@article {1409,
	title = {Estimating the number of species not yet described and their characteristics: the case of Western Palaearctic dung beetle species (Coleoptera, Scarabaeoidea)},
	journal = {Biodiversity and Conservation},
	volume = {12},
	number = {1},
	year = {2003},
	note = {Times Cited: 0Cited Reference Count: 99Cited References:      *STATS INC, 1999, STATISTICA COMP PROG     AKAIKE H, 1983, B INT STAT I, V50, P277     ALLSOPP PG, 1997, J BIOGEOGR, V24, P717     ANDERSON MJ, 1998, AUST J ECOL, V23, P158     ARNETT RH, 1967, ANN ENTOMOLOGICAL SO, V60, P162     ARNONE M, 1993, NATURALISTA SICILI 4, V17, P271     ARNONE M, 1995, NATURALISTA SICILIAN, V19, P447     BARAUD J, 1992, COLEOPTERES SCARABAE     BARAUD J, 1985, COLEOPTERES SCARABAE     BARROWCLOUGH GF, 1992, SYSTEMATICS ECOLOGY, P1     BENNETT KD, 1991, J BIOGEOGR, V18, P103     BLACKBURN TM, 1995, J BIOGEOGR, V22, P7     BORCARD D, 1992, ECOLOGY, V73, P1045     CARPANETO GM, 1985, ANIMALIA, V12, P87     CARPANETO GM, 1997, ANN MUSEO CIVICO STO, V91, P511     CARPANETO GM, 1995, BIOGEOGRAPHIA, V18, P441     CARPANETO GM, 1995, COLEOPTERA POLYPHAGA, V5     CARPANETO GM, 1979, FRAGMENTA ENTOMOLOGI, V15, P111     CARPANETO GM, 1976, FRAGMENTA ENTOMOLOGI, V12, P255     CARPANETO GM, 1980, LAVORI SOC ITALIANA, V8, P675     CHIKATUNOV V, 1997, KLAPALEKIANA, V33, P37     COPAS JB, 1983, J ROY STAT SOC B MET, V45, P311     COTTERILL FPD, 1995, BIODIVERS CONSERV, V4, P183     CRAWLEY MJ, 1993, GLIM ECOLOGISTS     DANIEL C, 1980, FITTING EQUATIONS DA     DELLACASA G, 1981, ANN MUSEO CIVICO STO, V83, P443     DELLACASA G, 1992, FRUSTULA ENTOMOLOGIC, V15, P119     DELLACASA G, 1990, FRUSTULA ENTOMOLOGIC, V13, P19     DELLACASA M, 1995, MEMORIE SOC ENTOMOLO, V74, P159     DELLACASA M, 1991, MEMORIE SOC ENTOMOLO, V70, P3     DELLACASA M, 1988, MEMORIE SOC ENTOMOLO, V67, P291     DELLACASA M, 1988, MEMORIE SOC ENTOMOLO, V66, P1     DELLACASA M, 1988, MEMORIE SOC ITALIANA, V67, P1     DELLACASA M, 1987, NATURALISTA SICILI 4, V11, P1     DERKSEN S, 1992, BRIT J MATH STAT PSY, V45, P265     DIAMOND JM, 1985, NATURE, V315, P538     DOBSON A, 1999, INTRO GEN LINEAR MOD     FLATHER CH, 1996, J BIOGEOGR, V23, P155     FRANK JH, 1979, COLEOPTERISTS B, V33, P133     GASTON KJ, 1994, BIODIV LETT, V2, P16     GASTON KJ, 1995, BIODIVERS CONSERV, V4, P119     GASTON KJ, 1996, BIODIVERSITY BIOL NU     GASTON KJ, 1994, BIOL CONSERV, V67, P37     GASTON KJ, 1995, BIOL J LINN SOC, V55, P225     GASTON KJ, 1991, CONSERV BIOL, V5, P283     GASTON KJ, 1991, ECOL ENTOMOL, V16, P505     GASTON KJ, 1993, HYMENOPTERA BIODIVER     GASTON KJ, 1992, NATURE, V356, P281     GASTON KJ, 1993, P ROY SOC LOND B BIO, V251, P139     HAMMOND P, 1992, GLOBAL BIODIVERSITY     HANSKI I, 1983, ACTA ENTOMOL FENN, V42, P36     HANSKI I, 1986, ACTA OECOL-OEC GEN, V7, P171     HANSKI I, 1991, DUNG BEETLE ECOLOGY, P75     HEARD SB, 2000, P ROY SOC LOND B BIO, V267, P613     HEWITT GM, 1996, BIOL J LINN SOC, V58, P247     HEYWOOD VH, 1995, GLOBAL BIODIVERSITY     HOLLANDE A, 1998, MONOGRAPH MUSEO REGI, V21     HUGHES JB, 1997, SCIENCE, V278, P689     ILLIES J, 1983, ANNU REV ENTOMOL, V28, P391     KRAL D, 1997, ACTA SOC ZOOLOGICAE, V61, P129     KRAL D, 1996, FOLIA HEYROVSKYANA, V4, P49     KRELL FT, 2000, NATURE, V405, P507     LAWTON JH, 1995, EXTINCTION RATES     LEGENDRE P, 1998, NUMERICAL ECOLOGY     LOBO JM, 2000, CAN ENTOMOL, V132, P307     LOBO JM, 1993, ECOL MED, V19, P29     LOBO JM, 1993, EUR J ENTOMOL, V90, P235     LOPEZCOLON JI, 1995, GIORNALE ITALIANO EN, V7, P355     LUMARET JP, 1987, ACTA ZOOL MEX, V24, P1     LUMARET JP, 1996, BIODIVERSITY LETT, V3, P192     LUMARET JP, 1996, COLLECTION PATRIMOIN, V26, P1     LUMARET JP, 1990, INVENTAIRES FAUNE FL, V1, P1     MACKAUER M, 1958, MEMORIE SOC ENTOMOLO, V37, P46     MACNALLY R, 2000, BIODIVERS CONSERV, V9, P655     MARTINPIERA F, 1992, NOUVELLE REV ENTOMOL, V9, P15     MAY RM, 1988, SCIENCE, V241, P1441     MAYR E, 1993, BIOL SYSTEMATICS STA     MCCULLAGH P, 1989, GEN LINEAR MODELS     MEDELLIN RA, 1999, CONSERV BIOL, V13, P143     MIELKE PW, 1974, WATER RESOUR