Phylogeography of Scarabaeus (Pachysoma) Macleay (Scarabaeidae: Scarabaeinae)

Abstract

Scarabaeus (Pachysoma) consists of 13 flightless dung beetle species endemic to the arid west coast of southern Africa. Scarabaeus (Pachysoma) are unique in their feeding and foraging habits, in that they randomly search for dry dung/detritus which, when found, is dragged forwards, and buried in a pre-constructed holding chamber, as opposed to the convention of rolling it backwards. This action is repeated to provision the chamber after which the nest is expanded to below the moisture line to allow the stored food to re-hydrate. Poor vagility, taxonomic contention - seen in Scarabaeus taxonomy - and conservation concern, made Scarabaeus (Pachysoma) an ideal group of beetles to study both the phylogenetics and potential influences that anthropogenic and environmental changes have had on structuring the species and populations thereof. Both molecular and morphological data were used as individual datasets and combined in a total evidence approach. Biogeographic inferences were made based on recent detailed Namib biogeography and the ages of the species were estimated using the molecular clock method. A phylogeographic study was done on three of the species of Scarabaeus (Pachysoma) – S. (P.) hippocrates, S. (P.) gariepinus and S. (P.) denticollis - that had previously shown south-north morphological clinal variation. Lastly, an attempt was made to isolate microsatellite loci for Scarabaeus, in the hope of characterising genetic diversity within and between populations of the same species. Scarabaeus (Pachysoma) was found to be monophyletic within Scarabaeus and was therefore classified as a derived subgenus thereof. Morphologically Scarabaeus (Pachysoma) was shown to have 13 species while at a molecular level strong resolution for 11 of the 13 was obtained. S. (P.) hippocrates and S. (P.) glentoni formed a species complex the hippocrates/glentoni complex. The combined phylogenetic tree showed good overall support for all 13 species. Both the morphological and molecular data partition phylogenies show congruence with the combined phylogeny, lending support for combining datasets. Scarabaeus (Pachysoma) appears to have arisen 2.9 million years ago. The formation of advective fog is a consistent water source for Desert dwelling organisms and appears to be associated with Scarabaeus (Pachysoma) radiation into inhospitable areas. Analysis of gene flow revealed large amounts of south-north movement, lending support for movement of psammophilous taxa with their substratum, the barchan dune. Population demographics of the three species, S. (P.) hippocrates, S. (P.) gariepinus and S. (P.) denticollis, chosen for this study differed greatly except in areas of geographic similarity. Major rivers appear to have acted as gene barriers, allowing for distinct genetic entities to be identified within the three species. Phylogeographic partitioning was supported by an AMOVA analysis. All three species were shown to have undergone historical population expansion dating back to the Pleistocene era. Nested Clade Analysis indicated that allopatric speciation; isolation by distance and continuous range expansion could be the factors having affected overall population structure. Recent events show that human induced factors, environmental barriers and reduced vagility have influenced the species population structure. Four potentially polymorphic loci were isolated for Scarabaeus using the FIASCO protocol. Identification of at least one additional locus is needed in order to obtain statistical significance for future studies directed at uncovering recent population dynamics.