Geographic variation in the skulls of the horseshoe bats, Rhinolophus simulator and R. cf. simulator: determining the relative contributions of adaptation and drift using geometric morphometrics.

Abstract

The relative contributions of adaptation and drift to morphological diversification of the crania of echolocating mammals was investigated using two horseshoe bat species, Rhinolophus simulator and R. cf. simulator as test cases. We used 3D geometric morphometrics to compare the shapes of skulls of the two lineages collected at various localities in southern Africa. Shape variation was predominantly attributed to selective forces; the between population variance (B) was not proportional to the within population variance (W). Modularity was evident in the crania of R. simulator but absent in the crania of R. cf. simulator and the mandibles of both species. The skulls of the two lineages thus appeared to be under different selection pressures, despite the overlap in their distributions. Selection acted mainly on the nasal dome region of R. cf. simulator whereas selection acted more on the cranium and mandibles than on the nasal domes of R. simulator. Probably the relatively higher echolocation frequencies used by R. cf. simulator, the shape of the nasal dome, which acts as a frequency dependent acoustic horn, is more crucial than in R. simulator, allowing maximization of the intensity of the emitted calls and resulting in comparable detection distances. In contrast, selection pressure is probably more pronounced on the mandibles and cranium of R. simulator to compensate for the loss in bite force because of its elongated rostrum. The predominance of selection probably reflects the stringent association between environment and the optimal functioning of phenotypic characters associated with echolocation and feeding in bats.