Male-male Contests Research Articles

Intrasexual Selection for Upper Limb Length in Homo sapiens.

Sexual selection via contest competition has equipped countless organisms with weaponry in their appendages to overpower their opponents. Here, we tested (1) whether greater upper limb length-measured as span controlling for biacromial width-confers an advantage in contest competition among adult humans, (2) several possible means by which upper limb length might increase success in intrasexual contest competition, and (3) whether, consistent with male-male contest competition creating stronger selection pressures than female-female contest competition, male Homo sapiens have greater upper limb length. We collected fight statistics and facial and body photographs from professional combatants (N = 715) in the Ultimate Fighting Championship (UFC; Study 1). Sexual dimorphism in upper limb length was then examined via diverse and demographically representative samples from four studies (total N = 6915), from Croatian adolescents and older Singaporean adults to United States Army personnel born across all major world regions (Studies 2a-2d). First, we found that greater upper limb length is associated with increased success in intrasexual contest competition, an effect driven by both the capacity to grapple opponents to submission and to knock opponents unconscious (Study 1). Second, we found unequivocal, cross-cultural evidence of unique sexual dimorphism in upper limb length after controlling for allometry: across four studies, men exhibited longer upper limbs than women (Studies 2a-2d). Upper limb length may have been shaped by intrasexual selection, with implications across the biological, anthropological, and psychological sciences.

First report of arm-span competition in buthid scorpions: male-male contest in Tityus cf. rosenbergi Pocock, 1898

First report of arm-span competition in buthid scorpions: male-male contest in Tityus cf. rosenbergi Pocock, 1898

Does losing reduce the tendency to engage with rivals to reach mates? An experimental test.

Male-male contests for access to females or breeding resources are critical in determining male reproductive success. Larger males and those with more effective weaponry are more likely to win fights. However, even after controlling for such predictors of fighting ability, studies have reported a winner-loser effect: previous winners are more likely to win subsequent contests, while losers often suffer repeated defeats. While the effect of winning-losing is well-documented for the outcome of future fights, its effect on other behaviors (e.g. mating) remains poorly investigated. Here, we test whether a winning versus losing experience influenced subsequent behaviors of male mosquitofish (Gambusia holbrooki) toward rivals and potential mates. We housed focal males with either a smaller or larger opponent for 24 h to manipulate their fighting experience to become winners or losers, respectively. The focal males then underwent tests that required them to enter and swim through a narrow corridor to reach females, bypassing a cylinder that contained either a larger rival male (competitive scenario), a juvenile or was empty (non-competitive scenarios). The tests were repeated after 1 wk. Winners were more likely to leave the start area and to reach the females, but only when a larger rival was presented, indicating higher levels of risk-taking behavior in aggressive interactions. This winner-loser effect persisted for at least 1 wk. We suggest that male mosquitofish adjust their assessment of their own and/or their rival's fighting ability following contests in ways whose detection by researchers depends on the social context.

Just a matter of size? Evaluating allometry and intersexual heterometry in Pagurus bernhardus using ratios and indices (Decapoda, Anomura).

Heterochely denotes the presence of dissimilarly sized chelipeds on opposite sides of the body, a prevalent occurrence in diverse crustaceans. Conversely, heterometry pertains to the quantifiable disparities in size between these chelipeds. Both chelipeds hold pivotal roles in activities such as foraging, mating, and defense. Consequently, individuals of both genders in heterochelic species exhibit this morphological pattern. Previous studies have identified sexual dimorphism in cheliped size, with males displaying larger major chelipeds compared to females, albeit solely relying on propodus length as a size proxy and focusing solely on the major cheliped. In our study, we meticulously examined 190 specimens of the common European hermit crab Pagurus bernhardus from two collections. We sought to elucidate allometric relationships and assess whether heterometry exhibited sex-based differences when adjusting for body size by using ratios. Our findings revealed that male chelipeds displayed hyperallometric growth relative to females, and all three calculated heterometry indices exhibited significant disparities between the sexes. Consequently, male specimens exhibited larger major and minor chelipeds, even when theoretically matched for body size with females. This phenomenon may be attributed, among other factors, to male-male contests. Should indirect mate selection favor males with larger chelipeds in proportion to their body size, this dynamic could potentiate sexual selection in their favor.

Inbreeding depression in a sexually selected weapon and the homologue in females.

