Dn. Tarkhnishvili et al., Morphological variation in brown frogs from the Caucasus and the taxonomy of the Rana macrocnemis group, HERPETOLOGI, 55(3), 1999, pp. 406-417
Multivariate analysis of body proportions, skin texture, and coloration cha
racteristics of brown frogs (Rana macrocnemis and R. camerani) from 14 Geor
gian populations confirms the existence of two, geographically distinct gro
ups of populations Frogs in the first group (R. macrocnemis) are characteri
zed by a smooth skin, a pale spotted pattern, absence of a mid-dorsal strip
e, and a small inner metatarsal tubercle. Frogs in the second group (known
as R. camerani) possess a rugose skin, a conspicuous pattern of dorsal spot
s, a mid-dorsal stripe with contrasting borders, and a large metatarsal tub
ercle. The first group occurs in the uplands of the Cleat Caucasus and othe
r forested areas while the second group occurs in the treeless uplands of t
he Near East, A stepped dine exists between them, with parallel variation i
n eight morphological characters. Other characters analyzed vary independen
tly of the dine. Fully diagnostic (fixed) morphological characters separati
ng the groups were not observed. Some characters previously used for taxono
mic purposes are shown to be associated with local ecological conditions. W
e conclude that R, macrocnemis is a single, though polytypic, species compo
sed of two interbreeding evolutionary lineages.