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9 - Horns and the role of development in the evolution of beetle contests

Published online by Cambridge University Press:  05 June 2013

By
Emilie C. Snell-Rood  and
Armin P. Moczek
Edited by
Ian C. W. Hardy  and
Mark Briffa
Ian C. W. Hardy
Affiliation:
University of Nottingham
Mark Briffa
Affiliation:
University of Plymouth
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Summary

Summary

Contests between beetles have attracted the attention of biologists and natural historians for a long time. Contests almost always involve males competing for females or for access to resources crucial to female reproduction. With only a few exceptions, beetle contests typically involve non-injurious trials of strength. In this chapter we begin by reviewing the literature on beetle contests with an emphasis on how ecological conditions and resource availability have shaped the nature of contests in diverse beetle families. We then focus on two inter-related aspects of the biology of beetle contests that have received particular and recent attention: the evolution of alternative reproductive tactics and the origin and diversification of weaponry. We describe the nature of alternative reproductive tactics in a range of beetle families, and explore the developmental and physiological mechanisms that have mediated contest diversification within and between species. We then explore the evolutionary developmental biology of weaponry in beetles, focusing on the elaborate horns of scarabs. Specifically, we describe how horns are built during development, and explore the relative contributions of conserved developmental mechanisms and novel developmental properties to the origin and diversification of horns. We then highlight several instances in which developmental studies have assisted our understanding of the evolution and ecology of contest behaviour, such as the ancestry of alternative phenotypes and sexual dimorphisms, the costs of weaponry and constraints and biases in the diversification of beetle weapons. Combining rich morphological and behavioural diversity with a diverse genetic and developmental toolbox, beetles emerge as a powerful model system in which to explore the causes, mechanisms and consequences of contest behaviour in nature.

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Animal Contests , pp. 178 - 198
Publisher: Cambridge University Press
Print publication year: 2013

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References

Abbott, JC, Dunbrack, RL & Orr, CD (1985) The interaction of size and experience in dominance relationships of juvenile steelhead trout(Salmo gairdneri). Behaviour, 92, 241–253.Google Scholar
Andersson, M (1994) Sexual Selection. Princeton, NJ: Princeton University Press.Google Scholar
Angelini, DR, Kikuchi, M & Jockusch, EL (2009) Genetic patterning in the adult capitate antenna of the beetleTribolium castaneum. Developmental Biology, 327, 240–251.CrossRefGoogle ScholarPubMed
Arrow, GJ (1951) Horned Beetles. The Hague: Junk Publishers.CrossRefGoogle Scholar
Brown, L & Bartalon, J (1986) Behavioral correlates of male morphology in a horned beetle. American Naturalist, 127, 565–570.CrossRefGoogle Scholar
Chamorro-Florescano, IA & Favila, ME (2008) Male reproductive status affects contest outcome during nidification in Canthon cyanellus cyanellus LeConte (Coleoptera: Scarabaeidae). Behaviour, 145, 1811–1821.CrossRefGoogle Scholar
Chase, ID, Bartolomeo, C & Dugatkin, LA (1994) Aggressive interactions and inter-contest interval – How long do winners keep winning?Animal Behaviour, 48, 393–400.CrossRefGoogle Scholar
