Phylogenetic relationships of the enigmatic land snail genus Prestonella (Mollusca: Stylommatophora) – the missing African element in the Gondwanan family Bulimulidae
The land snail superfamily Orthalicoidea, although generally assumed to be of Gondwanan origin, is considered by the majority of recent authors to be absent from the African continent. However, two poorly-known African genera, Aillya and Prestonella, have historically been referred to the orthalicoid family Bulimulidae s.l. Anatomical study of Aillya has subsequently shown it to be morphologically distinct from the Bulimulidae and referable to a family of its own, outside the Orthalicoidea, but Prestonella has remained an enigmatic taxon of unknown affinity. Using molecular and morphological evidence, we demonstrate conclusively that Prestonella is indeed a member of the Bulimulidae s.l. We thus confirm that this family is represented in Africa, and that it has a classical disjunct, tri-continental southern distribution. Thus, either the origin of the family must at the least predate the separation of Africa and South America in the Mid Cretaceous (under a vicariance scenario) or there must have been subsequent dispersal between the isolated Gondwanan fragments. In view of the limited dispersal ability of terrestrial snails, we consider the former more likely. Anatomically, Prestonella exhibits many character states thought to be plesiomorphic, suggesting a relationship with the subfamily Bulimulinae. Bayesian analysis of nuclear DNA sequence data places it as sister group (posterior probability = 1.0) to an Australasian clade comprising Bothriembryon and Placostylus. However, taxon sampling within the Orthalicoidea is currently inadequate to permit meaningful resolution of subfamilial affinity using molecular data. Similarly, although those orthalicoid taxa for which molecular data are available comprise a well-supported clade, the relationships of this clade to other stylommatophoran clades remain unresolved.