Phylogenetic analysis of the Camaenidae (Mollusca: Stylommatophora) with special emphasis on the American taxa

The monophyly of the land snail family Camaenidae has been in doubt due to a disjunct bihemispheric distributional pattern and to the lack of morphological synapomorphies. A cladistic analysis is presented using an ingroup composed of representatives of the three subfamilies distributed in Australia and 52 other species with American distribution. Bradybaenidae, Helicidae and Helminthoglyptidae were used as outgroups. Fifty morphological characters were treated as unordered and analysed using Pee-Wee ver. 2.9, a program for parsimony analysis using implied weights. The results of the analysis support Camaenidae as a monophyletic family (synapomorphies: oval genital orifice, absence of penial sheath). Two of the three Australasian subfamilies, Sinumeloninae and Camaeninae, are monophyletic in the strict consensus tree. The American taxa are classified in eight genera and arranged into two main clades. Caracolus is proposed as the sister group of the American Continental Camaenidae. The genus Solaropsis, previously excluded from this family by different authors, is reassigned to Camaenidae. Shell characters proved to be phylogenetically informative in defining Pleurodonte, Caracolus, Solaropsis, Isomeria and Labyrinthus.