Theory predicts that traits with heightened condition dependence, such as sexually selected traits, should be affected by inbreeding to a greater degree than other traits. The presence of environmental stress may compound the negative consequences of inbreeding depression. In this study, we examined inbreeding depression across multiple traits and whether it increased with a known form of environmental stress. We conducted our experiment using both sexes of the sexually dimorphic leaf-footed cactus bug, Narnia femorata (Hemiptera: Coreidae). Adult male cactus bugs have enlarged hind legs used as weapons in male-male contests; these traits, and their homologue in females, have been previously found to exhibit high condition dependence. In this study, we employed a small developmental group size as an environmental stress challenge. Nymph N. femorata aggregate throughout their juvenile stages, and previous work has shown the negative effects of small group size on survivorship and body size. We found evidence of inbreeding depression for survival and seven of the eight morphological traits measured in both sexes. Inbreeding depression was higher for the size of the male weapon and the female homolog. Additionally, small developmental group size negatively affected survival to adulthood. However, small group size did not magnify the effects of inbreeding on morphological traits. These findings support the hypothesis that traits with heightened condition dependence exhibit higher levels of inbreeding depression.

Alternative reproductive tactics of unflanged and flanged male orangutans revisited.

In many slowly developing mammal species, males reach sexual maturity well before they develop secondary sexual characteristics. Sexually mature male orangutans have exceptionally long periods of developmental arrest. The two male morphs have been associated with behavioral alternative reproductive tactics, but this interpretation is based on cross-sectional analyses predominantly of Northwest Sumatran populations. Here we present the first longitudinal analyses of behavioral changes of 10 adult males that have been observed in both unflanged and flanged morph. We also analyzed long-term behavioral data on an additional 143 individually identified males from two study sites, Suaq (Sumatra, Pongo abelii) and Tuanan (Borneo, Pongo pygmaeus wurmbii), to assess male mating tactics cross-sectionally in relation to population, male morph (unflanged and flanged), and other socio-ecological factors. Both our longitudinal and cross-sectional results confirm and refine previous cross-sectional accounts of the differences in mating tactics between the unflanged and the flanged male morphs. In the unflanged morph, males exhibit higher sociability, particularly with females, and higher rates of both copulation and sexual coercion than in the flanged morph. Based on our results and those of previous studies showing that females prefer flanged males, and that flanged males have higher reproductive success, we conclude that unflanged males face a trade-off between avoiding male-male contest competition and gaining mating access to females, and thus follow a "best-of-a-bad-job" mating strategy.

Juvenile leg autotomy predicts adult male morph in a New Zealand harvestman with weapon polymorphism.

Intraspecific weapon polymorphisms that arise via conditional thresholds may be affected by juvenile experience such as predator encounters, yet this idea has rarely been tested. The New Zealand harvestman Forsteropsalis pureora has three male morphs: majors (alphas and betas) are large-bodied with large chelicerae used in male-male contests, while minors (gammas) are small-bodied with small chelicerae and scramble to find mates. Individuals use leg autotomy to escape predators and there is no regeneration of the missing leg. Here, we tested whether juvenile experience affects adult morph using leg autotomy scars as a proxy of predator encounters. Juvenile males that lost at least one leg (with either locomotory or sensory function) had a 45 times higher probability of becoming a minor morph at adulthood than intact juvenile males. Leg loss during development may affect foraging, locomotion, and/or physiology, potentially linking a juvenile's predator encounters to their final adult morph and future reproductive tactic.

Female-female aggression in the Gila monster (Heloderma suspectum).

Historically, the role of aggression in the social lives of animals overwhelmingly focused on males. In recent years, however, female-female aggression in vertebrates, particularly lizards, has received increasing attention. This growing body of literature shows both similarities and differences to aggressive behaviours between males. Here, we document female-female aggression in captive Gila monsters (Heloderma suspectum). Based on four unique dyadic trials (eight adult female subjects), we developed a qualitative ethogram. Unexpected and most intriguing were the prevalence and intensity of aggressive acts that included brief and sustained biting, envenomation, and lateral rotation (i.e. rolling of body while holding onto opponent with closed jaws). Given specific behavioural acts (i.e. biting) and the results of bite-force experiments, we postulate that osteoderms (bony deposits in the skin) offer some degree of protection and reduce the likelihood of serious injury during female-female fights. Male-male contests in H. suspectum, in contrast, are more ritualized, and biting is rarely reported. Female-female aggression in other lizards has a role in territoriality, courtship tactics, and nest and offspring guarding. Future behavioural research on aggression in female Gila monsters is warranted to test these and other hypotheses in the laboratory and field.

