@article {2193,
	title = {Dung beetle (Scarabaeus (Pachysoma)) biology and immature stages: reversal to ancestral states under deset conditions (Coleoptera: Scarabaeidae)?},
	journal = {Biological Journal of the Linnean Society},
	volume = {83},
	year = {2004},
	pages = {453-460},
	abstract = {The unique feeding biology of the unusual flightless Namib Desert dung beetle species belonging to Scarabaeus ( Pachysoma ) MacLeay (Scarabaeidae: Scarabaeinae) has been studied in the past but in this report we document breeding biology, larval feeding and immature morphology for the first time. Females provision burrows with frag- ments of dry herbivore dung and detritus, in which the larvae develop and on which they feed. This is a novel devi- ation, and a probable reversal to the ancestral state, from the obligatory brood ball constructed from wet dung in which scarab larvae usually develop. The free-living larvae and pupae have several unique attributes that distin- guish them from relatives that develop within the confines of a brood ball and provide additional support for mono- phyly of the group. Many of these also appear as reversals to a probable ancestral condition. Unique larval characters include a left mandible with two teeth, the absence of a {\textquoteleft}coprine{\textquoteright} hump, small spiracles, and two- or indistinctly three- segmented antennae. Those unique to the pupa are the presence of peculiar prothoracic projections, the absence of lateral tergal supporting projections on the abdomen, and the perpendicular elytra and wings relative to the median bodyline.},
	keywords = {Ball-rollers, flightless, free-living, monophyletic, namib desert, unique, {\textendash}},
	author = {Scholtz, Clarke H. and Harrison, James du G. and Grebennikov, Vasily V.}
}
@article {2213,
	title = {Phylogeography of the Namib Desert dung beetles Scarabaeus (Pachysoma) MacLeay (Coleoptera: Scarabaeidae)},
	journal = {Journal of Biogeography},
	volume = {32},
	year = {2005},
	pages = {75-84},
	abstract = {Aim Namib biogeography in many instances remains reliant on advanced and detailed systematic studies. This study attempts to combine molecular phylogenetic data, geology and palaeo-climatic data to (i) resolve the relationships of the 13 morphological species of Scarabaeus (Pachysoma) and (ii) relate their evolution to past climatic and geological events. Location South Africa and Namibia. Methods Sequencing of a 1197 bp segment of the mitochondrial cytochrome oxidase I (COI) gene of the 13 species within Scarabaeus (Pachysoma) was undertaken. Analyses performed included Parsimony and Maximum Likelihood as well as imposing a molecular clock. Results The molecular phylogeny showed strong support for 11 of the 13 morphological species. The remaining two species, S. (P.) glentoni and S. (P.) hippocrates, formed a complex and could not be assigned specific status on the basis of the COI gene phylogeny. Strong support for the three species formerly classified within the genus Neopachysoma was consistently obtained. The subgenus appears to have arisen c. 2.9 Ma. Species within the subgenus arose at different times, with the common ancestor to Neopachysoma and the hippocrates complex having evolved 2.65 and 2.4 Ma, respectively. Scarabaeus (P.) denticollis, S. (P.) rotundigenus, S. (P.) rodriguesi and S. (P.) schinzi are some of the youngest species, having diverged between 2 million and 600,000 years ago. Main conclusions Scarabaeus (Pachysoma) is a derived monophyletic clade within the Scarabaeini. The subgenus appears to be young in comparison with the age of the Namib Desert, which dates back to the Miocene (c. 15 Ma). The psammophilous taxa are shown to disperse with their substratum and habitat, barchan dunes. Clear south/north evolutionary gradients can be seen within the species of this subgenus, which are consistent with the unidirectional wind regime. Species with a suite of mostly plesiomorphic characters have a southerly distribution while their derived psammophilous relatives have central to northern Namib distributions. Major rivers such as the Orange, Buffels and Holgat appear to be gene barriers to certain species as well as areas of origin of speciation events. },
	keywords = {Aptery, biogeography, COLEOPTERA, cytochrome oxidase I, endemic, mitochondrial DNA, namib desert, phylogeny, Scarabaeidae, Scarabaeus (Pachysoma).},
	author = {Sole, Catherine L. and Scholtz, Clarke H. and Bastos, Armanda D. S.}
}