Conner, J (1988) Field measurements of natural and sexual selection in the fungus beetleBolitotherus cornutus. Evolution, 42, 736–749.CrossRefGoogle ScholarPubMed
Conner, J (1989) Density-dependent sexual selection in the fungus beetle, Bolitotherus cornutus. Evolution, 43, 1378–1386.CrossRefGoogle ScholarPubMed
Convey, P (1989) Influences on the choice between territorial and satellite behaviour in male Libellula quadrimaculata Linn. (Odonata: Libellulidae). Behaviour, 109, 125–141.CrossRefGoogle Scholar
Cook, DF (1990) Differences in courtship, mating and postcopulatory behaviour between male morphs of the dung beetles Onthophagus binodis Thunberg (Coleoptera: Scarabaeidae). Animal Behaviour, 40, 428–436.CrossRefGoogle Scholar
Crespi, BJ (1988) Adaptation, compromise, and constraint: the development, morphometrics, and behavioral basis of a fighter–flier polymorphism in male Hoplothrips karnyi (Insecta: Thysanoptera). Behavioural Ecology and Sociobiology, 23, 93–104.CrossRefGoogle Scholar
Darwin, C (1874) The Descent of Man, and Selection in Relation to Sex. New York: A. L. Burt.CrossRefGoogle Scholar
Dominey, WJ (1984) Alternative mating tactics and evolutionarily stable strategies. American Zoologist, 24, 385–396.CrossRefGoogle Scholar
Eberhard, WG (1977) Fighting behavior of Golofa porter beetles (Scarabeidae: Dynastinae). Psyche, 83, 292–298.CrossRefGoogle Scholar
Eberhard, WG (1979) The function of horns in Podischnus agenor (Dynastinae) and other beetles. In: Blum, MS & Blum, NA (eds.) Sexual Selection and Reproductive Competition in Insects, pp. 231–258. New York, NY: Academic Press.Google Scholar
Eberhard, WG (1980) Horned beetles. Scientific American, 242, 166–182.CrossRefGoogle Scholar
Eberhard, WG (1981) The natural history of Doryphora sp. (Coleoptera, Chrysomelidae) and the function of its sterna horn. Annals of the Entomological Society of America, 74, 445–448.CrossRefGoogle Scholar
Eberhard, WG (1982) Beetle horn dimorphism: Making the best of a bad lot. American Naturalist, 119, 420–426.CrossRefGoogle Scholar
Eberhard, WG (1983) Behavior of adult bottle brush weevils (Rhinostomus barbirostris) (Coleoptera: Curculionidae). Revista de Biologia Tropical, 31, 233–244.Google Scholar
Eberhard, WG (1985) Sexual Selection and Animal Genitalia. Cambridge, MA: Harvard University Press.CrossRefGoogle Scholar
Eberhard, WG (1987) Use of horns in fights by the dimorphic males of Ageopsis nigricollis (Coleoptera, Scarabeidae, Dynastinae). Journal of the Kansas Entomological Society, 60, 504–509.Google Scholar
Eberhard, WG & Garcia-C, JM (2000) Ritual jousting by horned Parisoschoenus expositus weevils (Coleoptera: Curculionidae, Baridinae). Psyche, 103, 55–84.CrossRefGoogle Scholar
Eberhard, WG & Gutiérrez, EE (1991) Male dimorphism in beetles and earwigs and the question of developmental constraints. Evolution, 45, 18–28.CrossRefGoogle Scholar
Eberhard, WG, Garcia-C, JM & Lobo, J (2000) Size-specific defensive structures in a horned weevil confirm a classic battle plan: avoid fights with larger opponents. Proceedings of the Royal Society of London B, 267, 1129–1134.CrossRefGoogle Scholar
Emlen, DJ (1994) Environmental control of horn length dimorphism in the beetle Onthophagus acuminatus (Coleptera: Scarabaeidae). Proceedings of the Royal Society of London B, 256, 131–136.CrossRefGoogle Scholar
Emlen, DJ (1996) Artificial selection on horn length–body size allometry in the horned beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae). Evolution, 50, 1219–1230.CrossRefGoogle Scholar
Emlen, DJ (1997) Alternative reproductive tactics and male-dimorphism in the horned beetle Onthophagus acuminatus (Coleoptera: Scarabaeidae). Behavioral Ecology and Sociobiology, 41, 335–341.CrossRefGoogle Scholar