Trade-off between pre and post-copulatory traits depends on locomotor activity in male Tribolium castaneum beetles.

Locomotor performance is an indicator of dynamic exercise; thus, it is a central trait in many animal behaviours. Although higher locomotor endurance may increase male reproductive success (e.g., in mate searching and male-male contests), investment in other male reproductive traits (e.g., male attractiveness and sperm competition) may be decreased through energy consumption due to higher activity levels. Here, I investigated male attractiveness, mating success, and paternity success using males of the red flour beetle Tribolium castaneum selected for higher (H) and lower (L) locomotor endurance. Although there was no difference in male attractiveness between the selection regimes, H males had significantly higher mating success than L males. Conversely, L males had significantly higher paternity success than H males. Therefore, there was a trade-off between mating success and paternity success among the selection regimes, suggesting that locomotor endurance affects male reproduction in T.castaneum, and individual variation of locomotor endurance may be maintained within a population.

The Effect of Brief or Prolonged Bouts of Winning or Losing Male-Male Contests on Plasticity in Sexually Selected Traits.

AbstractFight outcomes often affect male fitness by determining their access to mates. Thus, "winner-loser" effects, where winners often win their next contest while losers tend to lose, can influence how males allocate resources toward pre- and postcopulatory traits. We experimentally manipulated the winning/losing experiences of pairs of size-matched male Gambusia holbrooki for 1 day, 1 week, or 3 weeks to test whether prior winning/losing experiences differentially affect the plasticity of male investment into either mating effort (precopulatory) or ejaculates (postcopulatory). When winner/loser pairs directly competed for a female, winners had better precopulatory outcomes than losers for three of the four traits we measured: mating attempts, successful attempts, and time spent with the female (but not aggression). However, winners and losers did not differ in either total sperm counts or sperm velocity. Interestingly, absolute male size, an important predictor of fighting success, mediated winner-loser effects on how long males then spent near a female. Compared with losers, smaller winners spent more time with the female than did larger winners, suggesting that how males respond to prior social experiences is size dependent. We discuss the general importance of controlling for inherent male condition when comparing male investment into condition-dependent traits.

Males armed with big weapons win fights at limited cost in ant-mimicking jumping spiders.

A core assumption of sexual selection theory is that sexually selected weapons, specialized morphological structures used directly in male contests, can improve an individual's reproductive success but only if the bearer can overcome associated costs, the negative effects on the bearer's fitness components. However, recent studies have shown that producing and wielding exaggerated weapons may not necessarily be costly. Rather, some traits can be selected for supporting, or compensating for, the expense of producing and wielding such exaggerated weapons. In the ant-mimicking jumping spider Myrmarachne gisti, exaggerated chelicerae are borne only by adult males and not females, showing sexual dimorphism and steep positive allometry with body size. Here, we determine the potential benefits of bearing exaggerated chelicerae during male contests and explore the potential for costs in terms of prey-capture efficiency and compensation between chelicera size and neighboring trait size. While males with longer chelicerae won most of their male-male contests, we found no significant differences in prey-capture efficiency between males and females regardless of whether prey was winged or flightless. Males' elongated chelicerae thus do not impede their efficiency at capturing prey. Furthermore, we found that the sizes of all neighboring traits are positively correlated with chelicera size, suggesting that these traits may be under correlational selection. Taken together, our findings suggest that M. gisti males armed with the exaggerated chelicerae that function as weapons win more fights at limited cost for performance in prey capture and compensate for neighboring structures.

Human third-party observers accurately track fighting skill and vigour along their unique paths to victory

Sexual selection via male-male contest competition has shaped the evolution of agonistic displays, weaponry, and fighting styles, and is further argued to have shaped human psychological mechanisms to detect, process, and respond appropriately to cues of fighting ability. Drawing on the largest fight-specific dataset to date across the sports and biological sciences (N = 2765 fights), we examined how different indicators of fighting ability in humans reflect unique paths to victory and indicate different forms of perceived and actual resource-holding power (RHP). Overall, we discovered that: (1) both striking skill and vigour, and grappling skill and vigour, individually and collectively predict RHP; (2) different RHP indicators are distinguished by a unique path to victory (e.g., striking skill is a knockout-typical strategy, whereas grappling vigour is a submission-typical strategy); and (3) third-party observers accurately track fighting skill and vigour along their unique paths to victory. Our argument that different measures of RHP are associated with unique paths to victory, and third-party observers accurately track fighting vigour and skill along their unique paths to victory, advance our understanding not only of human contest competition, but animal contest theory more broadly.

