RanidaeRanid Frogs, Ranids, Riparian Frogs, True Frogs

This spe­ciose, wide­spread, and di­verse fam­ily of frogs has un­der­gone such fre­quent re­de­f­i­n­i­tion that it seems likely that the cur­rent de­f­i­n­i­tion is nei­ther mono­phyletic, nor per­ma­nent. As de­fined here, there are two sub­fam­i­lies, con­tain­ing roughly 35 gen­era, and more than 600 species. Ranids are cos­mopoli­tan ex­cept for tem­per­ate South Amer­ica, most of Aus­tralia, the West In­dies, and most Oceanic is­lands. The sub­fam­ily Ran­i­nae has the same range as the fam­ily, while the Petro­pe­deti­nae is re­stricted to sub-Sa­ha­ran Africa. Species of the genus Rana are the only ranids found in the Amer­i­cas or Aus­tralia; most of the fam­ily's di­ver­sity is in the old world.

No char­ac­ters un­equiv­o­cally di­ag­nose Ranidae, lend­ing more strength to the po­si­tion that this is a pa­ra­phyletic group. Most ranids lack ribs, have den­tate upper jaws, eight holo­chordal-pro­coelous pre­sacral ver­te­brae, and an as­tra­galus and cal­ca­neum that are fused only at their ends. The omoster­num and ster­num are usu­ally os­si­fied, ex­cept in Petro­pe­detines, in which the ster­num tends to be car­ti­lagi­nous. Palatines are pre­sent; a parahy­oid is not. The pha­langeal for­mula is nor­mal, and the pupil is hor­i­zon­tal. Diploid num­ber ranges from 16 to 26; some species are poly­ploid.

Ranids tend to ex­hibit ax­il­lary am­plexus and in­di­rect de­vel­op­ment, lay­ing aquatic eggs that de­velop into type IV tad­poles. Ex­cep­tions to this rule in­clude some African petro­pe­detines, such as Arthroleptella, with non­feed­ing tad­poles, and An­hy­drophryne, which has di­rect de­vel­op­ment from ter­res­trial eggs. Some Asian ra­nine gen­era (e.g. Platy­man­tis) also ex­hibit di­rect de­vel­op­ment. Petro­pe­detines tend to be small (less than 25 mm snout-vent length), ter­res­trial frogs with slightly warty skin. Ra­nines vary widely, with sizes rang­ing from 20 - 300 mm snout-vent length (at 300 mm, Con­raua go­liath, is the largest known frog). Some ra­nines are ri­par­ian, with long legs and webbed feet (e.g. Oc­ci­dozyga); oth­ers are toad­like in ap­pear­ance and some­what fos­so­r­ial in habit (e.g. To­mopterna); still oth­ers are ar­bo­real (e.g. Platy­man­tis). The tad­poles of Amolops species are tor­rent-adapted, and have suc­to­r­ial disks on their bel­lies. The Eu­ro­pean Pelo­phy­lax lessonae is a hy­bri­do­ge­netic form that, in some pop­u­la­tions, ex­hibits a mod­i­fied form of cycli­cal partheno­gen­e­sis, in which sex­ual gen­er­a­tions al­ter­nate with asex­ual ones. Re­search on sev­eral species of Rana also form the basis for much that is known of am­phib­ian bi­ol­ogy, as well as pro­vid­ing ed­i­ble legs to gen­er­a­tions of gour­mands.

Ranids are Neo­ba­tra­chi­ans, but re­la­tion­ships among the fam­i­lies of these "ad­vanced" frogs are al­most wholly un­re­solved. Within the Neo­ba­tra­chia, ranids are mem­bers of the su­per­fam­ily Ra­noidea, a clade of de­rived forms that likely loses its mono­phyly if Den­dro­bati­dae is in­cluded. Fam­ily re­la­tion­ships among the ra­noids are in a state of chaos, and should be con­sid­ered un­known. The num­ber of sub­fam­i­lies de­scribed for Ranidae has fluc­tu­ated con­sid­er­ably over the years, co­in­ci­dent with new hy­pothe­ses of re­la­tion­ship. The for­mer sub­fam­i­lies Arthrolepti­nae and Asty­lostern­i­nae have here been re­moved from Ranidae, and placed in their own fam­ily (Arthrolep­ti­dae). Sim­i­larly, the once-ranid Hemisi­nae has been ac­corded fam­ily sta­tus (Hemisoti­dae). Fi­nally, the Man­tel­li­nae, which has been treated as a sub­fam­ily of the ranids by many au­thors, here is di­ag­nosed as a clade within Rha­cophori­dae. The sub­fam­ily Ran­i­nae, and its type genus Rana, are likely pa­ra­phyletic. Within the Ra­noidea, even this re­duced Ranidae is anal­o­gous to the Lep­to­dactyl­i­dae within the Bu­fonoidea: both fam­i­lies rep­re­sent a di­verse, pa­ra­phyletic group of non-treefrogs. The ranids are the pri­mar­ily old world coun­ter­parts to the new world lep­to­dactylids.

Many fos­sil ranids are known, in­clud­ing both ex­tant and ex­tinct species of Rana from the Ter­tiary and Qua­ter­nary in Eu­rope and North Amer­ica, and prob­a­ble Pty­chadena fos­sils from the Miocene of Mo­rocco.

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

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.

Ford, L. S., and D. Can­natella. 1993. The major clades of frogs. Her­peto­log­i­cal Mono­graphs 7:94-117.

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.