only a sin­gle species is pre­sent. (Collins and Co­nant, 1998)

In com­par­i­son to other frog species in the range, the gray treefrogs calls are shorter, only 0.5 to 3 sec­onds, yet sim­i­lar to the call of the Amer­i­can toad (Anaxyrus amer­i­canus). (Hard­ing, 1997)

In the lar­val state, Hyla ver­si­color uses chemore­cep­tion as its pri­mary method of com­mu­ni­ca­tion and de­fense against preda­tors. Preda­tory fish and sala­man­der lar­vae are de­tected via chemore­cep­tion. In­jured tad­poles also re­lease an "alarm sub­stance" to warn their con­specifics. (Du­ell­man and Trueb, 1986)

Adult gray treefrogs are very sen­si­tive to ground vi­bra­tions and pos­sess ex­cel­lent hear­ing. Yet, dur­ing hi­ber­na­tion they are un­re­spon­sive to most ex­ter­nal stim­uli. (Du­ell­man and Trueb, 1986)

Food Habits

As tad­poles, east­ern gray treefrogs begin life by graz­ing on algae and de­tri­tus in their pond. (Hard­ing, 1997)

After meta­mor­pho­sis, H. ver­si­color prey upon most types of in­sects and their lar­vae. Mites, spi­ders, plant lice, har­vest­men, and snails are also eaten. Gray treefrogs mostly hunt in­sects in the un­der­story of wooded areas in small trees and shrubs, where they may rely upon their cam­ou­flage with less risk of pre­da­tion. How­ever, like most frogs, H. ver­si­color is op­por­tu­ni­tis­tic and may also eat smaller frogs, in­clud­ing other tree frogs. (Collins and Co­nant, 1998)

  • Animal Foods
  • amphibians
  • insects
  • terrestrial non-insect arthropods
  • mollusks
  • terrestrial worms
  • Plant Foods
  • algae

Pre­da­tion

Many as­sorted species of birds, snakes, other frogs, and small mam­mals eat gray treefrogs. These frogs are ar­bo­real to avoid preda­tors, and ex­ploit new food re­sources. They also avoid the at­ten­tion of preda­tors by call­ing after dusk and being most ac­tive in the evening and night. They use cryp­tic col­oration and rarely leave the trees until the breed­ing sea­son. Their skin is able to as­sume most nat­ural col­ors in which it comes into con­tact.

Larger frogs, such as the bull­frog (Litho­bates cates­beianus) and green frog (Litho­bates clami­tans clami­tans, have been ob­served to con­sume gray treefrogs by stalk­ing call­ing males. In the water, giant wa­ter­bugs (Be­lostom­ati­dae) also at­tack Cope's gray treefrog. (Collins and Co­nant, 1998; Hard­ing, 1997)

In the lar­val state, gray treefrogs are sub­ject to pre­da­tion by fish and larger am­phib­ian lar­vae, such as the tiger sala­man­der (Am­bystomma tigrinum). When aquatic preda­tors are abun­dant, gray treefrog tad­poles re­duce their ac­tiv­ity and feed­ing. They grow more slowly, and meta­mor­phose at a smaller size. (Skelly, 1992)

  • Anti-predator Adaptations
  • cryptic

Ecosys­tem Roles

East­ern gray treefrogs can play a crit­i­cal role in the food­web of their ecosys­tems. As tad­poles they may graze enough algae to change the com­mu­nity of algal species in their ponds. Later, local pest pop­u­la­tions of mos­quito, gnats, and flies are re­duced in the ter­ri­tory of a sin­gle gray treefrog. In turn, Cope's gray treefrogs are the prey of larger frogs, car­niv­o­rous birds, and small mam­mals. H. ver­si­color are a sig­nif­i­cant link to sup­port the sur­vival of other an­i­mals in the ecosys­tem. (Hard­ing, 1997)

Like just about all an­i­mals, this species is host to par­a­sitic species. Among oth­ers, Poly­toma nearcticum is a flat­worm that lives in the gills of tad­poles and the blad­der of adults. Ne­ma­todes in the genus Strongy­loides are found in the di­ges­tive sys­tems of these frogs.