RES, V10, P223     NETER J, 1985, APPL LINEAR STAT MOD     NICHOLLS AO, 1989, BIOL CONSERV, V50, P51     OBRIEN CW, 1979, COLEOPTERISTS B, V33, P151     PALMER M, 1988, REUNION CIENTIFICA S, V28, P155     PATTERSON BD, 1994, BIODIVERSITY LETT, V2, P79     PATTERSON BD, 2000, DIVERSITY DISTRIBUTI, V6, P145     PITTINO R, 1996, FRAGMENTA ENTOMOLOGI, V27, P355     PURVIS A, 2000, SCIENCE, V288, P328     QINGHONG L, 1995, WATER AIR SOIL POLL, V85, P1587     SCHOLTZ CH, 1990, J NAT HIST, V24, P1027     SKIDMORE P, 1991, INSECTS BRIT COW DUN     SOBERON J, 1993, CONSERV BIOL, V7, P480     STEYSKAL GC, 1965, SCIENCE, V149, P880     SUTTON SL, 1991, CONSERVATION INSECTS, P405     TABERLET P, 1998, MOL ECOL, V7, P453     WHITE RE, 1979, COLEOPTERISTS B, V33, P167     WHITE RE, 1975, COLEOPTERISTS B, V29, P281     ZIANI S, 1997, B SOC ENTOMOLOGICA I, V128, P197     ZIANI S, 1997, BIOCOSME MESOGEEN, V14, P1EnglishArticle620EQBIODIVERS CONSERV},
	month = {Jan},
	pages = {147-166},
	abstract = {To determine the degree of completion of Western Palaearctic dung beetle species inventory, as well as to ascertain some of the main geographical and morphological characteristics associated with the probability of new species description, the available taxonomic and biogeographic information of the three Scarabaeoidea families that are associated with dung was analyzed. An asymptotic fit of the cumulative number of described species is used to estimate the approximate number of species not yet described. Variation partitioning and hierarchical decomposition techniques were used to explore the influence of body size, geographical range size and geographical location variables on the process of species description. Results indicate that the inventories of Western Palaearctic Scarabaeidae and Geotrupidae species are almost complete, but that around 16\% of Aphodiidae species have yet to be described. As the joint effect of geographical distribution and location is the main determinant of the species description rate in this family, it was predicted that new dung beetle species would be found in smaller areas of the southern and eastern Western Palaearctic region. The species body size accounted for a negligible fraction in the variation of the year of description. Both the geographical range size and the geographical location are essential variables negatively correlated with the year of species description, whose pure and combined effect can account for high percentages in the year of description variation (from 41 to 56\%). The location of most of the taxonomic workforce in areas where broader-ranged generalist species prevail in the assemblages is the probable reason for this pattern.},
	keywords = {accumulation functions, body-size, conservation, discovery, diversity, evolutionary history, extinction, geographic location, geographic range size, predi, probability, probability of, regression, species description rate, taxonomy, variation partitioning},
	url = {<Go to ISI>://000179521400010},
	author = {Cabrero-Sanudo, Francisco-Jose and Lobo, Jorge M.}
}
@article {1565,
	title = {Phylogenetic evidence for an association between tunneling behavior and the evolution of horns in dung beetles (Coleoptera : Scarabaeidae : Scarabaeinae)},
	journal = {Coleopterists Bulletin},
	volume = {60},
	number = {5},
	year = {2006},
	note = {Times Cited: 0Suppl. S},
	month = {Dec},
	pages = {47-56},
	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, D. J.}
}
@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 {1817,
	title = {Maternal and paternal effects on offspring phenotype in the dung beetle Onthophagus taurus},
	journal = {Evolution},
	volume = {54},
	number = {3},
	year = {2000},
	note = {Using Smart Source ParsingArticleEnglish},
	pages = {936-941},
