02112nas a2200229   4500008004100000245015000041210006900191300001200260490000700272520128500279653001701564653001501581653001601596653001701612653001701629653001101646653000801657100002401665700002701689700002801716856013801744       2004                        eng d00aDung beetle (Scarabaeus (Pachysoma)) biology and immature stages: reversal to ancestral states under deset conditions (Coleoptera: Scarabaeidae)?0 aDung beetle Scarabaeus Pachysoma biology and immature stages rev  a453-4600 v833 aThe 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 ‘coprine’ 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.10aBall-rollers10aflightless10afree-living10amonophyletic10anamib desert10aunique10a–1 aScholtz, Clarke, H.1 aHarrison, James, du G.1 aGrebennikov, Vasily, V.  uhttps://scarabnet.myspecies.info/dung-beetle-scarabaeus-pachysoma-biology-and-immature-stages-reversal-ancestral-states-under-deset-c