Condition- and context-dependent alternative reproductive tactic in Drosophila prolongata

Abstract Species with intense male–male competition for access to females often show alternative reproductive tactics (ARTs) where males of lower competitive ability adopt a sneaky behavior to gain access to mates. These ARTs are usually associated with intrasexual dimorphisms, in that males with distinct morphologies show different tactics. In some cases, however, males adopt different tactics without being dimorphic. Male Drosophila prolongata exhibit continuous variation in body size and shape, with enlarged forelegs that they use in male-male contests and in courtship, including stimulation of the female’s abdomen. During this “leg vibration,” however, nearby males can intercept the courted female and mate without their own courtship. Here, we studied the causes and consequences of these different mating tactics in competitive mating trials between males varying in their size and shape. We found that male mating tactics were condition-dependent. Whereas large, high-condition males were more likely to show leg vibration in their courtship, smaller, lower-condition males were more likely to intercept. However, the number of offspring produced was independent of male condition and reproductive tactic. We discuss possible scenarios for the evolution and maintenance of the ARTs and some future directions for the study of ARTs in this species and in general.

Testosterone, signal coloration, and signal color perception in male zebra finch contests.

Many animals use assessment signals to resolve contests over limited resources while minimizing the costs of those contests. The carotenoid-based orange to red bills of male zebra finches (Taeniopygia guttata) are thought to function as assessment signals in male-male contests, but behavioral analyses relating contest behaviors and outcomes to bill coloration have yielded mixed results. We examined the relationship between bill color and contests while incorporating measurements of color perception and testosterone (T) production, for an integrative view of aggressive signal behavior, production, and perception. We assayed the T production capabilities of 12 males in response to a gonadotropin-releasing hormone (GnRH) challenge. We then quantified the initiation, escalation, and outcome of over 400 contests in the group, and measured bill color using calibrated photography. Finally, because signal perception can influence signal function, we tested how males perceive variation in bill coloration, asking if males exhibit categorical perception of bill color, as has been shown recently in female zebra finches. The data suggest that males with greater T production capabilities than their rivals were more likely to initiate contests against those rivals, while males with redder bills than their rivals were more likely to win contests. Males exhibited categorical color perception, but individual variation in the effect of categorical perception on color discrimination abilities did not predict any aspects of contest behavior or outcomes. Our results are consistent with the hypotheses that T plays a role in zebra finch contests and that bill coloration functions as an aggressive signal. We suggest future approaches, based on animal contest theory, for how links among signals, perception, and assessment can be tested.

Behavioral repertoire of biparental care inApistogramma trifasciata(Pisces: Cichlidae)

Cichlid fish exhibit a complex behavioral repertoire, mainly regarding social and reproductive behaviors. Nevertheless, although dwarf cichlids exhibit a remarkable diversity of behaviors, only a few studies have been conducted with these fish. Apistogramma trifasciata is a common dwarf cichlid in the Neotropics and widely used as ornamental fish. Both males and females are territorial and engage in biparental care. Here we observed the behavior of A. trifasciata males and females to construct an ethogram. We registered the parental care of a couple since the spawn until the moment when the offspring became independent. Striking behaviors were observed, such as male-male contests involving circling displays and mouth wrestling, and parental care, which involved mouth brooding and parental color change. Finally, we suggest that our results may be used to establish this species as a model for parental care and territoriality studies.

Hybridization, Behavioral Patterns, and Pre- and Postzygotic Isolation Between Two Recently Derived Species of Walnut-Infesting Rhagoletis Fruit Flies in the Highlands of Mexico.

