In many animals, males interact with or without physical contact to contest the possession of mating sites. The winner in such interactions often has larger body size and is the individual that previously occupied the territory (the resident). However, there is little consensus among studies on territorial insect species concerning which traits determine individual fighting ability, and why residency increases the chances of victory. We studied the butterfly Actinote pellenea to evaluate the hypothesis that stronger males are more likely to become residents, and that traits functionally related to behaviors adopted during contests should be important determinants of fighting ability. Males of A. pellenea grapple in the air and fall to the ground during combat. Therefore, we predicted that traits related to physical strength should be the most important determinants of the outcome of contests. To test our hypothesis, we observed fights between resident and intruder males, and fights in which both rivals were induced to behave as residents. We found that physical fighting was more likely when both rivals behave as residents. In addition, wing length and body mass were more closely related to contest success than age. Our results indicate that previous residency increases male motivation to fight, and that stronger individuals become residents. Although butterflies are considered weaponless, we suggest that traits important to contest resolution change according to behaviors adopted during disputes.
Sexual selection Mating territories Competition Rendezvous sites RHP Lek
We thank the Fundação José Pedro de Oliveira and Parque Estadual da Serra do Mar that permitted the field studies and the Fundação de Amparo à Pesquisa do Estado de São Paulo (Fapesp) for fellowships. We also thank Darrel Kemp, Martin Bergman, and three anonymous referees for their many important contributions.
This study was funded by the Fundação de Amparo à Pesquisa do Estado de São Paulo—FAPESP (grant number 08/52354-2R).
Compliance with ethical standards
Conflict of interest
The authors declare that they have no competing interests.
This study was authorized by the Brazilian governmental agency called ICMBio (license 18904-1), in accordance with Brazilian environmental laws.
This article does not contain any studies with human participants performed by any of the authors.
Kokko H (2013) Dyadic contests: modelling fights between two individuals. In: Hardy ICW, Briffa M (eds) Animal contests. Cambrigde University Press, Cambrigde, pp 5–32CrossRefGoogle Scholar
Koskimäki J, Rantala MJ, Taskinen J, Tynkkynen K, Suhonen J (2004) Immunocompetence and resource holding potential in the damselfly, Calopteryx virgo L. Behav Ecol 15:169–173CrossRefGoogle Scholar
Lailvaux SP, Irschick DJ (2006) A functional perspective on sexual selection: insights and future prospects. Anim Behav 72:263–273CrossRefGoogle Scholar
Leimar O, Enquist M (1984) Effects of asymmetries in owner-intruder conflicts. J Theor Biol 111:475–491CrossRefGoogle Scholar
Maynard-Smith J (1982) Evolution and the theory of games. Cambridge University Press, Cambridge, UKCrossRefGoogle Scholar
Minor WC (1950) Footprints in the trail. Erle Press, ChicagoGoogle Scholar
Oliveira R, Schlindwein C (2010) Experimental demonstration of alternative mating tactics of male Ptilothrix fructifera (Hymenoptera, Apidae). Anim Behav 80:241–247CrossRefGoogle Scholar
Orr AG (1999) The big greasy, Cressida cressida (Papilionidae). In: Kirchng RL, Scheermeyer E, Jones RE, Pierce NE (eds) Biology of Australian butterflies. CSIRO, CanberraGoogle Scholar
Peixoto PEC, Benson WW (2008) Body mass and not wing length predicts territorial success in a tropical satyrine butterfly. Ethol 114:1069–1077CrossRefGoogle Scholar
Peixoto PEC, Benson WW (2009) Daily activity patterns of two co-ocurring tropical satyrine butterflies. J Insect Sci 9:54Google Scholar
Peixoto PEC, Benson WW (2011) Seasonal effects of density on territory occupation by males of the satyrine butterfly Paryphthimoides phronius (Butler 1867). J Ethol 27:489–496CrossRefGoogle Scholar
Peixoto PEC, Benson WW (2012) Influence of previous residency and body mass in the territorial contests of the butterfly Hermeuptychia fallax (Lepidoptera: Satyrinae). J Ethol 30:61–68CrossRefGoogle Scholar
Peixoto PEC, Medina A, Mendoza-Cuenca L (2014) Do territorial butterflies show a macroecological fighting pattern in response to environmental stability? Behav Process 109:14–20CrossRefGoogle Scholar
Pinheiro CEG (1991) Territorial hilltopping behavior of three swallowtail butterflies (Lepidoptera, Papilionidae) in western Brazil. J Res Lep 29:134–142Google Scholar
Rutowski RL (1991) The evolution of male mate-locating behavior in butterflies. Am Nat 138:1121–1139CrossRefGoogle Scholar
Rutowski RL (1992) Male mate-locating behavior in the common eggfly, Hypolimnas bolina (Nymphalidae). J Lep Soc 46:24–38Google Scholar
Stuart-Fox D (2006) Testing game theory models: fighting ability and decision rules in chameleon contests. Proc R Soc Lond B 273:1555–1561CrossRefGoogle Scholar
Switzer PV (2002) Territory quality, habitat selection, and competition in the amberwing dragonfly, Perithemis tenera (Say) (Odonata: Libellulidae): population patterns as a consequence on individual behavior. J Kansas Entomol Soc 75:154–157Google Scholar
Takeuchi T (2006) Matter of size or matter of residency experience? Territorial contest in a green hairstreak, Chrysozephyrus smaragdinus (Lepidoptera: Lycaenidae). Ethology 112:293–299CrossRefGoogle Scholar
Takeuchi T, Honda K (2009) Early comers become owners: effect of residency experience on territorial contests dynamics in a lycaenid butterfly. Ethology 115:767–773CrossRefGoogle Scholar
Temeles EJ (1994) The role of neighbours in territorial systems: when are they ‘dear enemies’? Anim Behav 47:339–350CrossRefGoogle Scholar
Tyler H, Brown KSJ, Wilson K (1994) Swallowtail butterflies of the Americas. Scientific Publishers, GainesvilleGoogle Scholar
Vieira MC, Peixoto PEC (2013) Winners and losers: a meta-analysis of functional determinants of fighting ability in arthropod contests. Funct Ecol 27:305–313CrossRefGoogle Scholar
Waage JK (1988) Confusion over residence and the escalation of damselfly territorial disputes. Anim Behav 36:586–595CrossRefGoogle Scholar
Wickman PO, Wiklund C (1983) Territorial defence and its seasonal decline in the speckled wood butterfly (Pararge aegeria). Anim Behav 31:1206–1216CrossRefGoogle Scholar