The role of body size and fighting experience in predicting contest behaviour in the black field cricket, <Emphasis Type="Italic">Teleogryllus commodus</Emphasis>

Body size strongly predicts fighting behaviour and outcome in many species, with the larger opponent usually winning contests. However, recent fighting experience can have a strong influence on the establishment of dominance hierarchies, with recent winners being more likely to win subsequent contests, while recent losers are more likely to lose. Recent fighting experience therefore has the potential to modify the effect of body size in determining contest behaviour and outcome. Here, we investigate whether recent fighting experience weakens the role of body size in predicting contest behaviour in the black field cricket, Teleogryllus commodus. We compared the role of body size in determining contest outcome during initial non-physical encounters and escalated aggressive physical encounters (grapples), as well as the probability of escalation occurring, in contests involving either a naïve or experienced smaller male against a naïve larger male. We found that recent fighting experience only affected contest outcome during non-physical encounters. Once a contest had escalated into grappling, the effect of previous wins and losses was no longer apparent and body size strongly predicted contest outcome. Thus, once males can directly assess their opponent's fighting ability, recent fighting experience did not alter the effect of body size on contest behaviour and outcome.     

Effect of losing on male fights of broad-horned flour beetle, Gnatocerus cornutus

Experience-dependent tactics of males trying to gain access to females were examined in the beetle Gnatocerus cornutus, which engages in male–male fighting for mates. In male fights, subsequent behavior is modified by winning and losing experiences. Males, therefore, may choose between several behavioral tactics to obtain a mate, based on his previous fighting experience. In G. cornutus, we examined for how long aggressive behaviors were modified by experiences of winning or losing, i.e., the duration of the prior experience effect. Losing decreased a male's frequency of fighting for 4 days, and few defeated males fought any male regardless of the opponent's size. By the fifth day, this effect disappeared. No modulation of male behavior due to winning was observed. Furthermore, the experience of losing not only decreased a male's aggressiveness but also switched the male behavior from fighting to dispersal from the fight site to another site. In future, it is necessary to clarify why the optimal term of the losing experience is 4 days in this beetle.     

Fighting behaviour as a correlate of male mating success in black grouse Tetrao tetrix

Fighting is a fundamental determinant of male fitness in species where females prefer socially dominant males as mates or where dominants can prevent subordinates from mating. This in turn can lead to the evolution of honest inter- and intra-sexual cues of male dominance. Fighting as a behaviour comprises both fighting rate (number of fights per unit of time) and fighting performance (success in winning fights), but it is not always clear which of these components are important for female choice and how they link to signals of male quality. To quantify the relative importance of fighting as a cue for females, we recorded detailed behavioural data from male black grouse Tetrao tetrix at leks. We explored the relationship between phenotypic traits (body mass, eye comb size, tail (lyre) length and blue chroma colouration) and fighting performance and rates and how these were related to male mating success. In older males' pairwise fights, winners had lower blue chroma than losers, but there were no differences in other morphological traits. In yearlings, no morphological trait predicted success in pairwise contests. Both fighting rate and performance were positively related to the number of copulations acquired by a male; however, when controlled for lek centrality, fighting performance and not fighting rate was significantly related to mating success. Our results indicate that females may be using components of fighting behaviour as cues for mate choice.     

Previous experience matters in the stalk-eyed fly <Emphasis Type="Italic">Teleopsis dalmanni</Emphasis>

Previous experiences can play a significant role in determining future behaviors. Winner and loser effects, where the outcome of previous aggressive encounters influences the behavioral approach to and outcomes of future conflicts, have been documented in many taxa and illustrate this phenomenon. These effects are prevalent in species that interact frequently because modulation of these potentially costly social interactions may influence fitness. Stalk-eyed flies of the dimorphic species Teleopsis dalmanni engage in frequent fights over food resources, as well as over access to harems of females, with larger males typically prevailing when size disparities exist. However, whether and how prior experience influences fighting decisions and outcomes remains unexplored. To test for winner and loser effects in stalk-eyed flies, sexually mature flies were paired in size-mismatched dyads to establish winning and losing experiences. After their first contest, the flies were paired with size-matched individuals and allowed to interact. We determined whether an initial winning or losing experience significantly altered the outcome probabilities in the second size-matched encounter. Initial winning experience did not significantly affect the second interaction, providing no evidence for a winner effect. However, initial losers were significantly more likely to lose a subsequent interaction which provides evidence for a loser effect in stalk-eyed flies. In addition, smaller males experienced an increased probability of losing their second interaction regardless of prior winning or losing experience. This effect was not seen in large males. Our data suggest that the loser effects we observed, which were more pronounced in small males, could result from the energetic costs of fighting that they were less able to absorb than large males.     

