Search

Aim: The expansion of open habitats during the mid-Miocene has been hypothesized as a driver of allopatric speciation for many African taxa. This habitat-dependent mode of diversification has been implicated in the shift from C3 (e.g. forest/woodland) to C4 dominated systems (i.e. open savanna, grasslands) in a number of African squamates. We examined this hypothesis using a genus of African viperid snakes (Bitis) with both open habitat and forest-dwelling representatives. \ud \ud Location: Africa. \ud \ud Methods: A comprehensive multilocus dataset was used to generate a calibrated species tree using a multispecies coalescent model. Individual gene trees and patterns of nuclear allele sharing were used to assess species monophyly and isolation. To test the habitat-dependent evolution hypothesis, we generated an ancestral character state reconstruction for open and closed habitats using the dated phylogeny. This was related to the timing of open habitat expansion and forest/woodland contraction in Africa. \ud \ud Results: The genus Bitis originated in the Oligocene, with species level diversification in the late Miocene/Pliocene. Four well-supported clades correspond to the recognized subgenera Bitis, Keniabitis, Macrocerastes and Calechidna. Several previously unrecognized lineages potentially represent cryptic species. \ud \ud Main conclusions Habitat-dependent evolution does not appear to have been a main driver for generic level viperine diversification: the ancestral state for Bitis was open habitat and at least one clade moved into forest in the Miocene, long after forest had contracted and fragmented. Forest-dependent species diversified only in the late Miocene, presumably as forest became further reduced in extent, fitting an allopatric model of speciation. Although our results do not favour a general pattern of habitat-dependent diversification in Bitis, cladogenesis within the subgenus Calechidna for “arenicolous” species (Bitis caudalis complex) and “rupicolous” species (B. Atropos-cornuta complex), corresponds to the aridification of southwest Africa. This suggests there are subtleties not captured in the broad open habitat category, which are relevant for understanding the role of habitat-dependent evolution.

Social integration is an important factor when reintroducing group-living species, but examples of the formation of social groups before reintroduction are largely lacking. African wild dogs Lycaon pictus are endangered, and reintroductions have become a routine part of range expansion in South Africa. Wild dogs form packs that are essential to their reproduction and survival, and artificial pack formation is often required before reintroduction. We examined the proximate (i.e. social integration) and ultimate (i.e. reproduction) success of 43 artificial pack formation attempts in the South African managed metapopulation, between 1995 and 2018. The top (and dominant) model for proximate success included an interaction between total group size and an initial separation fence. Larger groups took longer to integrate, irrespective of initial separation, whereas smaller groups brought together immediately integrated faster than those that were initially separated. The top models for ultimate success included an interaction between the proportion of males and number of days spent in the pre-release enclosure, the total number of days in the enclosure and an interaction between the proportion of captive-sourced individuals and the total number of days in the enclosure. Ultimate success increased when packs spent less time in the enclosure, especially if those packs had a low proportion of males (i.e. female biased) or included >25% captive-sourced individuals. Neither the size of the artificially created pack nor the season in which the pack was released affected ultimate success. The success of social integration and reproductive success of artificially formed packs in this study was higher than for natural pack formations. We provide guidelines for optimizing future artificial pack formation in wild dogs for reintroduction success. Our results serve as an example of the practical importance of social behavior in successfully implementing conservation measures for group-living species.