Emlen, DJ (2001) Costs and the diversification of exaggerated animal structures. Science, 291, 1534–1536.CrossRefGoogle ScholarPubMed
Emlen, DJ (2008) The evolution of animal weapons. Annual Review of Ecology Evolution and Systematics, 39, 387–413.CrossRefGoogle Scholar
Emlen, DJ & Nijhout, HF (1999) Hormonal control of male horn length dimorphism in the dung beetle Onthophagus taurus (Coleoptera: Scarabaeidae). Journal of Insect Physiology, 45, 45–53.CrossRefGoogle Scholar
Emlen, DJ & Nijhout, HF (2000) The development and evolution of exaggerated morphologies in insects. Annual Review of Entomology, 45, 661–708.CrossRefGoogle ScholarPubMed
Emlen, DJ & Nijhout, HF (2001) Hormonal control of male horn length dimorphism in Onthophagus taurus (Coleoptera: Scarabaeidae): A second critical period of sensitivity to juvenile hormone. Journal of Insect Physiology, 47, 1045–1054.CrossRefGoogle ScholarPubMed
Emlen, DJ & Philips, TK (2006) Phylogenetic evidence for an association between tunneling behavior and the evolution of horns in dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Coleopterists Society Monograph, 5, 47–56.Google Scholar
Emlen, DJ, Hunt, J & Simmons, LW (2005a) Evolution of sexual dimorphism and male dimorphism in the expression of beetle horns: Phylogenetic evidence for modularity, evolutionary lability, and constraint. American Naturalist, 166, S42–S68.CrossRefGoogle ScholarPubMed
Emlen, DJ, Marangelo, J, Ball, B & Cunningham, CW (2005b) Diversity in the weapons of sexual selection: Horn evolution in the beetle genus Onthophagus (Coleoptera: Scarabaeidae). Evolution, 59, 1060–1084.CrossRefGoogle Scholar
Emlen, ST & Oring, LW (1977) Ecology, sexual selection, and the evolution of mating systems. Science, 197, 215–223.CrossRefGoogle ScholarPubMed
Enquist, M & Leimar, O (1983) Evolution of fighting behaviour, decision rules and assessment of relative strength. Journal of Theoretical Biology, 102, 387–410.CrossRefGoogle Scholar
Enquist, M & Leimar, O (1987) Evolution of fighting behaviour – The effect of variation in resource value. Journal of Theoretical Biology, 127, 187–205.CrossRefGoogle Scholar
Enquist, M & Leimar, O (1990) The evolution of fatal fighting. Animal Behaviour, 39, 1–9.CrossRefGoogle Scholar
Espmark, Y (1964) Studies in dominance–subordinate relationships in a group of semi-domestic reindeer (Rangifer tarandus L.). Animal Behaviour, 12, 420–426.CrossRefGoogle Scholar
Franz, NM (2003) Mating behaviour of Staminodeus vectoris (Coleoptera: Curculionidae), and the value of systematics in behavioral studies. Journal of Natural History, 37, 1727–1750.CrossRefGoogle Scholar
Gadgil, M (1972) Male dimorphism as a consequence of sexual selection. American Naturalist, 106, 574–580.CrossRefGoogle Scholar
Goldsmith, SK (1987) The mating system and alternative reproductive behaviours of Dendrobias mandibularis (Coleoptera: Cerambycidae). Behavioural Ecology and Sociobiology, 20, 111–115.CrossRefGoogle Scholar
Gross, MR (1996) Alternative reproductive strategies and tactics: Diversity within sexes. Trends in Ecology and Evolution, 11, 92–98.CrossRefGoogle ScholarPubMed
Hamilton, WD (1979) Wingless and fighting males in fig wasps and other insects. In: Blum, MS & Blum, NA (eds.) Sexual Selection and Reproductive Competition in Insects, pp. 167–220. New York, NY: Academic Press.Google Scholar
Hammerstein, P (1981) The role of asymmetries in animal contests. Animal Behaviour, 29, 193–205.CrossRefGoogle Scholar
Hanley, RS (2001) Mandibular allometry and male dimorphism in a group of obligately mycophagous beetles (Insecta: Coleoptera: Staphylinidae: Oxyporinae). Biological Journal of the Linnean Society, 72, 451–459.CrossRefGoogle Scholar
