Lithobates palustrisPickerel Frog

Ge­o­graphic Range

The pick­erel frog (Litho­bates palus­tris) is found through­out much of the east­ern United States and parts of south­east­ern Canada. It is found as far north as New Brunswick, Nova Sco­tia, and the Upper Penin­sula of Michi­gan. The range ex­tends as far south as South Car­olina and west­ward into north­ern Al­abama, Mis­sis­sippi, and east­ern Texas. The pick­erel frog is abun­dant along the east coast and is found as far west as Mis­souri. While it is found in north­ern Geor­gia, it is al­most en­tirely ab­sent from Florida. (IUCN SSC Am­phib­ian Spe­cial­ist Group, 2015; Schaff and Smith, 1970)

Habi­tat

The pick­erel frog is found in a va­ri­ety of tem­per­ate fresh­wa­ter habi­tats. In north­ern re­gions, the pick­erel frog can be found in clear, cool wa­ters of lakes, ponds, streams, or pools. In south­ern re­gions, it can be found in warmer wa­ters of swamps and rivers. The dis­tri­b­u­tion in many south­ern and mid­west­ern states is al­most en­tirely de­pen­dent on lime­stone cave habi­tats, with­out which the pick­erel frog likely could not sur­vive in these areas. The pick­erel frog has been re­ported to over­win­ter in these cave habi­tats, which sup­ply a source of cool and clear water and pro­vide pro­tec­tion from harsh en­vi­ron­men­tal con­di­tions. In some states, the pick­erel frog has also been found over­win­ter­ing in aban­doned mines. In warmer months, the pick­erel frog can be found in some ter­res­trial habi­tats such as fields, grass­lands, or wooded areas. (De­Graff and Rudis, 1983; Schaff and Smith, 1970)

  • Aquatic Biomes
  • lakes and ponds
  • rivers and streams
  • temporary pools
  • Other Habitat Features
  • caves

Phys­i­cal De­scrip­tion

Adult pick­erel frogs gen­er­ally have seven pairs of black or brown dor­sal spots be­tween their dor­so­lat­eral folds, as well as a spot on their snout. Vari­a­tion is pre­sent across ge­o­graphic gra­di­ents, and can in­clude dif­fer­ent num­bers of dor­sal spots, pres­ence or ab­sence of a snout spot, the de­gree of fu­sion of dor­sal spots, and the over­all pat­tern of dor­sal spots. Gen­eral body color is gray or tan, while the ven­tral side of the body is white or yel­low. There is a white line pre­sent on the upper lip, and hind legs are usu­ally banded with black or dark brown stripes. The av­er­age size of adults is be­tween 44 to 75 mm SVL (snout-vent length), with a max SVL of 87 mm. There is some sex­ual di­mor­phism be­tween males and fe­males, as fe­males are gen­er­ally re­ported to be larger in size, though spe­cific mea­sure­ments are not known. Dur­ing mat­ing sea­son, males have en­larged thumb pads, which makes them eas­ily iden­ti­fi­able.

Litho­bates palus­tris tad­poles may ex­hibit a va­ri­ety of col­orations, in­clud­ing green, brown, or gray. They may or may not have spots scat­tered along the dor­sal side, but usu­ally have a light or iri­des­cent ven­tral side. Length of tad­poles ranges from 5 to 8 cm. (Hunter, et al., 1999; Schaff and Smith, 1970; Trauth, et al., 2004)

  • Sexual Dimorphism
  • female larger
  • Range length
    87 (high) mm
    3.43 (high) in
  • Average length
    60 mm
    2.36 in

De­vel­op­ment

The eggs of the pick­erel frog are laid in clus­ters on sub­merged twigs or stems, and can de­velop in wa­ters as cool as 8°C and as warm as 29°C. Clus­ters are firm and are 5-10cm across. In­di­vid­ual eggs are en­closed in gel and mea­sure be­tween 3.6 and 5mm in di­am­e­ter. Eggs hatch be­tween 11 and 21 days. The pick­erel frog re­mains a tad­pole for roughly 3 months, at which point meta­mor­pho­sis oc­curs. Length of tad­poles ranges from 5 to 8 cm. The pick­erel frog grows in­de­ter­mi­nately, with adults reach­ing up to 87 mm SVL (snout-vent length). (Hunter, et al., 1999)