Com­men­sal/Par­a­sitic Species
  • Gas­troin­testi­nal ne­ma­tode (Strongy­loides)
  • a mono­ge­nean flat­worm, Poly­toma nearcticum

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Peo­ple ben­e­fit from the sub­stan­tial amount of in­sect pests that are eaten by H. ver­si­color. The spring breed­ing cho­rus also pro­vides evening en­ter­tain­ment to re-af­firm our con­nec­tion with na­ture. We also use the pres­ence of east­ern gray treefrogs as a sci­en­tific tool to in­di­cate the over­all bio­di­ver­sity and the level of con­t­a­m­i­nants in a re­gion. Over­all, the east­ern gray treefrog plays an im­por­tant role in the eco­log­i­cal bal­ance of wooded farm­lands and res­i­den­tial areas and con­tributes to our own well-be­ing. (Hard­ing, 1997)

  • Positive Impacts
  • controls pest population

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

There are no known ad­verse ef­fects of Hyla ver­si­color on hu­mans. (Hard­ing, 1997)

Con­ser­va­tion Sta­tus

Hyla ver­si­color is not cur­rently clas­si­fied as en­dan­gered or of spe­cial con­cern. How­ever, habi­tat de­struc­tion and human pol­lu­tants are con­tribut­ing to the over­all de­cline of am­phib­ians, in­clud­ing frog and toad species. Pub­lic sup­port of habi­tat areas in state parks, na­ture re­serves, and pri­vate prop­erty con­tin­ues to pro­mote the sur­vival of am­phib­ian species. On­go­ing sci­en­tific re­search also im­proves our un­der­stand­ing of this dy­namic species. (Hard­ing, 1997)

Other Com­ments

Cope's gray treefrog (H. chrysoscelis) and the East­ern gray treefrog (H. ver­si­color) are a unique ex­am­ple of spe­ci­a­tion in ac­tion. The two species have very sim­i­lar genes, and ap­pear iden­ti­cal; how­ever, the east­ern gray treefrog pos­sesses a sec­ond chro­mo­some set, twice the num­ber of chro­mo­somes as Cope's gray treefrog. Cope's (H. chrysoscelis) is called a diploid, and the east­ern (H. ver­si­color) is called a tetraploid. The only re­li­able ways to dis­tin­guish be­tween the species are by the calls of the males or by mi­cro­scopic ex­am­i­na­tion of their chro­mo­somes. (Collins and Co­nant, 1998)

H. ver­si­color is thought to have evolved from H. chrysoscelis when an extra chro­mo­so­mal set was passed to sev­eral sur­viv­ing egg masses some­time early in the Pleis­tocene epoch, com­monly known as the "Ice Age." At this time, pop­u­la­tions of H. chrysoscelis were iso­lated by in­ter­ven­ing areas of ex­treme low tem­po­er­a­tures. By the time the cli­mate warmed and the glac­i­ers re­treated, the two pop­u­la­tions had evolved in dif­fer­ent di­rec­tions, and though they now occur to­gether, they no longer in­ter­breed and are dif­fer­ent species. (Ger­hardt, et al., 1994; Hard­ing, 1997)

The west­ern fox snake (Pan­therophis vulpi­nus) and east­ern fox snake (Pan­therophis gloydi) of the Great Lakes Basin in North Amer­ica are an­other ex­am­ple of this pat­tern of spe­ci­a­tion. The habi­tats and prey vary greatly be­tween these two species, yet ex­cept for min­i­mal vari­a­tion of their scale pat­terns they al­most ap­pear iden­ti­cal. (Hard­ing, 1997)

Con­trib­u­tors

George Ham­mond (ed­i­tor), An­i­mal Di­ver­sity Web.

Lee A. Mueller (au­thor), Michi­gan State Uni­ver­sity, James Hard­ing (ed­i­tor, in­struc­tor), Michi­gan State Uni­ver­sity.

