@article {1438,
	title = {Patterns of daily flight activity in onitine dung beetles (Scarabaeinae: Onitini)},
	journal = {Oecologia},
	volume = {103},
	number = {4},
	year = {1995},
	pages = {444-452},
	keywords = {Diel activity, Dung beetles, Endothermy, Flight behaviour},
	author = {Caveney, Stanley and Scholtz, Clarke H. and McIntyre, Peter}
}
@article {1452,
	title = {Morphological correlates of flightlessness in southern African Scarabaeinae (Coleoptera: Scarabaeidae): testing a condition of the water-conservation hypothesis},
	journal = {Canadian Journal of Zoology},
	volume = {76},
	year = {1998},
	month = {Jan},
	pages = {1123-1133},
	abstract = {Flightlessness in desert beetles is thought to have evolved either as a response to decreased environmental heterogeneity or directly to reduce water loss. The water-conservation hypothesis rests on three conditions: that spiracular transpiration is greater than cuticular transpiration; that cuticular transpiration rates are lower in desert species; and that changes in body form associated with flightlessness lead to an overall reduction in water loss rates. The extreme form of the morphological-convergence condition suggests that this change in body shape should be most pronounced in desert-dwelling taxa. The morphological-convergence condition was examined using a morphometric analysis of body shape in flying and nonflying dung beetle species from two southern African tribes occurring in arid and mesic habitats. Although the Canthonini have a more rounded body than the Scarabaeini, flightless species in both tribes have a more rounded body than the flying ones, except at the smallest body sizes. This rounding is more pronounced in flightless, desert-dwelling Scarabaeini than in flightless species from more mesic habitats. All three conditions required by the water-conservation hypothesis are met in various beetle taxa, but the hypothesis and its conditions have yet to be tested on a single, monophyletic taxon.},
	author = {Chown, Steven L. and Pistorius, Pierre and Scholtz, Clarke H.}
}
@article {1461,
	title = {Twilight orientation to polarised light in the crepuscular dung beetle Scarabaeus zambesianus},
	journal = {The Journal of Experimental Biology},
	volume = {206},
	year = {2003},
	month = {Feb},
	pages = {1535-1543},
	abstract = {The polarisation pattern of skylight offers many arthropods a reference for visual compass orientation. The dung beetle Scarabaeus zambesianusstarts foraging at around sunset. After locating a source of fresh droppings, it forms a ball of dung and rolls it off at high speed to escape competition at and around the dung pile. Using behavioural experiments in the field and in the laboratory, we show that the beetle is able to roll along a straight path by using the polarised light pattern of evening skylight. The receptors used to detect this skylight cue can be found in the ommatidia of the dorsal rim area of the eye, whose structures differ from the regular ommatidia in the rest of the eye. The dorsal rim ommatidia are characterised by rhabdoms with microvilli oriented at only two orthogonal orientations. Together with the finding that the receptors do not twist along the length of the rhabdom, this indicates that the photoreceptors of the dorsal rim area are polarisation sensitive. Large rhabdoms, a reflecting tracheal sheath and a lack of screening pigments make this area of the eye well adapted for polarised light detection at low light levels. The fan-shaped arrangement of receptors over the dorsal rim area was previously believed to be an adaptation to polarised light analysis, but here we argue that it is simply a consequence of the way that the eye is built.},
	keywords = {dung beetle, ommatidia, orientation, polarisation pattern, receptor, rhabdom, Scarabaeus zambesianus, skylight.},
	author = {Dacke, Marie and Nordstrom, Peter and Scholtz, Clarke H.}
}
@article {1462,
	title = {A specialized dorsal rim area for polarized light detection in the compound eye of the scarab beetle Pachysoma striatum},
	journal = {Journal of Comp. Physiology A},
	volume = {188},
	year = {2002},
	pages = {211-216},
	keywords = {COLEOPTERA, compound eye, Polarized light detection, vision},
	author = {Dacke, Marie and Nordstrom, Peter and Scholtz, Clarke H. and Warrant, E. J.}
}
@article {1504,
	title = {A dung beetle survey of selected Gauteng nature reserves: implications for conservation of the provincial scarabaeine fauna},
	journal = {African Entomology},
	volume = {13},
	number = {1},
	year = {2005},
	pages = {1-16},
