@article {1506,
	title = {Cladistic, phenetic and biogeographical analysis of the flightless dung beetle genus, Gyronotus van Lansberge (Scarabaeidae: Scarabaeinae), in threatened eastern Afrotropical forests},
	journal = {Journal of Natural History},
	volume = {35},
	year = {2001},
	pages = {1607-1625},
	author = {Davis, Adrian L. V. and Scholtz, Clarke H. and Harrison, James du G.}
}
@article {1748,
	title = {Phylogeny of Scarabaeus (Pachysoma MacLeay) stat. nov., and related flightless Scarabaeini (Scarabaeidae: Scarabaeinae)},
	journal = {Annals of the Transvaal Museum},
	volume = {40},
	year = {2003},
	pages = {47-71},
	abstract = {ThemonophylyofPachysomaMacLeayisassessedviacladisticanalysis,basedonsixty-fouradultcharactersfrom thirty-seventaxacomprisingall currentlyandpreviously recognizedgenerainScarabaeini, except Madateuchus Paulian.PachysomacomprisesamonophyleticcladeofderivedspeciesofScarabaeusandNeopachysomaFerreira isthesisterlineageofPachysomasensustricto.Scarabaeus(Pachysoma)stat. nov. isconsideredasubgenusof ScarabaeusLinnaeuswhileNeopachysomaFerreiraismaintainedasasynonymof Pachysomasensulato. NeomnematiumJanssenstogether withMnematiumMacLeayispolyphyletic, whilethelatter isparaphyletic. MnematiumMacLeayandNeomnematiumJanssensaremaintainedassynonymsofScarabaeus,andtheirspecies includedinsubgeneraScarabaeus(Scarabaeus) andScarabaeus(Scarabaeolus).DrepanopodusJanssens, KheperJanssensandSceliagesWestwood,asderivedcladesofScarabaeus,couldalsoberecognizedassubgen- eratomaintainamonophyleticScarabaeus,but aremaintainedasgenerapendingfurther work. Aridificationof theNamibDesertispostulatedtohaveinitiatedtheevolutionofScarabaeus(Pachysoma),whichfeedondrydungor detritusandarederivedfromaball-rollingandwet-dung-feedingScarabaeus-likeancestor. Theevolutionof flightlessnessandfoodrelocationstrategy,biogeography,andadditionaldistributionalandbiologicalinformationare presented for the group. },
	keywords = {Aptery, Biology, Drepanopodus, Kheper, Pachylomerus, Pachysoma, phylogeny, Psammophily, Scarabaeolus, Scarabaeus, Sceliages},
	author = {Harrison, James du G. and Philips, T. Keith}
}
@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.}
}