Re­pro­duc­tion

Male and fe­male pick­erel frogs gather in large groups to mate. Males croak on land by the water's edge as well as un­der­wa­ter to at­tract fe­males. When a male has suc­cess­fully at­tracted a fe­male, he will hold the fe­male in am­plexus, an em­brace in which the male grasps the fe­male with his front limbs from be­hind and ex­ter­nally fer­til­izes the eggs as they are re­leased. In the case of the pick­erel frog, am­plexus is pec­toral. (Brandt, 1936; Hunter, et al., 1999)

Pick­erel frogs breed yearly in pools, ponds, lakes or other bod­ies of stand­ing water be­tween Feb­ru­ary and May. Dur­ing mat­ing, fe­males cling to sub­merged twigs and stems and de­posit be­tween 2,000 and 3,000 eggs in a clump on the twig or stem, which are fer­til­ized by males dur­ing am­plexus. Eggs hatch in 11 to 21 days, and in­di­vid­u­als re­main in the tad­pole stage for 3 months. Be­cause pick­erel frogs are con­sid­ered to pro­vide lit­tle to no care for off­spring, it is as­sumed that tad­poles are in­de­pen­dent at hatch­ing. After trans­for­ma­tion, pick­erel frogs are gen­er­ally 2.5 cm in length. (Given, 2005; Hunter, et al., 1999; Wright and Wright, 1949)

  • Breeding interval
    Once yearly
  • Breeding season
    February-May
  • Range number of offspring
    2,000 to 3,000
  • Range time to hatching
    11 to 21 days
  • Average time to independence
    0 days
  • Average age at sexual or reproductive maturity (female)
    unknown years
  • Average age at sexual or reproductive maturity (male)
    unknown years

There is no in­for­ma­tion on the level of parental care pro­vided by the pick­erel frog, but most anu­ran species are thought to pro­vide lit­tle to no care. (Beck, 1998; El­liott, 2004; Stu­art, et al., 2008)

  • Parental Investment
  • no parental involvement

Lifes­pan/Longevity

The lifes­pan of the pick­erel frog is un­known. Sim­i­lar frogs in the Litho­bates genus, such as the north­ern leop­ard frog (Litho­bates pip­i­ens), are known to live up to 5 years in the wild and 9 years in cap­tiv­ity. (Lan­noo, 2005)

Be­hav­ior

The pick­erel frog con­gre­gates near bod­ies of water after hi­ber­na­tion and re­mains there dur­ing the mat­ing sea­son. After the mat­ing sea­son ends, the pick­erel frog may mi­grate out into fields and woods where it lives a mostly soli­tary lifestyle. Dur­ing mat­ing sea­son, the pick­erel frog is most ac­tive at night, but may be ac­tive dur­ing the day after mat­ing sea­son ends. The pick­erel frog hi­ber­nates dur­ing win­ter months, al­though it goes into hi­ber­na­tion much later than the vi­su­ally-sim­i­lar north­ern leop­ard frog (Litho­bates pip­i­ens) and may be some­what mo­bile dur­ing win­ter months. The pick­erel frog is usu­ally found in the sub­strate of ponds dur­ing hi­ber­na­tion, al­though it can over­win­ter in aban­doned mines or caves as well. In gen­eral, the pick­erel frog is not ter­ri­to­r­ial, but males may call ag­gres­sively to other males while de­fend­ing call­ing spots dur­ing the mat­ing sea­son. (Brandt, 1936; De­Graff and Rudis, 1983; Given, 2005; Hunter, et al., 1999)

Home Range

There are no stud­ies that name a spe­cific home range for the pick­erel frog.

Com­mu­ni­ca­tion and Per­cep­tion

The call of the pick­erel frog is de­scribed as a short snor­ing sound, last­ing an av­er­age two sec­onds in length at 1222 Hz. The male pick­erel frog will vo­cal­ize calls to at­tract fe­males dur­ing mat­ing sea­son. Like many anu­ran species, the pick­erel frog uti­lizes the same call­ing spot each night dur­ing mat­ing sea­son, and call­ing cho­ruses can last up­wards of five hours. Typ­i­cally, the male pick­erel frog will broad­cast into the air, but some in­stances of un­der­wa­ter call­ing have been doc­u­mented in dis­turbed habi­tats.