Hongo, Y (2003) Appraising behavior during male–male interaction in the Japanese horned beetle Trypoxylus dichotomus septentrionalis (Kono). Behaviour, 140, 501–517.CrossRefGoogle Scholar
Howard, RS (1955) The occurrence of fighting behavior in the grain beetle Tenebrio molitor with the possible formation of a dominance hierarchy. Ecology, 36, 281–285.CrossRefGoogle Scholar
Hsu, YY & Wolf, LL (2001) The winner and loser effect: What fighting behaviours are influenced?Animal Behaviour, 61, 777–786.CrossRefGoogle Scholar
Hughes, AL & Hughes, MK (1987) Asymmetric contests among sawyer beetles (Cerambycidae: Monochamus notatus and Monochamus scutellatus). Canadian Journal of Zoology, 65, 823–827.CrossRefGoogle Scholar
Hunt, J & Simmons, LW (2000) Maternal and paternal effects on offspring phenotype in the dung beetle Onthophagus taurus. Evolution, 54, 936–941.CrossRefGoogle ScholarPubMed
Hunt, J & Simmons, LW (2001) Status-dependent selection in the dimorphic beetle Onthophagus taurus. Proceedings of the Royal Society of London B, 268, 2409–2414.CrossRefGoogle ScholarPubMed
Johnson, LK (1982) Sexual selection in a brentid weevil. Evolution, 36, 251–262.CrossRefGoogle Scholar
Judge, KA & Bonanno, VL (2008) Male weaponry in a fighting cricket. PLoS ONE, 3, 10.CrossRefGoogle Scholar
Karino, K, Niiyama, H & Chiba, M (2005) Horn length is the determining factor in the outcomes of escalated fights among male Japanese horned beetles, Allomyrina dichotoma L. (Coleoptera : Scarabaeidae). Journal of Insect Behavior, 18, 805–815.CrossRefGoogle Scholar
Kawano, K (2002) Character displacement in giant rhinoceros beetles. American Naturalist, 159, 255–271.CrossRefGoogle ScholarPubMed
Kawano, K (2006) Sexual dimorphism and the making of oversized male characters in beetles (Coleoptera). Annals of the Entomological Society of America, 99, 327–341.CrossRefGoogle Scholar
Kijimoto, T, Andrews, J & Moczek, AP (2010) Programmed cell death shapes the expression of horns within and between species of horned beetles. Evolution and Development, 12, 449–458.CrossRefGoogle ScholarPubMed
King, A & Fashing, N (2007) Infanticidal behavior in the subsocial beetle Odontotaenius disjunctus (Illiger) (Coleoptera: Passalidae). Journal of Insect Behavior, 20, 527–536.CrossRefGoogle Scholar
Kruse, KC & Switzer, PV (2007) Physical contests for females in the Japanese beetle, Popillia japonica. Journal of Insect Science, 7, 34.Google ScholarPubMed
Longair, RW (2004) Tusked males, male dimorphism and nesting behavior in a subsocial Afrotropical wasp, Synagris cornuta, and weapons and dimorphism in the genus (Hymenoptera: Vespidae: Eumeninae). Journal of the Kansas Entomological Society, 77, 528–557.CrossRefGoogle Scholar
Lloyd, JE (1979) Sexual selection in luminescent beetles. In: Blum, MS & Blum, NA (eds.) Sexual Selection and Reproductive Competition in Insects, pp. 293–342. New York, NY: Academic Press.Google Scholar
Madewell, R & Moczek, AP (2006) Horn possession reduces maneuverability in the horn-polyphenic beetle, Onthophagus nigriventris. Journal of Insect Science, 6, article 21.Google ScholarPubMed
Mason, LG (1980) Sexual selection and the evolution of pair bonding in solider beetles. Evolution, 34, 174–180.CrossRefGoogle Scholar
Maynard Smith, JM (1974) Theory of games and evolution of animal conflicts. Journal of Theoretical Biology, 47, 209–221.CrossRefGoogle Scholar
Maynard Smith, JM (1979) Game theory and the evolution of behaviour. Proceedings of the Royal Society of London B, 205, 475–488.CrossRefGoogle Scholar
McLachlan, AJ & Allen, DF (1987) Male mating success in Diptera: Advantages of small size. Oikos, 48, 11–14.CrossRefGoogle Scholar