Episodes of isolation and secondary contact among populations of insects of Nearctic origin during Pleistocene glacial/postglacial climatic cycles had a strong evolutionary influence on the diversity of flies in the genus Rhagoletis in mountainous areas of Mexico. As a series of experiments undertaken to gather support for phylogenetic hypotheses on the origin of three walnut-infesting species in the suavis group, we examined pre- and postzygotic isolation between Rhagoletis completa Cresson, 1929 and R. ramosae Hernández-Ortiz, 1985. Despite morphological, biological, and behavioral differences, these two species were found to be capable of hybridization. Mating experiments in large enclosures revealed asymmetric sexual isolation. There were notable differences in male sexual behavior. While R. ramosae males mated exclusively on host fruit, R. completa males used fruit and alternative mating locations. During fruit-guarding and male-male contests, R. completa and R. ramosae males adopted markedly different wing postures. R. completa females were more reluctant to copulate with heterospecific males than R. ramosae females. During no choice crosses in small enclosures, there was a reduction of egg hatch for the hybrid cross of R. completa males × R. ramosae females. Our results and previous studies on reproductive isolation between other species pairs in the suavis group support a clade in which R. ramosae, R. zoquiBush, 1966, and R. completa are close relatives all still capable of hybridizing.

The hidden cost of group living for aggregating juveniles in a sexually dimorphic species

Abstract The number of conspecifics present during the juvenile stages can have profound consequences on development rates and adult body size, traits often closely related to fitness. Conspecifics can have direct negative effects on each other due to resource competition, and also direct positive effects due to benefits like improved thermoregulation. We investigated morphological and developmental consequences of juvenile group size in the leaf-footed cactus bug Narnia femorata (Hemiptera: Coreidae). These insects are ideal to test the consequences of social environment during development because nymphs naturally aggregate in groups of varying size. Furthermore, the sexual dimorphism of this species allowed us to test for sex-specific effects of developmental density. Males possess enlarged hind legs used as weapons in male-male contests, yet females are physically larger. We found insects from smaller groups had 43% higher mortality than those from the larger groups. On average, adult body and hind leg sizes did not differ across densities for either sex. Interestingly, we found that those first to mature into adults within a sibling group became the largest adults. The largest, fastest males to adulthood also wielded the biggest weapons due to the positive allometry of this trait.

Musculoskeletal mass and shape are correlated with competitive ability in male house mice (Mus musculus).

Intense physical competition between males for mating opportunities is widespread among mammals. In such agonistic encounters, males with combinations of morphological, physiological and behavioral characters that allow them to dominate an opponent have greater fitness. However, the specific physical traits associated with competitive ability are poorly understood. Larger body size is often correlated with fitness in mammals. Interestingly, fitness is maximized at intermediate body masses in male house mice (Mus musculus), a species with a polygynous mating system in which males compete physically for access to reproductive resources. Here, we used competition trials in semi-natural, mixed-sex population enclosures to directly measure competitive ability in male house mice based on control of a preferred nesting site. We tested the hypothesis that the musculoskeletal systems of male mice demonstrating high competitive ability are more specialized for competition by comparing the masses of 10 major muscle groups and eight bones as well as a set of 12 skeletal shape indices associated with anatomical specialization for fighting performance in a set of nine winners and 20 losers. Winning males possessed several traits hypothesized to enhance performance in male-male contests: relatively greater mass in several muscle groups and bones of the forelimb and hindlimb and larger scapular surface area. Unexpectedly, no measurements of the head and neck differed significantly between winners and losers. These results identify musculoskeletal traits associated with competitive ability in male house mice and suggest that our current understanding of mammalian fighting performance is incomplete and more nuanced than previously considered.

Structural coloration predicts the outcome of male contests in the Amazonian damselfly Chalcopteryx scintillans (Odonata: Polythoridae)

Structural coloration predicts the outcome of male contests in the Amazonian damselfly Chalcopteryx scintillans (Odonata: Polythoridae)

Low intrasexual competitiveness and decreasing testosterone in human males (Homo sapiens): the adaptive meaning

Abstract The development of indirect mechanisms of intrasexual competition (e.g., visual identification of possible rivals) could be related to personality traits such as aggressiveness and self-esteem. However, the study of endocrine changes associated to indirect mechanisms of intrasexual competition is scarce. The aim of this study was to investigate the changes in testosterone levels after a rival choice test in men and how intrasexual competitiveness, aggressiveness, and self-esteem modulate these changes. A group of 160 healthy men answered four personality questionnaires, participated in a rival choice test, and donated saliva samples to measure the changes in their testosterone levels. We found a significant decrease in testosterone levels of men with lower intrasexual competitiveness, but testosterone levels remained stables in competitive men. Non-significant results were found for aggressiveness and self-esteem. These decreases in testosterone levels could be interpreted as an adaptation aimed to reduce costs in male-male contests in Western modern societies.