The bigger individual in a fight usually wins unless the smaller individual is a resident or has recently won a fight. I conducted three experiments on the effects of body size, residency, and fight history on the outcome of male-male fights in a burying beetle. Fights were staged between an intruding male and the male of a male-female pair. When males differed in size, the larger male usually won regardless of residency or individual fight histories. Residents and winners of previous fights won only when competing males were similar in size. Hence, male body size largely determines the outcomes of fights in this beetle. 1. Introduction Fighting between males over mating opportunities is a widespread phenomenon in the animal kingdom and has received much empirical attention [1]. Fighting ability is often correlated with morphological and physiological attributes such as body size, weaponry, and ornaments [1], but these are not the only attributes that may determine fighting ability. It has been hypothesized that the “prior residence effect” can also affect the outcome of fighting contests in accordance with the convention “resident wins, intruder retreats” [2]. A third effect is the “winner-loser effect,” in which winners are more likely to win again and losers are more likely to lose again [3]. These two effects sometimes counteract morphological and physiological attributes (e.g., [4]). The complex parental behaviour of burying beetles (Nicrophorus: Silphidae) has been well-studied (reviewed in [5, 6]). Nicrophorus exploits small vertebrate carrions as food for its young. A male-female pair prepares a carcass by burying it, removing its hair, and rounding it into a ball. Eggs are then laid in the soil adjacent to the carrion ball. After hatching, the larvae crawl to the carrion ball, where they are fed by parental regurgitations. Nicrophorus is generally monogamous [7–9], and both sexes display intense intrasexual competition [10, 11]. Two or more individuals of both sexes often locate the same carcass, but usually only a single dominant pair eventually occupies the carcass. Resident males are more likely to be injured than resident females [12], and males have a greater tendency to guard [13, 14]. Contests between males are expected to be more intense than those between females. Larger individuals of Nicrophorus usually win contests among conspecifics in N. humator [10] or in N. quadripunctatus [11]. However, the presence of the winner effect is supported by a previous study of N. humator [15], and it is possible that other attributes affect the
References
[1]
F. A. Huntingford and A. K. Turner, Animal Conflict, Chapman & Hall, New York, NY, USA, 1987.
[2]
J. Maynard Smith and G. A. Parker, “The logic of asymmetric contests,” Animal Behaviour, vol. 24, no. 1, pp. 159–175, 1976.
[3]
Y. Hsu and L. L. Wolf, “The winner and loser effect: integrating multiple experiences,” Animal Behaviour, vol. 57, no. 4, pp. 903–910, 1999.
[4]
K. Okada and T. Miyatake, “Effect of losing on male fights of broad-horned flour beetle, Gnatocerus cornutus,” Behavioral Ecology and Sociobiology, vol. 64, no. 3, pp. 361–369, 2010.
[5]
A. K. Eggert and J. K. Müller, “Biparental care and social evolution in burying beetles: lessons from the larder,” in The Evolution of Social Behavior in Insects and Arachnids, J. C. Choe and B. J. Crespi, Eds., pp. 216–236, Cambridge University Press, Cambridge, UK, 1997.
[6]
M. P. Scott, “The ecology and behavior of burying beetles,” Annual Review of Entomology, vol. 43, pp. 595–618, 1998.
[7]
A.-K. Eggert and S. K. Sakaluk, “Benefits of communal breeding in burying beetles: a field experiment,” Ecological Entomology, vol. 25, no. 3, pp. 262–266, 2000.
[8]
S. T. Trumbo, “Monogamy to communal breeding: exploitation of a broad resource base by burying beetles (Nicrophorus),” Ecological Entomology, vol. 17, no. 3, pp. 289–298, 1992.
[9]
S. T. Trumbo and A. K. Eggert, “Beyond monogamy: territory quality influences sexual advertisement in male burying beetles,” Animal Behaviour, vol. 48, no. 5, pp. 1043–1047, 1994.
[10]
M. Otronen, “The effect of body size on the outcome of fights in burying beetles (Nicrophorus),” Annales Zoologici Fennici, vol. 25, no. 2, pp. 191–201, 1988.
[11]
S. Suzuki, M. Nagano, and S. T. Trumbo, “Intrasexual competition and mating behavior in Ptomascopus morio (Coleoptera: Silphidae Nicrophorinae),” Journal of Insect Behavior, vol. 18, no. 2, pp. 233–242, 2005.
[12]
S. T. Trumbo, “Defending young biparentally: female risk-taking with and without a male in the burying beetle, Nicrophorus pustulatus,” Behavioral Ecology and Sociobiology, vol. 61, no. 11, pp. 1717–1723, 2007.
[13]
I. A. Fetherston, M. P. Scott, and J. F. A. Traniello, “Parental care in burying beetles: the organization of male and female brood-care behavior,” Ethology, vol. 85, pp. 177–190, 1990.
[14]
I. A. Fetherston, M. P. Scott, and J. F. A. Traniello, “Behavioural compensation for mate loss in the burying beetle Nicrophorus orbicollis,” Animal Behaviour, vol. 47, no. 4, pp. 777–785, 1994.
[15]
M. Otronen, “The effect of prior experience on the outcome of fights in the burying beetle, Nicrophorus humator,” Animal Behaviour, vol. 40, no. 5, pp. 980–982, 1990.
[16]
S. Suzuki, “Biparental burying beetles, Nicrophorus quadripunctatus (Coleoptera, Silphidae), do not show a division of labor in their carrion-burial behavior,” Elytra New Series, vol. 1, pp. 41–45, 2011.
[17]
M. P. Scott, “Brood guarding and the evolution of male parental care in burying beetles,” Behavioral Ecology and Sociobiology, vol. 26, no. 1, pp. 31–39, 1990.
[18]
S. T. Trumbo, “Reproductive benefits of infanticide in a biparental burying beetle Nicrophorus orbicollis,” Behavioral Ecology and Sociobiology, vol. 27, no. 4, pp. 269–273, 1990.
[19]
S. A. Safryn and M. P. Scott, “Sizing up the competition: do burying beetles weigh or measure their opponents?” Journal of Insect Behavior, vol. 13, no. 2, pp. 291–297, 2000.
[20]
S. Suzuki, “Cooperative defence of brood in Nicrophorus quadripunctatus (Coleoptera: Silphidae),” European Journal of Entomology, vol. 108, no. 4, pp. 561–563, 2011.
[21]
S. T. Trumbo, “Infanticide, sexual selection and task specialization in a biparental burying beetle,” Animal Behaviour, vol. 72, no. 5, pp. 1159–1167, 2006.
[22]
Y. Hsu, R. L. Earley, and L. L. Wolf, “Modulation of aggressive behaviour by fighting experience: mechanisms and contest outcomes,” Biological Reviews of the Cambridge Philosophical Society, vol. 81, no. 1, pp. 33–74, 2006.
[23]
C. Rutte, M. Taborsky, and M. W. G. Brinkhof, “What sets the odds of winning and losing?” Trends in Ecology and Evolution, vol. 21, no. 1, pp. 16–21, 2006.