McLain, DK (1982) Behavioral and morphological correlates of male dominance and courtship persistence in the blister beetle Epicauta pennsylvanica (Coleoptera: Meoloidae). American Midland Naturalist, 107, 396–403.CrossRefGoogle Scholar
Miller, KB & Wheeler, QD (2005) Asymmetrical male mandibular horns and mating behavior in Agathidium Panzer (Coleoptera: Leiodidae). Journal of Natural History, 39, 779–792.CrossRefGoogle Scholar
Moczek, AP (2003) The behavioral ecology of threshold evolution in a polyphenic beetle. Behavioral Ecology, 14, 841–854.CrossRefGoogle Scholar
Moczek, AP (2005) The evolution and development of novel traits, or how beetles got their horns. BioScience, 11, 935–951.Google Scholar
Moczek, AP (2006a) Integrating micro- and macroevolution of development through the study of horned beetles. Heredity, 97, 168–178.CrossRefGoogle Scholar
Moczek, AP (2006b) Pupal remodeling and the development and evolution of sexual dimorphism in horned beetles. American Naturalist, 168, 711–729.CrossRefGoogle ScholarPubMed
Moczek, AP (2009) Developmental plasticity and the origins of diversity: a case study on horned beetles. In: Ananthakrishnan, TN & Whitman, D (eds.) Phenotypic Plasticity in Insects: Mechanisms and Consequences, pp. 27–80. Plymouth, UK: Science Publishers Inc.Google Scholar
Moczek, AP & Emlen, DJ (1999) Proximate determination of male horn dimorphism in the beetle Onthophagus taurus (Coleoptera: Scarabaeidae). Journal of Evolutionary Biology, 12, 27–37.CrossRefGoogle Scholar
Moczek, AP & Emlen, DJ (2000) Male horn dimorphism in the scarab beetle, Onthophagus taurus: Do alternative reproductive tactics favour alternative phenotypes?Animal Behaviour, 59, 459–466.CrossRefGoogle ScholarPubMed
Moczek, AP & Nagy, LM (2005) Diverse developmental mechanisms contribute to different levels of diversity in horned beetles. Evolution and Development, 7, 175–185.CrossRefGoogle ScholarPubMed
Moczek, AP & Nijhout, HF (2002) Developmental mechanisms of threshold evolution in a polyphenic beetle. Evolution and Development, 4, 252–264.CrossRefGoogle Scholar
Moczek, AP & Nijhout, HF (2003) Rapid evolution of a polyphenic threshold. Evolution and Development, 5, 259–268.CrossRefGoogle ScholarPubMed
Moczek, AP & Nijhout, HF (2004) Trade-offs during the development of primary and secondary sexual traits in a horned beetle. American Naturalist, 163, 184–191.CrossRefGoogle Scholar
Moczek, AP & Rose, DJ (2009) Differential recruitment of limb patterning genes during development and diversification of beetle horns. Proceedings of the National Academy of Sciences USA, 106, 8992–8997.CrossRefGoogle ScholarPubMed
Moczek, AP, Brühl, CA & Krell, F-T (2004) Linear and threshold-dependent expression of secondary sexual traits in the same individual: Insights from a horned beetle (Coleoptera: Scarabaeidae). Biological Journal of the Linnean Society, 83, 473–480.CrossRefGoogle Scholar
Moczek, AP, Rose, D, Sewell, W, & Kesselring, BR (2006a) Conservation, innovation, and the evolution of horned beetle diversity. Development Genes and Evolution, 216, 655–665.CrossRefGoogle ScholarPubMed
Moczek, AP, Cruickshank, TE, & Shelby, JA (2006b) When ontogeny reveals what phylogeny hides: Gain and loss of horns during development and evolution of horned beetles. Evolution, 60, 2329–2341.CrossRefGoogle ScholarPubMed
Nijhout, HF & Emlen, DJ (1998) Competition among body parts in the development and evolution of insect morphology. Proceedings of the National Academy of Sciences USA, 95, 3685–3689.CrossRefGoogle ScholarPubMed
Okada, K & Miyatake, T (2007) Ownership-dependent mating tactics of minor males of the beetle Librodor japonicus (Nitidulidae) with intra-sexual dimorphism of mandibles. Journal of Ethology, 25, 255–261.CrossRefGoogle Scholar
