Anura

Anu­rans rep­re­sent, by far, the most spe­ciose, di­verse, and wide­spread of the three ex­tant am­phib­ian or­ders. They are found through­out most of the world, ex­cept in polar re­gions, and some oceanic is­lands and ex­tremely xeric deserts. Anu­ran di­ver­sity is great­est in the trop­ics. Twenty-five fam­i­lies are cur­rently rec­og­nized, rep­re­sent­ing more than 4,000 species, with more being dis­cov­ered reg­u­larly. This clade is also re­ferred to as Ba­tra­chia and is placed in su­per­order Salien­tia.

There is no sci­en­tific dis­tinc­tion be­tween "frogs" and "toads," al­though most anu­rans are usu­ally re­ferred to as one or the other. Anu­rans have sev­eral synapo­mor­phies that dis­tin­guish them from other am­phib­ians. The name, Anura, mean­ing "with­out tail," iden­ti­fies one of these: with one ex­cep­tion (As­ca­phus), adult frogs do not have tails. Anu­rans also have nine or fewer pre­sacral ver­te­brae (usu­ally eight), and the three or four pos­te­rior to the sacrum are fused into a rod called the urostyle. Both the ra­dius and ulna (fore­arm bones), and the tibia and fibula (shank bones), are fused to each other. Fur­ther­more, the as­tra­galus and cal­ca­neum, ankle bones, are greatly elon­gate, pro­vid­ing an ad­di­tional level in the legs of frogs, which they uti­lize in jump­ing. The salta­tory lo­co­mo­tion by which many rec­og­nize frogs is aided, and per­haps even per­mit­ted, by these many mor­pho­log­i­cal adap­ta­tions. Sev­eral skull bones are lack­ing in frogs, al­though their heads re­main highly os­si­fied. The den­tary usu­ally lacks teeth, but the max­illa and pre­max­illa are usu­ally tooth-bear­ing. The tongue is often large, and free pos­te­ri­orly. Males of most species have vocal sacs. The tad­pole stage of many anu­ran life his­to­ries is also unique to frogs, with sev­eral spe­cial­iza­tions such as in­ter­nal gills and the ab­sence of true teeth.

Most anu­rans have ex­ter­nal fer­til­iza­tion, and adopt a mat­ing pos­ture called am­plexus to in­sure con­tact be­tween eggs and sperm. There are ex­cep­tions even to this rule of anu­ran re­pro­duc­tion, which points to the ex­treme di­ver­sity of re­pro­duc­tive modes found in this an­i­mals. The prim­i­tive mode in­volves large num­bers of aquatic eggs and feed­ing lar­vae, but al­ter­nate modes in­clude but are not lim­ited to ter­res­trial eggs with aquatic lar­vae, di­rect de­vel­op­ment (in which there is no tad­pole stage), vi­vipar­ity, and non-feed­ing aquatic lar­vae. Ap­prox­i­mately 10% of anu­rans ex­hibit some form of parental care. Ad­di­tion­ally, many species are highly ter­ri­to­r­ial, de­fend­ing nests, ovipo­si­tion sites, or other re­sources. Many tem­per­ate species breed "ex­plo­sively," con­gre­gat­ing in large num­bers around water for only a night or two each year, dur­ing which time all mat­ing takes place. Some trop­i­cal species breed year-round. Anu­rans are found from trop­i­cal rain­forests to dry moun­tain­tops, from deserts to swamps. Adults may be ar­bo­real, ter­res­trial, aquatic, or fos­so­r­ial. Un­like cae­cil­ians and sala­man­ders, no anu­rans are fully pae­do­mor­phic. Most anu­rans are noc­tur­nal. In the win­ter, many tem­per­ate anu­rans enter a state of tor­por to avoid freez­ing; in arid re­gions, frogs may bury them­selves un­der­ground, or min­i­mize the cu­ta­neous res­pi­ra­tion (and thus the water loss) that usu­ally iden­ti­fies frogs.

De­spite ex­ten­sive re­search on the evo­lu­tion­ary his­tory of am­phib­ians, phy­lo­ge­netic re­la­tion­ships among the three or­ders of ex­tant am­phib­ians re­main prob­lem­atic. Of three pos­si­ble his­to­ries, the only one that has not been se­ri­ously con­sid­ered is an Anura - Gymnophiona (cae­cil­ians) sis­ter re­la­tion­ship, with Cau­data (sala­man­ders) sis­ter to that group. A sala­man­der-cae­cil­ian clade (with Anura sis­ter to that) is sup­ported by soft anatom­i­cal char­ac­ters and ri­bo­so­mal DNA se­quences. Os­te­o­log­i­cal char­ac­ters sup­port a sala­man­der-frog clade, as does a com­bi­na­tion of mor­pho­log­i­cal and mol­e­c­u­lar ev­i­dence. There is lit­tle doubt that Anura, and Salien­tia, are mono­phyletic. Each group is sup­ported by sev­eral synapo­mor­phies. Within Anura, how­ever, and es­pe­cially among the neo­ba­tra­chi­ans ("higher frogs"), very few his­tor­i­cal re­la­tion­ships are re­solved. Anu­ran phy­lo­ge­nies re­main highly con­tentious.

Fos­sil anu­rans are known from the Juras­sic in Eu­rope, North Amer­ica, and South Amer­ica, ex­tend­ing through the Pleis­tocene. Given the ex­ten­sive ra­di­a­tion of many anu­ran clades, the fos­sil record is some­what poor, with sev­eral fam­i­lies lack­ing any fos­sil rep­re­sen­ta­tives. Sev­eral fos­sil gen­era have not been as­signed to rec­og­nized fam­i­lies. The su­per­order Salien­tia in­cludes the fos­sil "proto-frog" Tri­ado­ba­tra­chus, from Mada­gas­car, and Anura.

Can­natella, D., L. Ford, and L. Bock­stanz. 1996. Salien­tia: Tree of Life. (Web­site.) http://​tolweb.​org/​tree?​group=Salientia&​contgroup=Living_​Amphibians

Cog­ger, H. G., and R. G. Zweifel, ed­i­tors. 1998. En­cy­clo­pe­dia of Rep­tiles and Am­phib­ians, 2nd edi­tion. Aca­d­e­mic Press, San Diego.

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

Pough, F. H., R. M. An­drews, J. E. Cadle, M. L. Crump, A. H. Sav­itzky, and K. D. Wells. 1998. Her­petol­ogy. Pren­tice-Hall, Inc., Upper Sad­dle River, NJ.

Steb­bins, R. C., and N. W. Cohen. 1995. A nat­ural his­tory of am­phib­ians. Prince­ton Uni­ver­sity Press, Prince­ton.

Zug, G. R. 1993. Her­petol­ogy: an in­tro­duc­tory bi­ol­ogy of am­phib­ians and rep­tiles. Aca­d­e­mic Press, San Diego.

Con­trib­u­tors

Heather Hey­ing (au­thor).

Glossary

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.

ectothermic

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

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.