	abstract = {Aquantitativedungbeetlesurvey(Coleoptera:Scarabaeidae:Scarabaeinae)wasconducted insixGautengnaturereserves(Tswaing,Leeuwfontein,Roodeplaat,Ezemvelo,AbeBailey, Suikerbosrand) representativeof theprovincial rangeinenvironmental conditions. The studyprovidedaprovincial species inventorythat has beentestedfor completeness bycomparisonwithmuseumreferencematerial. It alsopermittedananalysisof major influencesonregional(altitude,annualrainfall)andlocal(soilandvegetationtype)patterns of speciesabundance. Thesurveyrecordedatotal of 152species. Althoughafurther29 specieswererepresentedinreferencecollections,theirabsencefromthepresentworkwas probablyduetohabitat,food,ortemporalspecializations.Multivariateanalyses(clustering, MDS)ofspeciesabundancedataformedsixclusters,eachcomprisingexclusivelythesitesin singlereserves. Thisindicatesthatregionalbetween-reservefaunaldifferencesaregreater thanlocal withinreservedifferences, thusdemonstratingthevalueof eachreserve. Ina hierarchical analysisofobliquefactors, fiveoutofsevenstatisticallydefinedclusterscom- prisedexclusivelythesitesinsinglereserveswithSuikerbosrandsplitintwo.Theseclusters werevariouslycorrelatedwithnineextendedfactors.Alongsevenfactors,correlationswere uniqueoressentiallyuniquetosinglereservesandwerefairlyhigh,particularlyatinterme- diatealtitude. Alongtheremainingtwosharedfactors, therewereopposingtrendsinthat correlationseitherdecreasedwithhigheraltitudeordecreasedwithloweraltitude. Thus, eachclustershowsthreecoefficientsofdetermination, onerepresentingtheproportionof varianceduetosharedhighlandfaunalinfluence,onerepresentingthatduetosharedlow- landfaunal influenceandthethirdrepresentingthatduetouniquelocal faunal composi- tion. Theanalysesidentifythetransitionfromlowlandtohighland-dominatedfaunas, the relativefaunal distinctivenessofeachreserve, andomissionsfromtheprovincial reserve system.},
	keywords = {conservation, Dung, Gauteng, nature reserve, Scarabaeinae, SouthAfrica, survey},
	author = {Davis, A. L. V. and Scholtz, Clarke H. and Deschodt, C.}
}
@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 {1701,
	title = {Larvae of Ceratocanthidae and Hybosoridae (Coleoptera: Scarabaeoidea): study of morphology, phylogenetic analysis and evidence of paraphyly of Hybosoridae},
	journal = {Systematic Entomology},
	volume = {29},
	year = {2004},
	pages = {524-543},
	abstract = {Larvae of the scarabaeoid genera Germarostes Paulian, Cyphopisthes Gestro, Paulianostes Ballerio, Ceratocanthus White, Pterorthochaetes Gestro, Madrasostes Paulian, Astaenomoechus Mart{\i} ́nez \& Pereira (Ceratocanthidae) and Hybosorus Macleay, Phaeochrous Castelnau, and Anaides Westwood (Hybo- soridae) are described, keyed and illustrated with fifty-seven drawings. A phylo- genetic analysis of these two families based on larval morphology is presented. Fifty-four larval morphological and three biological characters from twenty-seven taxa revealed nineteen equally parsimonious cladograms. The monophyly of (Ceratocanthidae {\th} Hybosoridae) is supported by four unambiguous unique syna- pomorphies: dorsal medial endocarina on cranium extended anteriorly into fron- tal sclerite; presence of large membranous spot on apical antennomere; labium dorsally with four pores in centre (secondarily reduced to two pores in some groups); and presence of stridulatory organ on fore- and middle legs (secondarily reduced in some groups). Our analysis suggests that the family Hybosoridae is paraphyletic with respect to Ceratocanthidae. The clade comprising the hybosorid genera Hybosorus and Phaeochrous is the sister group of the remaining Hybo- soridae plus Ceratocanthidae. It is supported by two unambiguous synapomorphies: two apical antennomeres completely joined and the stridulatory organ represented by seven to nine large teeth anteriorly on the middle leg. The hybosorid genus Anaides is a sister group to the remaining Hybosoridae plus Ceratocanthidae (without Hybosorus and Phaeochrous) and the ceratocanthid genus Germarostes is a sister group to the remaining Hybosoridae plus Ceratocanthidae (without Hybosorus, Phaeochrous and Anaides). The ceratocanthid genera Cyphopisthes, Astaenomoechus, Paulianostes, Pterorthochaetes, and Madrasostes constitute a sister group to the hybosorid genus Cryptogenius and are supported by the presence of two reversions: two dorsal pores on labium and completely reduced stridulatory organs on fore- and middle legs. },
	author = {Grebennikov, Vasily V. and Ballerio, Alberto and Ocampo, Federico C. and Scholtz, Clarke H.}
}
@article {1702,
	title = {The basal phylogeny of Scarabaeoidea (Insecta: Coleoptera) inferred from larval morphology},
