Caudata

Sala­man­ders are dis­trib­uted through­out most of the ho­l­arc­tic re­gion, with the great­est di­ver­sity in the United States. Of ten cur­rently rec­og­nized ex­tant fam­i­lies (com­pris­ing more than 420 ex­tant species), all but one (Hyno­bi­idae) are rep­re­sented in North Amer­ica. Five fam­i­lies are re­stricted to the tem­per­ate West­ern hemi­sphere. Rep­re­sen­ta­tives of the re­main­ing four fam­i­lies are also found in Eu­rope and/or Asia, and one fam­ily has ra­di­ated into trop­i­cal Amer­ica (Pletho­d­on­ti­dae).

Cau­dates are eas­ily dis­tin­guished from other am­phib­ians on the basis of sev­eral mor­pho­log­i­cal char­ac­ters, in­clud­ing the pres­ence of a tail in all lar­vae, ju­ve­niles and adults. Limbs ex­tend at right an­gles from the elon­gate, cylin­dri­cal body, and hindlimbs and fore­limbs tend to be of ap­prox­i­mately equal size (with some no­table ex­cep­tions, such as Sirenidae, which lack hindlimbs). The otic notch and mid­dle ear are ab­sent, as are sev­eral skull bones, in­clud­ing but not lim­ited to the pos­tor­bital, post­pari­etal, jugal, ba­sioc­cip­i­tal, and ec­toptery­goid. When pre­sent, aquatic lar­vae have true teeth on both upper and lower jaws, gill slits and ex­ter­nal gills. Sala­man­ders have the largest genomes of any tetrapods. Adult sizes range from less than one inch to al­most six feet.

Sala­man­ders ex­hibit a wide range of life his­tory and courtship pat­terns. Basal, so-called "prim­i­tive" sala­man­ders have ex­ter­nal fer­til­iza­tion (Cryp­to­bran­choidea, and prob­a­bly Sirenoidea), but the ma­jor­ity of ex­tant fam­i­lies (Sala­man­droidea) have in­ter­nal fer­til­iza­tion. Cop­u­la­tory or­gans do not exist in any sala­man­ders. In in­ter­nally fer­til­iz­ing sala­man­ders, spe­cial­ized cloa­cal glands in males pro­duce sper­matophores, pyra­mi­dal struc­tures capped with sperm. Fe­males pick the sperm cap up, and eggs are usu­ally fer­til­ized as they pass through the cloaca. Most sala­man­ders de­posit aquatic eggs, which hatch into aquatic lar­vae. Some have di­rect de­vel­op­ment of ter­res­trial eggs, and even ovo­vi­vip­ary and true vi­vipar­ity are known in cau­dates. Sev­eral sala­man­ders ex­hibit parental care, which usu­ally takes the form of egg guard­ing by one or both par­ents. Sala­man­ders run the gamut of pos­si­ble de­vel­op­men­tal pat­terns, from com­plete, ob­lig­ate meta­mor­pho­sis, to fac­ul­ta­tive or par­tial meta­mor­pho­sis, to ob­lig­ate pae­do­mor­pho­sis, in which adults re­tain lar­val char­ac­ters through­out their lives (all cryp­to­branchids, sirenids, am­phi­u­mids, and pro­teids are ob­lig­ate pae­do­morphs).

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 (frogs and toads) - Gymnophiona (cae­cil­ians) sis­ter re­la­tion­ship, with Cau­data sis­ter to that group. A sala­man­der-cae­cil­ian clade is sup­ported by soft anatom­i­cal char­ac­ters and ri­bo­so­mal DNA se­quences. Os­te­o­log­i­cal char­ac­ters sup­port a sala­man­der-frog clade, and com­bined mor­pho­log­i­cal and mol­e­c­u­lar ev­i­dence mar­gin­ally sup­port the sala­man­der-frog sis­ter re­la­tion­ship. There is no con­tro­versy as what is or is not a cau­date, al­though some fam­ily de­f­i­n­i­tions re­main trou­bling (e.g. Pro­tei­dae), and there is still con­sid­er­able dis­agree­ment over the iden­tity of sub­or­ders within Cau­data.

Fos­sil sala­man­ders are known from most ex­tant fam­i­lies, as well as four ex­tinct fam­i­lies. The fos­sil record re­veals sala­man­ders be­gin­ning in the upper Juras­sic of North Amer­ica and Eura­sia, ex­tend­ing through the Pleis­tocene. Some fos­sil cau­dates have not yet been as­signed to fam­i­lies. While the terms Cau­data and Urodela are usu­ally used in­ter­change­ably, some au­thors have sug­gested using Urodela to de­scribe only ex­tant forms, while re­tain­ing Cau­data as the more in­clu­sive group in­clud­ing all known ex­tant and fos­sil species.

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.

Lar­son, A., D. Heyse, T. Jack­man, D. Mad­di­son, F. Mof­fitt, and T. Titus. 1996. Cau­data: Tree of life. (Web­site.) http://​tolweb.​org/​tree?​group=Caudata&​contgroup=Living_​Amphibians

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.

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.