![]() Such data may include more accurate quantification of RHP-related traits or whole-body measures of fitness ( Briffa, Hardy, and Mowles 2013). 2011 Briffa, Hardy, and Mowles 2013 Palaoro and Briffa 2017). Several authors have suggested that collecting data on a broad suite of traits used in contests may provide a potential solution to this challenge ( Briffa and Elwood 2009 Kasumovic et al. Identifying the assessment strategy that is operational in a particular study system has, however, proven to be challenging across different animal groups (e.g., Turner and Huntingford 1986 Palaoro and Briffa 2017 Green and Patek 2018 Walker and Holwell 2018). 2009 Elwood and Prenter 2013 Walker and Holwell 2018). 2003), ungulates (see review in Jennings and Gammell 2013), and many studies in insects and spiders (e.g., Parker and Thompson 1980 Kelly 2006 Small et al. Models of animal contest assessment have been applied to empirical studies for a range of taxa to assess the applicability of these models, for example, newts ( Verrell 1986), fiddler crabs ( Pratt et al. Three classes of competitive assessment models have been developed, each predicting different correlations between the RHP and contest cost (generally contest duration) of a contestant: pure self-assessment, cumulative self-assessment, and mutual assessment strategies ( Arnott and Elwood 2009 for predictions and assumptions of the assessment models, see Table 1 of Painting and Holwell 2014 and Table 1 of Green and Patek 2018). This development has been highly influential on the growth and direction of research on animal contest systems, giving rise to more refined evolutionary game theory models ( Parker and Rubenstein 1981 Austad 1989 Kokko et al. Maynard Smith and Price (1973) published the first applications of game theory to the study of decision-making and assessment strategies in animal contests. RHP may be influenced by several factors, including weapon and body size, and quantifying the relative importance of each is integral to increasing our understanding of decision-making by combatants ( Vieira and Peixoto 2013 Palaoro and Briffa 2017). The physical contests in which these weapons are wielded may incur significant costs to contestants ( Enquist and Leimar 1987 Briffa and Sneddon 2007 Lane and Briffa 2017), and they are generally won by the individual with the greatest fighting ability, often referred to as the greatest “resource holding power” (RHP sensu Parker 1974). Intrasexual contest competition among males for mates has driven the evolution of highly exaggerated “weaponized” traits in a diverse range of animal taxa ( Kodric-Brown and Brown 1984 Blanckenhorn 2005 Plard et al. Such inconclusive findings are not uncommon in animal contest assessment studies, even when contest cost and resource holding power data are contextualized with behavioral and ecological data. We found mixed support for both pure self-assessment and mutual assessment contest strategies. Contest outcome was determined by hind femur size and strongly influenced by residency effects. We found that males of all sizes engaged aggressively in intrasexual contests for mating access to sedentary females, utilizing their hypertrophied hind legs as weapons. The present study integrates data on contest behavior, weapon morphology, residency effects, cost accumulation, and correlates of contest success to test game theory-informed models of competitive assessment strategies in the sexually dimorphic monkey beetle, Heterochelus chiragricus. In view of this, some authors have suggested that utilizing a broad suite of data concerning multiple facets of the study system may assist in gaining clearer insights into animal contests and assessment strategies. Conclusions from empirical studies about the means of assessment in their study systems have not, however, always been clear. Game theory has been used as a basis to develop models of the competitive assessment strategies that may be used by males either to judge their probability of winning by comparing their own fighting ability to that of their opponents or to persist in contests for a period determined only by their own fighting ability. Some of the most striking examples of intrasexual contest competition are to be found in the insects, whose weaponry and contest behaviors have become highly intricate and diverse.
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