	journal = {Invertebrate Systematics},
	volume = {18},
	year = {2004},
	pages = {321-348},
	keywords = {Geotrupidae, glaphyridae, hybosoridae, Scarabaeidae},
	author = {Grebennikov, Vasily V. and Scholtz, Clarke H.}
}
@article {1806,
	title = {Changes in a Texas USA Dung Beetle Community between 1975 and 1985 Coleoptera Scarabaeidae Scarabaeinae},
	journal = {The Coleopterists Bulletin},
	volume = {40},
	number = {4},
	year = {1986},
	note = {Using Smart Source Parsing(Recd. 1987)ArticleEnglish},
	pages = {313-316},
	abstract = {A quantitative survey of dung beetles in May 1985 at Welder Wildlife Refuge, Sinton, Texas, resulted in 15 species with the two most common species, the introduced African Onthophagus gazella (Fab.) and the native O. alluvius H. \& C., comprising 49\% of the total samples. Neither of these was recorded during a similar survey 10 years previously. Three uncommon species recorded in the earlier survey were not taken in 1985. The appearance at Welder of O. gazella since 1975 illustrates a successful introduction, whereas the appearance of O. alluvius seems to be the effect of a well-documented increase in annual rainfall.},
	keywords = {Animalia-, Animals-, Arthropoda-, Arthropods-, Coleoptera-: Insecta-, Insects-, Invertebrata-, Invertebrates-, onthophagus-gazella onthophagus-alluvius species diversity species abundance importation rainfall},
	author = {Howden, Henry F. and Scholtz, Clarke H.}
}
@article {1891,
	title = {Survival and reproduction of Euoniticellus intermedius (Coleoptera: Scarabaeidae) in dung following application of cypermethrin and flumethrin pour-ons to cattle},
	journal = {Bulletin of Entomological Research},
	volume = {89},
	year = {1999},
	pages = {543-548},
	abstract = {Cattle were treated with cypermethrin and flumethrin pour-ons to determine possible adverse effects of residues in dung on the survival, fecundity and fertility of Euoniticellus intermedius (Reiche). Dung from both cypermethrin- and flumethrin-treated cattle was collected 1, 2, 3, 5, 7, 14, 21 and 28 days after treatment, with an untreated control for each. Mortality of adult E. intermedius exposed to dung collected from cypermethrin-treated cattle two to seven days after treatment ranged between 80\% and 100\%. Cypermethrin had no effect on adult and immature survival or on fecundity and fertility of E. intermediusexposed to dung collected 1, 14, 21 and 28 days after treatment. Dung collected from flumethrin-treated cattle had no apparent effect on adult survival, egg production, immature survival, adult emergence, fecundity and fertility of E. intermedius, except at seven days after treatment, when fewer brood balls were produced and consequently fewer adults emerged from dung from treated animals compared with the control.},
	author = {Kruger, K. and Lukhele, O. M. and Scholtz, Clarke H.}
}
@article {1892,
	title = {Lethal and sublethal effects of ivermectin on the dung-breeding beetles Euoniticellus intermedius (Reiche) and Onitis alexis Klug (Coleoptera: Scarabaeidae)},
	journal = {Agriculture, Ecosystems and Environment},
	volume = {61},
	year = {1997},
	pages = {123-131},
	keywords = {avermectins, dung beetle, fecundity, fertility, larval development},
	author = {Kruger, K. and Scholtz, Clarke H.}
}
@article {1893,
	title = {Possible risk posed by drift of the insect growth regulator pyriproxyfen to the rare dung beetle Circellium bacchus (F.) in a national park},
	journal = {Journal of Insect Conservation},
	volume = {1},
	year = {1997},
	pages = {215-220},
	abstract = {Aerial drift of the juvenile hormone analogue (JHA), pyriproxyfen (Nemesis 100 ec{\textregistered}), used to control red scale on citrus on farms close to the Addo Elephant National Park in South Africa, was suspected of causing a decline in a population of the rare dung beetle species Circellium bacchus(F.) in the Park. The effect of pyriproxyfen on fertility and egg viability, as well as larval, pupal and callow adult development of C. bacchuswas studied in a laboratory assay. Adult beetles, soil surface and the initial dung supply were exposed to pyriproxyfen applied as a spray at 10 times less than the concentration used commercially to simulate spray drift. Exposure of adult beetles to pyriproxyfen did not affect egg production or the viability of eggs, nor did the compound have adverse effects on immature development, indicating that pyriproxyfen is unlikely to be the cause of the observed population depression of C. bacchus.},
	keywords = {development, juvenile hormone analogue, reproduction, Scarabaeidae, spray drift},
