@article {1570,
	title = {Phylogenetic evidence for an association between tunneling behavior and the evolution of horns in dung beetles (Coleoptera : Scarabaeidae : Scarabaeinae)},
	journal = {The Coleopterists Bulletin},
	volume = {60},
	number = {5},
	year = {2006},
	note = {Times Cited: 0Suppl. S},
	month = {Dec},
	pages = {47-56},
	type = {monograph},
	abstract = {Dung beetles employ numerous behavioral strategies to sequester dung away from other insects, and these have been broadly grouped into two categories: species that dig tunnels beneath the dung (tunnelers) and species that roll dung on the surface of the soil (rollers). Many species also are armed with rigid exoskeletal outgrowths called horns. Horns function as weapons, and horn sizes can be extreme. One widespread pattern within dung beetles is that tunneling species often have horns, whereas rolling species almost always do not, suggesting that residing (and fighting) inside tunnels at the dung deposition site may be an important ecological prerequisite for the evolution of horns in dung beetles. Here, we test explicitly for an historical association between tunneling behavior and the evolution of horns using a recent phylogeny for the scarabaeine dung beetles. We show that all eight of the independent gains of horns included in our analyses occurred on branches of the phylogeny reconstructed as tunneling, and that one of the three evolutionary losses of horns occurred on a branch that had lost tunneling behavior. We interpret this as evidence for a biologically meaningful association between tunneling behavior and the evolution of enlarged or exaggerated weapons such as horns, supporting the ideas of Eberhard and others that beetle horns may be most "beneficial" when used within the confines of restricted spaces such as burrows or tunnels.},
	keywords = {ALTERNATIVE REPRODUCTIVE TACTICS, body-size, DIMORPHISM, FEMALE CHOICE, FIDDLER-CRAB, FIGHTS, FOOD RELOCATION, onthophagus-taurus, SELECTION, SEXUAL, TRADE-OFFS},
	isbn = {0010-065X},
	url = {<Go to ISI>://000243611100005 },
	author = {Emlen, Douglas J. and Philips, T. Keith}
}
@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 {2087,
	title = {Food relocation and nesting behavior of the Argentinian dung beetle genus Eucranium and comparison with the southwest African Scarabaeus (Pachysoma) (Coleoptera: Scarabaeidae: Scarabaeinae)},
	journal = {Revista Sociedad Entomologica Argentina},
	volume = {64},
	number = {1-2},
	year = {2005},
	pages = {53-59},
	abstract = {The food relocation and nesting behavior of the South American genus Eucranium Brullé is described based on observations carried out under natural conditions. Food preference, food transportation, period of surface activity, and habitat preference are described for two species, E. arachnoides Brullé and E. planicolle Burmeister. The biology and food relocation behavior of Eucranium is compared with that of the morphologically similar South African subgenus Scarabaeus (Pachysoma) MacLeay. The convergent adaptations to psamophilous habitats of Eucranium and S. (Pachysoma) are discussed. },
	keywords = {behavior, Dung beetles, endemism, FOOD RELOCATION, xeric habitat},
	author = {Ocampo, Federico C. and Philips, T. Keith}
}
@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.}
}