Okada, K & Miyatake, T (2009) Genetic correlations between weapons, body shape and fighting behaviour in the horned beetleGnatocerus cornutus. Animal Behaviour, 77, 1057–1065.CrossRefGoogle Scholar
Okada, K & Miyatake, T (2010) Effect of losing on male fights of broad-horned flour beetle, Gnatocerus cornutus. Behavioral Ecology and Sociobiology, 64, 361–369.CrossRefGoogle Scholar
Okada, K, Miyanoshita, A & Miyatake, T (2006) Intra-sexual dimorphism in male mandibles and male aggressive behavior in the broad-horned flour beetle Gnatocerus cornutus (Coleoptera: Tenebrionidae). Journal of Insect Behavior, 19, 457–467.CrossRefGoogle Scholar
Okada, K, Miyatake, T, Nomura, Y & Kuroda, K (2008) Fighting, dispersing, and sneaking: Body-size dependent mating tactics by male Librodor japonicus beetles. Ecological Entomology, 33, 269–275.CrossRefGoogle Scholar
O’Neill, KM, Evans, HE & O’Neill, RP (1989) Phenotypic correlates of mating success in the sand wasp Bembecinus quinquespinosus (Hymenoptera: Sphecidae). Canadian Journal of Zoology, 67, 2557–2568.CrossRefGoogle Scholar
Otronen, M (1988a) The effect of body size on the outcome of fights in burying beetles (Nicrophorus). Annales Zoologici Fennici, 25, 191–201.Google Scholar
Otronen, M (1988b) Intra- and intersexual interactions at breeding burrows in the horned beetle, Coprophanaeus ensifer. Animal Behaviour, 36, 741–748.CrossRefGoogle Scholar
Palmer, TJ (1978) A horned beetle which fights. Nature, 274, 583–584.CrossRefGoogle Scholar
Panhuis, TM, Butlin, R, Zuk, M, et al. (2001) Sexual selection and speciation. Trends in Ecology and Evolution, 16, 364–371.CrossRefGoogle ScholarPubMed
Parker, GA (1974) Assessment strategy and evolution of fighting behavior. Journal of Theoretical Biology, 47, 223–243.CrossRefGoogle Scholar
Parzer, HF & Moczek, AP (2008) Rapid antagonistic coevolution between primary and secondary sexual characters in horned beetles. Evolution, 62, 2423–2428.CrossRefGoogle ScholarPubMed
Pizzo, A, Roggero, A, Palestrini, C, et al. (2008) Rapid shape divergences between natural and introduced populations of a horned beetle partly mirror divergences between species. Evolution and Development, 10, 166–175.CrossRefGoogle ScholarPubMed
Pomfret, JC & Knell, RJ (2006) Sexual selection and horn allometry in the dung beetleEuoniticellus intermedius. Animal Behaviour, 71, 567–576.CrossRefGoogle Scholar
Potten, C & Wilson, J (2004) Apoptosis: The Life and Death of Cells. Cambridge: Cambridge University Press.Google Scholar
Rasa, OAE (1999) Division of labour and extended parenting in a desert tenebrionid beetle. Ethology, 105, 37–56.CrossRefGoogle Scholar
Rasmussen, JL (1994) The influence of horn and body size on the reproductive behavior of the horned rainbow Scarab beetle Phanaeus difformis (Coleoptera: Scarabaeidae). Journal of Insect Behavior, 7, 67–82.CrossRefGoogle Scholar
Rowland, JM & Emlen, DJ (2009) Two thresholds, three male forms result in facultative male trimorphism in beetles. Science, 323, 773–776.CrossRefGoogle ScholarPubMed
Rubenstein, DI (1984) Resource acquisition and alternative mating strategies in water striders. American Zoologist, 24, 345–353.CrossRefGoogle Scholar
Rudinsky, JA & Ryker, LC (1976) Sound production in Scolytidae: rivalry and pre-mating stridulation of male Douglas-fir beetles. Journal of Insect Physiology, 22, 997–1003.CrossRefGoogle Scholar
Schuett, GW (1997) Body size and agonistic experience affect dominance and mating success in male copperheads. Animal Behaviour, 54, 213–224.CrossRefGoogle ScholarPubMed
Schuster, JC (1983) Acoustical signals of Passalid beetles: Complex repertoires. Florida Entomologist, 66, 486–496.CrossRefGoogle Scholar
Schuster, JC & Schuster, LB (1985) Social behavior in Passalid beetles (Coleoptera: Passalidae): Cooperative brood care. Florida Entomologist, 68, 266–272.CrossRefGoogle Scholar