	author = {Kruger, K. and Scholtz, Clarke H.}
}
@article {1894,
	title = {Changes in the structure of dung insect communities after ivermectin usage in a grasslansd  ecosystem. II. Impact of ivermectin under high-rainfall conditions},
	journal = {Acta Oecologica},
	volume = {19},
	number = {5},
	year = {1998},
	pages = {439-451},
	keywords = {avermectin, dung beetle, ecotoxicity, longer-term study, non-target organisms},
	author = {Kruger, K. and Scholtz, Clarke H.}
}
@article {1895,
	title = {Biology and ecology of Circellium bacchus (Fabricius 1781) (Coleoptera, Scarabaeidae), a South African dung beetle of conservation concern},
	journal = {Tropical Zoology},
	volume = {19},
	year = {2006},
	pages = {185-207},
	abstract = {The dung beetle Circelliuin bacchus (Fabricius 1781) was once widespread in southern Abica but is now restricted to a few isolated fragments in the south of the Western and Eastern Cape Provinces of South Africa, with the largest pop- ulation present in the Addo Elephant National Park (AENP). The beetles{\textquoteright} activity is governed by precipitation and temperature; 18-26 "C and high relative humid- ity favour activity. The beetles are generalist dung feeders and breeders but have clear preferences for elephant dung for feeding, and buffalo dung for breeding. Fecundity is the lowest recorded for a dung beetle, with a maximum of two but on average only one progeny produced per year. The species is unique amongst ball-rolling (ielecoprid) dung beetles in that females initiate, form and roll brood balls, something carried out exclusively by males in other species. The beetles are habitat specialisrs prefening dense undisturbed vegetation to more open, dis- turbed vegetation. The species should be considered rare because of its narrow geographical range, restricted biotope specificity and biological attributes. Fur- thermore, ii complies with most of the characteristics that increase its likelihood of endangerment or extinction and qualifies as "vulnerable" according to IUCN criteria of threatened species.},
	keywords = {Circellrum bacchus, conservation, ecology, scarab, vulnerable},
	author = {Kryger, U. and Cole, K. S. and Tukker, R. and Scholtz, Clarke H.}
}
@article {1896,
	title = {Effects of cattle treatment with a cypermethrin/cymiazol spray on survival and reproduction of the dung beetle species Euoniticellus intermedius (Coleoptera: Scarabaeidae)},
	journal = {Bulletin of Entomological Research},
	volume = {96},
	year = {2006},
	pages = {597-603},
	abstract = {In a bioassay to determine non-target ecotoxicological effects of a pyrethroid spray (Ektoban1) on dung beetles, dung from both cypermethrin/cymiazol- treated and control cattle was collected one, two, three, five, seven, 14, 21 and 28 days after treatment and fed to a treatment and control group (respectively) of beetles of the species Euoniticellus intermedius (Reiche). This was done to assess whether a spray formulation of cypermethrin may affect dung beetles differently than previously tested pour-on formulations. Following three beetle generations for two weeks each, the experiment retrieved no significant differences in adult or larval survival, egg production, fecundity and fertility between the control and treatment group. These results demonstrated that the used spray formulation of cypermethrin is likely to be far less detrimental to dung beetles than previously tested pour-ons.},
	keywords = {antiparasiticide, cymiazol, cypermethrin, Dung beetles, ecotoxicological effect, Euoniticellus intermedius, pyrethroid},
	author = {Kryger, U. and Deschodt, C. and Davis, A. L. and Scholtz, Clarke H.}
}
@article {1897,
	title = {Effects of cattle treatment with a fluazuron pour-on on survival and reproduction of the dung beetle species Onthophagus gazella (Fabricius)},
	journal = {Veterinary Parasitology},
	volume = {143},
	year = {2007},
	pages = {380-384},
	abstract = {While resistance against many other classes of acaricides has been described, products containing benzoylphenyl urea are currently still successfully used against the pesticide-resistant blue tick (Boophilus decoloratus) in South Africa. In order to assess any adverse impact of these tickicides on the important dung beetle (Coleoptera: Scarabaeidae) fauna, a bioassay was undertaken on the ecotoxicological effects of a fluazuron (benzoylphenyl urea) pour-on formulation (Acatak1) on the survival and reproduction of the African dung beetles species Onthophagus gazella (Fabricius). The experiment yielded no significant differences in adult or larval survival, egg production, fecundity and fertility between the control and treatment group following three beetle generations over. These results suggested that treatment of cattle with the fluazuron pour-on formulation Acatak1 was not detrimental to the selected dung beetle species in any notable way.},