In ad­di­tion to a mat­ing call, the pick­erel frog has been doc­u­mented uti­liz­ing two ag­gres­sive vo­cal­iza­tions in male-to-male in­ter­ac­tions. These calls have been de­scribed by Given (2004) as a "growl" and a "snicker." Both calls are shorter in du­ra­tion than the mat­ing call, last­ing an av­er­age of 1.3 sec­onds or 0.21 sec­onds, re­spec­tively. (Given, 2005; Given, 2008; Van­De­Walle, 2011)

Food Habits

Pick­erel frogs are car­niv­o­rous, and are gen­er­ally known to rely on ter­res­trial arthro­pods as a main food source, specif­i­cally in­sects and arach­nids. Other food sources in­clude snails, cray­fish, isopods and aquatic am­phipods, but these or­gan­isms only make up roughly 5% of their diet. In cave sys­tems in the Ozark Moun­tains, pick­erel frogs have been ob­served eat­ing stream isopods, al­though they are not con­sid­ered to be a sig­nif­i­cant threat to stream isopods in these caves. (De­Graff and Rudis, 1983; Feno­lio, et al., 2005)

  • Animal Foods
  • insects
  • terrestrial non-insect arthropods
  • other marine invertebrates

Pre­da­tion

Data on pick­erel frog pre­da­tion are lim­ited, but there are recorded in­stances of the pick­erel frog being used as bait for pick­erel (Esox spp.). There have also been ob­ser­va­tions of pre­da­tion by bald ea­gles (Hali­aee­tus leu­co­cephalus) and minks (Neo­vi­son vison), as well as bull­frogs (Litho­bates cates­beianus) and green frogs (Litho­bates clami­tans) in cap­tiv­ity. De­spite the yel­low warn­ing col­oration on the thighs of the pick­erel frog, which is ex­hib­ited due to toxic skin se­cre­tions, it is sus­pected that the pick­erel frog is sub­ject to other ver­te­brate preda­tors. Tad­poles are also likely to be prey for many fish and in­ver­te­brates. (Hunter, et al., 1999; Lan­noo, 2005; Van­De­Walle, 2011)

Ecosys­tem Roles

Like many anu­rans, the pick­erel frog is sub­ject to in­fec­tion by the am­phib­ian chytrid fun­gus (Ba­tra­chochytrium den­dro­ba­tidis). The pick­erel frog is also sus­cep­ti­ble to var­i­ous par­a­sites in­clud­ing pro­to­zoans (Myx­i­d­ium serot­inum, Nyc­totherus cordi­formis, Opalina species, and Ichthyophonus species), flukes (such as Echi­nos­toma triv­o­lvis, Brachy­coelium sala­man­drae and Glypthelmins qui­eta), par­a­sitic flat­worms (Mesoces­toides species), ne­ma­todes (Ab­bre­vi­ata species, Cos­mo­cer­coides vari­abilis, and Os­wal­docruzia pip­i­ens), and trom­bi­culid mites (Han­ne­ma­nia species). Stud­ies on the rate of in­fec­tion in the pick­erel frog are lim­ited, but pick­erel frog pop­u­la­tions are gen­er­ally sta­ble. The pick­erel frog pro­vides food web sta­bil­ity be­cause of its role as an in­sec­ti­vore and as a prey item for fish, birds, and other preda­tors. (David­son and Cham­bers, 2011; Jones, et al., 2013; Lan­noo, 2005; Orlofske, et al., 2009)

Com­men­sal/Par­a­sitic Species
  • pro­to­zoans (Myx­i­d­ium serot­inum)
  • pro­to­zoans (Nyc­totherus cordi­formis)
  • pro­to­zoans (Opalina species)
  • pro­to­zoans (Ichthyophonus species)
  • flukes (Echi­nos­toma triv­o­lvis)
  • flukes (Brachy­coelium sala­man­drae)
  • flukes (Glypthelmins qui­eta)
  • par­a­sitic flat­worms (Mesoces­toides species)
  • ne­ma­todes (Ab­bre­vi­ata species)
  • ne­ma­todes (Cos­mo­cer­coides vari­abilis)
  • ne­ma­todes (Os­wal­docruzia pip­i­ens)
  • trom­bi­culid mites (Han­ne­ma­nia species)

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

The pick­erel frog has been used as fish­ing bait by hu­mans. Like most other anu­rans, it is likely that the pick­erel frog pro­vides reg­u­lat­ing ser­vices in the form of pest con­trol be­cause it is an in­sec­ti­vore, and may po­ten­tially con­tribute to dis­ease con­trol through con­sump­tion of mos­qui­toes. How­ever, such con­tri­bu­tions have not been sci­en­tif­i­cally demon­strated. (Lan­noo, 2005)

  • Positive Impacts
  • body parts are source of valuable material

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

The pick­erel frog is poi­so­nous. It se­cretes tox­ins through its skin, which can be harm­ful to hu­mans and pets if in­gested. (Lan­noo, 2005; Schaff and Smith, 1970)