	abstract = {Parents often have important influences on the development of traits in their offspring. One mechanism by which parents are able to influence offspring phenotype is through the level of care they provide. In onthophagine dung beetles, parents typically provision their offspring by packing dung fragments into a brood mass. Onthophagus taurus males can be separated into two discrete morphs: Large, "major" males have head horns, whereas "minor" males are hornless. Here we show that a switch in parental provisioning strategies adopted by males coincides with the switch in male morphology. Male provisioning results in the production of heavier brood masses than females will produce alone. However, unlike females in which the level of provisioning increases with body size in a continuous manner, the level of provisioning provided by males represents an "all-or-none" tactic with all major males providing a fixed level of provisioning irrespective of their body size. Offspring size is determined largely by the quantity of dung provided to the developing larvae so that paternal and maternal provisioning affects the body size and horn size of offspring produced. The levels of provisioning by individual parents are significantly repeatable, suggesting paternal and maternal effects as candidate indirect genetic effects in the evolution of horn size in the genus Onthophagus.},
	keywords = {Animalia-, Animals-, Arthropoda-, Arthropods-, Behavior-, body-size, brood-mass, Coleoptera-: Insecta-, Evolution-and-Adaptation, horn-: size-, Insects-, Invertebrata-, Invertebrates-, male-, male-p, Onthophagus-taurus [dung-beetle] (Coleoptera-): female-},
	author = {Hunt, J. and Simmons, L. W.}
}
@article {2033,
	title = {Proximate determination of male horn dimorphism in the beetle Onthophagus taurus (Coleoptera: Scarabaeidae)},
	journal = {Journal of Evolutionary Biology},
	volume = {12},
	number = {1},
	year = {1999},
	note = {Jan., 1999ArticleEnglish},
	pages = {27-37},
	abstract = {The existence of discrete phenotypic variation within one sex poses interesting questions regarding how such intrasexual polymorphisms are produced and modified during the course of evolution. Approaching these kinds of questions requires insights into the genetic architecture underlying a polymorphism and an understanding of the proximate mechanisms determining phenotype expression. Here we explore the genetic underpinnings and proximate factors influencing the expression of beetle horns - a dramatic sexually selected trait exhibiting intramale dimorphism in many species. Two relatively discrete male morphs are present in natural populations of the dung beetle Onthophagus taurus (Scarabaeidae, Onthophagini). Males exceeding a critical body size develop a pair of long, curved horns on their heads, while those smaller than this critical body size remain essentially hornless. We present results from laboratory breeding experiments designed to assess the relative importance of inherited and environmental factors as determinants of male morphology. Using father-son regressions, our findings demonstrate that horn length and body size of male progeny are not predicted from paternal morphology. instead, natural variation in an environmental factor, the amount of food available to larvae, determined both the body sizes exhibited by males as adults and the presence or absence of horns. The nonlinear scaling relationship between the body size and horn length of males bred in the laboratory did not differ from the pattern of variation present in natural populations, suggesting that nutritional conditions account for variation in male morphology in natural populations as well. We discuss our results by extending ideas proposed to explain the evolution of conditional expression of alternative phenotypes in physically heterogeneous environments toward incorporating facultative expression of secondary sexual traits. We use this synthesis to begin characterizing the potential origin and subsequent evolution of facultative horn expression in onthophagine beetles.},
	keywords = {allometry-, Animalia-, Animals-, Arthropoda-, Arthropods-, body-size, Coleoptera-: Insecta-, Evolution-and-Adaptation, horn-dimorphism, Insects-, Invertebrata-, Invertebrates-, mor, Onthophagus-taurus [dung-beetle] (Coleoptera-): male-, Population-Studies},
	author = {Moczek, A. P. and Emlen, D. J.}
}