	keywords = {Acaricide, Benzoylphenyl urea, Boophilus decoloratus, dung beetle, ecotoxicological effect, Fluazuron, Onthophagus gazella},
	author = {Kryger, U. and Deschodt, C. and Davis, A. L. V. and Scholtz, Clarke H.}
}
@article {1908,
	title = {Pre- and post-copulatory mate selection mechanisms in an african dung beetle, Circellium bacchus (Coleoptera: Scarabaeidae)},
	journal = {Journal of Insect Behavior},
	volume = {21},
	year = {2008},
	pages = {111-122},
	abstract = {Females of most taxa mate selectively. Mate selection may be: (1) pre- copulatory, involving active female choice and male-male competition, and (2) post- copulatory, with cryptic female choice and sperm competition. Because female dung beetles (Circellium bacchus) invest heavily in parental care by ball-rolling and remaining with developing larva they are, therefore, expected to be highly selective when mating. Mate choice in this species was investigated via behavioral observations and investigations of genital allometry of both sexes, leading to conclusions about the mechanisms of, and male characteristics important in, female choice. Male{\textendash}male competition seems to be crucial in mate selection of C. bacchus, although the females appeared to show no active mate choice. There is a negative allometric relationship between genital size and body size of males as predicted by the {\textquoteleft}one size fits all{\textquoteright} hypothesis (where males have genitalia that fit average-sized females). For the females, no relationship was found between genital size and body size. This might be as a result of the non-sclerotized nature of female genitalia, which may allow for greater morphological plasticity.},
	keywords = {genital allometry ., male, mate choice ., Sexual selection ., {\textendash}male competition ., {\textquoteleft}one size fits all{\textquoteright} hypothesis},
	author = {Le Roux, Elizabeth and Scholtz, Clarke H. and Kinahan, A. A. and Bateman, P. W.}
}
@article {1776,
	title = {Are ball-rolling (Scarabaeini, Gymnopleurini, Sisyphini) and tunnelling scarabaeine dung beetles equally choosy about the size of ingested dung particles?},
	journal = {Ecological Entomology},
	volume = {30},
	year = {2005},
	pages = {700-705},
	abstract = {1. The maximum size of ingested particles was determined in 11 species of ball-rolling, adult dung beetle (Scarabaeidae: Scarabaeinae) by mixing small latex or glass balls of known diameter into their food. The tribes Scarabaeini, Gymnopleurini, and Sisyphini (four, four, and three species respec- tively) were represented, with mean body sizes ranging from 0.33 to 4.0 g fresh weight. 2. Only particles with maximum diameters of 4{\textendash}85 mm were ingested. Hence rollers, like other known beetles feeding on fresh dung, filter out larger, indiges- tible plant fragments and confine ingestion to small particles of higher nutritional value. 3. The maximum diameter of ingested particles increased significantly with body weight, whereas taxon (tribe) had no additional effect. Because big rollers accept larger particles than do tunnellers (which make dung stores for feeding and breeding in the soil immediately below the pat) of similar weight, the slope of the diameter-against-weight regression for rollers was significantly higher than that found earlier for tunnellers. 4. An explanation could be that a typical food ball made by a roller is con- siderably smaller than the amount of dung available to a feeding tunneller of the same size. If the roller were as choosy about particle size as the tunneller, it might not get enough food. This applies to large rollers in particular because their food balls contain a higher proportion of coarse fibres than those made by small species. },
	keywords = {Ball-rollers, Dung beetles, food exploitation, particle feeding, rollers, Scarabaeidae, Scarabaeinae, telecoprids},
	author = {Holter, P. and Scholtz, Clarke H.}
}
@article {1777,
	title = {What do dung beetles eat?},
	journal = {Ecological Entomology},
	volume = {32},
	year = {2007},
	pages = {690-697},
	abstract = {    1.   Most adult coprophagous beetles feed on fresh dung of mammalian herbivores, confining ingestion to small particles with measured maximum diameters from 2 {\textendash} 5 to 130  },