Con­ser­va­tion Sta­tus

The pick­erel frog is listed by the IUCN as "Least Con­cern." The pick­erel frog is not given any spe­cial con­ser­va­tion sta­tus on the US Fed­eral List or CITES. With thou­sands of sub-pop­u­la­tions and a wide range of habi­tats, the pick­erel frog pop­u­la­tion is con­sid­ered sta­ble, al­though there are lo­cal­ized pick­erel frog de­clines in many states in­clud­ing Iowa, Wis­con­sin, Min­nesota, and Kansas, as well as On­tario, Canada. The pick­erel frog is sub­ject to in­fec­tion by am­phib­ian chytrid fun­gus (Ba­tra­chochytrium den­dro­ba­tidis) as well as sev­eral ranaviruses, but these dis­eases have not re­sulted in sig­nif­i­cant pop­u­la­tion de­clines. Pick­erel frog habi­tat is sub­ject to clearcut­ting and ur­ban­iza­tion, which likely af­fects sub-pop­u­la­tions. While species-spe­cific leg­is­la­tion is not in place for the pick­erel frog, it seems to ben­e­fit in­di­rectly from some gov­ern­ment pro­grams such as the Wet­land Re­serve Pro­gram. (IUCN SSC Am­phib­ian Spe­cial­ist Group, 2015; Lan­noo, 2005; Wad­dle, et al., 2012)

Con­trib­u­tors

Annie Rudasill (au­thor), Rad­ford Uni­ver­sity, Cari Mc­gre­gor (ed­i­tor), Rad­ford Uni­ver­sity, Zeb Pike (ed­i­tor), Rad­ford Uni­ver­sity, Karen Pow­ers (ed­i­tor), Rad­ford Uni­ver­sity, April Tin­gle (ed­i­tor), Rad­ford Uni­ver­sity, Jacob Vaught (ed­i­tor), Rad­ford Uni­ver­sity, Tanya Dewey (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

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

aposematic

having coloration that serves a protective function for the animal, usually used to refer to animals with colors that warn predators of their toxicity. For example: animals with bright red or yellow coloration are often toxic or distasteful.

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.

bog

a wetland area rich in accumulated plant material and with acidic soils surrounding a body of open water. Bogs have a flora dominated by sedges, heaths, and sphagnum.

carnivore

an animal that mainly eats meat

choruses

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

diurnal
  1. active during the day, 2. lasting for one day.
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.

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.

indeterminate growth

Animals with indeterminate growth continue to grow throughout their lives.

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).

marsh

marshes are wetland areas often dominated by grasses and reeds.

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.

migratory

makes seasonal movements between breeding and wintering grounds

motile

having the capacity to move from one place to another.

natatorial

specialized for swimming

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.

poisonous

an animal which has a substance capable of killing, injuring, or impairing other animals through its chemical action (for example, the skin of poison dart frogs).

polygynandrous

the kind of polygamy in which a female pairs with several males, each of which also pairs with several different females.

saltatorial

specialized for leaping or bounding locomotion; jumps or hops.

seasonal breeding

breeding is confined to a particular season

sexual

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

solitary

lives alone

swamp

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

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.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

Ref­er­ences

Beck, C. 1998. Mode of fer­til­iza­tion and parental care in anu­rans. An­i­mal Be­hav­ior, 55/2: 439-449.

Brandt, B. 1936. The frogs and toads of east­ern North Car­olina. Copeia, 1936/4: 215-223.

Cassie, B. 1999. Na­tional Audubon So­ci­ety First Field Guide: Am­phib­ians. New York, NY: Na­tional Audubon So­ci­ety.

David­son, S., D. Cham­bers. 2011. Oc­cur­rence of Ba­tra­chochytrium den­dro­ba­tidis in am­phib­ians of Wise County, Vir­ginia, USA. Her­peto­log­i­cal Re­view, 42/2: 214-215.

De­Graff, R., D. Rudis. 1983. Am­phib­ians and Rep­tiles of New Eng­land, Habi­tats and Nat­ural His­tory. Amherst, MA: Uni­ver­sity of Mass­a­chu­setts Press.

Du­ell­man, W., L. Trueb. 1986. Bi­ol­ogy of Am­phib­ians. Bal­ti­more, MD: The Johns Hop­kins Uni­ver­sity Press.

El­liott, L. 2004. The Calls of Frogs and Toads. Me­chan­ics­burg, PA: Stack­pole Books.

Feno­lio, D., G. Graen­ing, J. Stout. 2005. Sea­sonal move­ment pat­tern of pick­erel frogs (Rana palus­tris) in an Ozark cave and eco­log­i­cal im­pli­ca­tions sup­ported by sta­ble iso­tope ev­i­dence. The South­west­ern Nat­u­ral­ist, 50/3: 385-389.