Glossary

Nearctic

living in the Nearctic biogeographic province, the northern part of the New World. This includes Greenland, the Canadian Arctic islands, and all of the North American as far south as the highlands of central Mexico.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

carnivore

an animal that mainly eats meat

chemical

uses smells or other chemicals to communicate

choruses

to jointly display, usually with sounds, at the same time as two or more other individuals of the same or different species

crepuscular

active at dawn and dusk

cryptic

having markings, coloration, shapes, or other features that cause an animal to be camouflaged in its natural environment; being difficult to see or otherwise detect.

detritus

particles of organic material from dead and decomposing organisms. Detritus is the result of the activity of decomposers (organisms that decompose organic material).

ectothermic

animals which must use heat acquired from the environment and behavioral adaptations to regulate body temperature

external fertilization

fertilization takes place outside the female's body

fertilization

union of egg and spermatozoan

forest

forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.

freshwater

mainly lives in water that is not salty.

herbivore

An animal that eats mainly plants or parts of plants.

heterothermic

having a body temperature that fluctuates with that of the immediate environment; having no mechanism or a poorly developed mechanism for regulating internal body temperature.

hibernation

the state that some animals enter during winter in which normal physiological processes are significantly reduced, thus lowering the animal's energy requirements. The act or condition of passing winter in a torpid or resting state, typically involving the abandonment of homoiothermy in mammals.

insectivore

An animal that eats mainly insects or spiders.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

metamorphosis

A large change in the shape or structure of an animal that happens as the animal grows. In insects, "incomplete metamorphosis" is when young animals are similar to adults and change gradually into the adult form, and "complete metamorphosis" is when there is a profound change between larval and adult forms. Butterflies have complete metamorphosis, grasshoppers have incomplete metamorphosis.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

nocturnal

active during the night

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

polygynous

having more than one female as a mate at one time

riparian

Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).

saltatorial

specialized for leaping or bounding locomotion; jumps or hops.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

solitary

lives alone

suburban

living in residential areas on the outskirts of large cities or towns.

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

tactile

uses touch to communicate

temperate

that region of the Earth between 23.5 degrees North and 60 degrees North (between the Tropic of Cancer and the Arctic Circle) and between 23.5 degrees South and 60 degrees South (between the Tropic of Capricorn and the Antarctic Circle).

terrestrial

Living on the ground.

territorial

defends an area within the home range, occupied by a single animals or group of animals of the same species and held through overt defense, display, or advertisement

vibrations

movements of a hard surface that are produced by animals as signals to others

visual

uses sight to communicate

Ref­er­ences

Collins, J., R. Co­nant. 1998. A Field Guide to Rep­tiles and Am­phib­ians of East Cen­tral North Amer­ica. 3rd edi­tion, ex­panded. New York: Houghton Mif­flin Co.

Du­ell­man, W., L. Trueb. 1986. Bi­ol­ogy of Am­phib­ians. New York: Mc­Graw-Hill Book Com­pany.

Ger­hardt, H., M. Ptacek, R. Sage. 1994. Spe­ci­a­tion By Poly­poidy in Treefrogs: Mul­ti­ple Ori­gins of the Tetraploid, Hyla ver­si­color. Evo­lu­tion, 48: 898-908.

Hard­ing, . 1997. Am­phib­ians and Rep­tiles of the Great Lakes Re­gion. Ann Arbor, Michi­gan: Uni­ver­sity of Michi­gan Press.

Haus­fater, G., C. Ger­hardt, G. Klump. 1990. Par­a­sites and mate choice in gray treefrogs, Hyla ver­si­color. Amer­i­can Zo­ol­ogy, 30: 299-331.

Skelly, D. 1992. Field Ev­i­dence for a Cost of Be­hav­ioral An­tipreda­tor Re­sponse in a Lar­val Am­phib­ian. Ecol­ogy, 73/2: 704-708.

Steb­bins, R., N. Cohen. 1995. A Nat­ural His­tory of Am­phib­ians. Prince­ton, New Jer­sey: Prince­ton Uni­ver­sity Press.

Wal­lace, H., B. Bad­away, B. Wal­lace. 1999. Am­phib­ian sex de­ter­mi­na­tion and sex re­ver­sal. Cel­lu­lar and Mol­e­c­u­lar Life Sci­ences, 55/6-7: 901-909.