	keywords = {Assimilable carbon, C/N ratio, coprophagous, food exploitation, nitrogen, selective feeding},
	author = {Holter, Peter and Scholtz, Clarke H.}
}
@article {1778,
	title = {Dung feeding in adult scarabaeines (tunnellers and endocoprids): even large dung beetles eat small particles.},
	journal = {Ecological Entomology},
	volume = {27},
	number = {2},
	year = {2002},
	note = {digital copy},
	pages = {169-176},
	keywords = {dung beetle, ecological function, ecosystem service, ecosystem services, feeding, Scarabaeidae},
	author = {Holter, Peter and Scholtz, Clarke H. and Wardhaugh, Keith G.}
}
@article {2105,
	title = {A phylogenetic analysis of dung beetles (Scarabaeinae: Scarabaeidae): unrolling an evolutionary history},
	journal = {Invertebrate Systematics},
	volume = {18},
	year = {2004},
	pages = {53-88},
	author = {Philips, T. Keith and Pretorius, Etheresia and Scholtz, Clarke H.}
}
@article {2115,
	title = {Geometric morphometrics, the metendosternite and its use in phylogenetics of the Scarabaeinae (Coleoptera)},
	journal = {Elytron},
	volume = {14},
	year = {2000},
	pages = {125-148},
	abstract = {Results from a geometric morphometric and cladistic analysis of 20 species (including 4 flightless species) of the Scarabaeinae (Scarabaeidae: Coleoptera) were compared. The metendosternite has largely been neglected in past morphological studies but proved to be very useful because of its easily identifiable landmarks. These landmarks were analyzed in a geometric morphometric analysis, where a Procrustes distance matrix and a dendrogram were produced. A relative warp analysis was also performed on the data and the results of the first two relative warps plotted against one another. The dendrogram clearly distinguished the flightless species from the flighted and it is hypothesized that three lineages of flightless taxa convergently underwent similar alterations in the reduction in metendosternite size. The cladistics study utilized 200 internal and external morphological characters, including 20 based on the metendosternite. Comparison of the cladogram with the dendrogram indicates that geometric morphometrics is a useful technique to hypothesize relationships, especially with structures that may be difficult to interpret in a cladistic analysis. },
	keywords = {cladistic analysis, COLEOPTERA, geometric morphometrics, landmark data, metendosternite, Procrustes distance., relative warp, Scarabaeinae},
	isbn = {0214-1353},
	author = {Pretorius, Resia and Philips, T. Keith and Scholtz, Clarke H.}
}
@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.}
}
@article {2247,
	title = {Trophic associations of a dung beetle assemblage (Scarabaeidae: Scarabaeinae) in a woodland savanna of Botswana},
	journal = {Environmental Entomology},
	volume = {37},
	number = {2},
	year = {2008},
	pages = {431-441},
	abstract = {Species richness and abundance of dung beetles were assessed across a range of bait types that acted as surrogates for the food resources available in Chobe National Park, Botswana. These bait types were comprised of the dung of pig (omnivore), cattle (ruminant herbivore dropping {\TH}ne-{\TH}berd pads), sheep (pellet-dropping ruminant herbivore), and elephant (monogastric, nonru- minant herbivore producing coarse-{\TH}bered droppings), and chicken livers (carrion). Species richness was similar between traps baited with pig, cattle, and elephant dung but was relatively lower in those baited with sheep dung and carrion. In traps baited with pig dung, abundance was relatively greater than in all other bait types. A cluster analysis of species abundance distributions for the 30 most abundant species identi{\TH}ed four different patterns of bait type association at a 60\% level of similarity. All but 1 of the 15 species in cluster A were attracted primarily to the dung of omnivores and pad-dropping ruminant herbivores (pig and cattle). All seven species of cluster B were attracted primarily to coarse-{\TH}bered, nonruminant herbivore dung (elephant). All four species of cluster C were primarily carrion and pig dung associated, whereas all four species of cluster D were carrion specialists. In conclusion, the most abundant species were attracted to all bait types, but most species were largely specialized to different dung types or carrion, with dung attracting the majority of the fauna in terms of both species richness and abundance.},
	keywords = {African savannas, bait types, Botswana, Scarabaeinae, trophic associations},
	author = {Tshikae, B. Power and Davis, Adrian L. V. and Scholtz, Clarke H.}
}