Given, M. 2008. Does phys­i­cal or acousti­cal dis­tur­bance cause male pick­erel frogs, Rana palus­tris, to vo­cal­ize un­der­wa­ter?. Am­phibia-Rep­tilia, 29/2: 177-184.

Given, M. 2005. Vo­cal­iza­tions and re­pro­duc­tive be­hav­ior of male pick­erel frogs, Rana palus­tris. Jour­nal of Her­petol­ogy, 39/2: 223-233.

Hardy, L., L. Ray­mond. 1991. Ob­ser­va­tions on the ac­tiv­ity of the pick­erel frog, Rana palus­tris, in north­ern Louisiana. Jour­nal of Her­petol­ogy, 25/2: 220-222.

Hock­ing, D., K. Bab­bitt. 2014. Am­phib­ian con­tri­bu­tions to ecosys­tem ser­vices. Her­peto­log­i­cal Con­ser­va­tion and Bi­ol­ogy, 9/1: 1-17.

Hov­er­man, J., M. Gray, D. Miller. 2010. Anu­ran sus­cep­ti­bil­i­ties to ranaviruses: role of species iden­tity, ex­po­sure route, and a novel virus iso­late. Dis­eases of Aquatic Or­gan­isms, 89/1: 97-107.

Hunter, M., A. Cal­houn, M. Mc­Col­lough. 1999. Maine Am­phib­ians and Rep­tiles. On­rono, ME: Uni­ver­sity of Maine Press.

IUCN SSC Am­phib­ian Spe­cial­ist Group, 2015. "Litho­bates palus­tris" (On-line). The IUCN Red List of Threat­ened Species 2015. Ac­cessed Jan­u­ary 29, 2016 at http://​www.​iucnredlist.​org/​details/​58690/​0.

Jones, E., K. Hyde, K. Pang. 2013. Fresh­wa­ter Fungi and Fun­gal-like Or­gan­isms. Berlin, Ger­many: De­Gruyter.

Lan­noo, M. 2005. Am­phib­ian De­clines: The Con­ser­va­tion Sta­tus of United States Species. Oak­land, CA: Uni­ver­sity of Cal­i­for­nia Press.

McAl­lis­ter, C., S. Trauth, C. Bursey. 1995. Par­a­sites of the pick­erel frog, Rana palus­tris (Anura: Ranidae) from the south­ern part of its range. The South­west­ern Nat­u­ral­ist, 40/1: 111-116.

Moore, J. 1939. Tem­per­a­ture tol­er­ance and rates of de­vel­op­ment in eggs of am­phibia. Ecol­ogy, 20/4: 459-478.

Orlofske, S., L. Belden, W. Hop­kins. 2009. Mod­er­ate Echi­nos­toma triv­o­lvis in­fec­tion has no ef­fects on phys­i­ol­ogy and fit­ness-re­lated traits of lar­val pick­erel frogs (Rana palus­tris). Jour­nal of Par­a­sitol­ogy, 95/4: 787-792.

Schaff, R., P. Smith. 1970. Ge­o­graphic vari­a­tion in the pick­erel frog. Her­peto­log­ica, 26/2: 240-254.

Schaff, R., P. Smith. 1971. Rana palus­tris (LeConte) pick­erel frog. Cat­a­log of Amer­i­can Am­phib­ians and Rep­tiles, 117/1: 1-3.

Stu­art, S., M. Hoff­man, J. Chan­son, N. Cox, R. Berridge, P. Ra­mani, B. Young. 2008. Threat­ened Am­phib­ians of the World. Barcelona, Spain: Lynx Edi­cions.

Trauth, S., H. Ro­bi­son, M. Plum­mer. 2004. The Am­phib­ians and Rep­tiles of Arkansas. Fayet­teville, AR: The Uni­ver­sity of Arkansas Press.

Van­De­Walle, T. 2011. Frogs and Toads in Your Pocket: A Guide to Am­phib­ians of the Upper Mid­west. Iowa City, IA: Uni­ver­sity of Iowa Press.

Wad­dle, J., B. Glo­rioso, S. Faulkner. 2012. A quan­ti­ta­tive as­sess­ment of the con­ser­va­tion ben­e­fits of the Wet­lands Re­serve Pro­gram to am­phib­ians. Restora­tion Ecol­ogy, 21/2: 200-206.

Wright, A., A. Wright. 1949. Hand­book of Frogs and Toads of the United States and Canada. Ithaca, NY: Com­stock Pub­lish­ing Co.