Setsuda, K, Tsuchida, K, Watanabe, H, et al. (1999) Size dependent predatory pressure in the Japanese horned beetle, Allomyrina dichotoma L. (Coleoptera; Scarabaeidae). Journal of Ethology, 17, 73–77.CrossRefGoogle Scholar
Shelby, JA, Madewell, R & Moczek, AP (2007) Juvenile hormone mediates sexual dimorphism in horned beetles. Journal of Experimental Zoology, 308B, 417–427.CrossRefGoogle Scholar
Simmons, LW & Emlen, DJ (2006) Evolutionary trade-off between weapons and testes. Proceedings of the Royal Society of London B, 103, 16346–16351.Google ScholarPubMed
Simmons, LW & Garcia-Gonzalez, F (2008) Evolutionary reduction in testes size and competitive fertilization success in response to the experimental removal of sexual selection in dung beetles. Evolution, 62, 2580–2591.CrossRefGoogle ScholarPubMed
Simmons, LW, Tomkins, JL & Hunt, J (1999) Sperm competition games played by dimorphic male beetles. Proceedings of the Royal Society of London B, 266, 145–150.CrossRefGoogle Scholar
Siva-Jothy, MT (1987) Mate securing tactics and the cost of fighting in the Japanese Horned Beetle, Allomyrina dichotoma L. (Scarabaeidae). Journal of Ethology, 5, 165–172.CrossRefGoogle Scholar
Snell-Rood, EC, Cash, A, Han, MV, et al. (2011) Developmental decoupling of alternative phenotypes: Insights from the transcriptomes of horn-polyphenic beetles. Evolution, 65, 231–245.CrossRefGoogle ScholarPubMed
Svacha, P (1992) What are and what are not imaginal discs: Reevaluation of some basic concepts (Insecta, Holometabola). Developmental Biology, 154, 101–117.CrossRefGoogle Scholar
Tomkins, JL, Kotiaho, JS & LeBas, NR (2005) Matters of scale: Positive allometry and the evolution of male dimorphisms. American Naturalist, 165, 389–402.CrossRefGoogle ScholarPubMed
Thornhill, R & Alcock, J (1983) The Evolution of Insect Mating Systems. New York, NY: iUniverse.com, Inc.CrossRefGoogle Scholar
Trumbo, ST (2007) Defending young biparentally: Female risk-taking with and without a male in the buyring beetle, Nicrophorus pustulatus. Behavioral Ecology and Sociobiology, 61, 1717–1723.CrossRefGoogle Scholar
Wasik, BR, Rose, DJ & Moczek, AP (2010) Beetle horns are regulated by the Hox gene, sex combs reduced, in a species- and sex-specific manner. Evolution and Development, 12, 353–362.CrossRefGoogle Scholar
Watson, NL & Simmons, LW (2010a) Mate choice in the dung beetle Onthophagus sagittarius: Are female horns ornaments? Behavioral Ecology, 21, 424–430.CrossRefGoogle Scholar
Watson, NL & Simmons, LW (2010b) Reproductive competition promotes the evolution of female weaponry. Proceedings of the Royal Society of London B, 277, 2035–2040.CrossRefGoogle ScholarPubMed
Wells, MS (1988) Effects of body size and resource value on fighting behaviour in a jumping spider. Animal Behaviour, 36, 321–326.CrossRefGoogle Scholar
West-Eberhard, MJ (1983) Sexual selection, social competition, and speciation. Quarterly Review of Biology, 58, 155–183.CrossRefGoogle Scholar
West-Eberhard, MJ (2003) Developmental Plasticity and Evolution. Oxford: Oxford University Press.Google Scholar
Whiting, MF, Bradler, S & Maxwell, T (2003) Loss and recovery of wings in stick insects. Nature, 421, 264–267.CrossRefGoogle ScholarPubMed
Windsor, DM (1987) Natural history of a subsocial tortoise beetle Acromis sparsa Boheman (Chrysomelidae: Cassidinae) in Panama. Psyche, 94, 127–150.CrossRefGoogle Scholar
Yamane, T, Okada, K, Nakayama, S, et al. (2010) Dispersal and ejaculatory strategies associated with exaggeration of weapon in an armed beetle. Proceedings of the Royal Society of London B, 277, 1705–1710.CrossRefGoogle Scholar

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