A lover, not a fighter: mating causes male crickets to lose fights

Both resource-holding potential (RHP) and experience in aggressive contests are known to affect future aggressive behaviour. However, few studies have examined the effects of mating experience on agonistic behaviour, despite the fact that dominant males usually acquire more matings. We investigated the effect of mating experience on male aggressive behaviour including the relationship between RHP and fighting success in the fall field cricket, Gryllus pennsylvanicus. We formed pairs of size- and age-matched males that varied in RHP (relative weapon size) and conducted two experiments. In the first, we varied male mating experience by allowing one male in a pair to either be (a) ‘mated’: court, be mounted and copulate with a virgin female or (b) ‘experienced’: court, be mounted, but prevented from copulating. The second experiment varied postcopulatory experience where the male was allowed (‘contact’) or prevented from (‘no-contact’) continued contact with his recent mate. Following treatment, experimental males engaged in an aggressive contest with the naïve size- and age-matched male. In our first experiment, we found that mated and experienced males were equally likely to escalate contests to combat with a naïve opponent, but mated males were less likely than experienced males to win. There was no effect of mating on the relationship between RHP and fighting success. In our second experiment, we found no effect of maintaining contact with the female on the tendency to escalate or the probability of winning. As in the first experiment, males with relatively larger heads again won more fights and this relationship was unaffected by male experience. These results suggest that mating is itself detrimental to male success in aggressive contests, but that this effect is not sufficient to eliminate the effect of RHP on fighting success.     

Lack of innate preference for morph and species identity in mate-searching <Emphasis Type="Italic">Enallagma</Emphasis> damselflies

Insect mate recognition is often viewed as stereotypic, innate, and species-specific. However, male damselflies can learn to identify female-specific color morphs as potential mates. A suite of male mimicry hypotheses assume that heteromorphic females, which differ from males in color pattern, are more easily recognized as “female” and thus lack the inherent, anti-harassment advantage that the more male-like signal provides for andromorphs. Using two measures of male preference, we investigated whether naïve males have a preexisting sensory bias for a given morph color in Enallagma civile, a species that appeared to exhibit extreme plasticity in morph expression across generations within a breeding season. E. civile males raised in the absence of females exhibited no preference for either morph, whereas males raised with one female type exhibited a learned sensory bias for that morph. Male Enallagma also lacked a bias toward conspecific females over a congeneric sister species. In a naturally naïve population of Enallagma ebrium, males reacted sexually to both morphs of Enallagma hageni as often as they did to conspecific females, whose thoracic spectra were nearly identical with those of E. hageni. Moreover, despite the similar thoracic spectra of males and andromorphs, both of which reflected UV, males rarely reacted sexually to other males. Our results falsified implicit assumptions of male mimicry hypotheses, supported learned mate recognition, and suggested a scenario for speciation via sexual conflict.     

The effects of male–male contests and female eavesdropping on female mate choice and male mating success in the jumping spider, <Emphasis Type="Italic">Thiania bhamoensis</Emphasis> (Araneae: Salticidae)

In experiments that comprised of three phases (fight, choice, and mating) under “seen” and “unseen” conditions, we examined the effects of the outcomes of male–male contests and female eavesdropping on female mate choice and male mating success in the fighting spider, Thiania bhamoensis (Salticidae). The results revealed female eavesdropping on agonistic interactions. Females that had watched an aggressive interaction showed no distinctive preference for the winner over the loser, but they preferred the loser when they had not observed a fight. Winners, however, achieved a greater mating success than did losers during the mating phase. Gaining access to females was based on the insistence of the winners in courtship in terms of the number of quivers, rather than on the fighting behavior of the males. Hence, the outcome of male–male contests may not be an important determinant of a male’s mating success in T. bhamoensis. Instead, courtship display plays an important role in determining the success of male mating in this species. This study also suggests that female mate preference may not be a good indicator of eventual female mate choice and male mating success. Thus, a causal relationship between female mate preference and male mating success cannot be inferred. Joanna P. Y. Chan, Pei Rong Lau, and Ai Jie Tham contributed equally.     

Resource-holding power asymmetries,the prior residence effect,and reproductive payoffs in male northern elephant seal fights

The effect of resource-holding power (RHP) and prior residency asymmetries on fight outcome and subsequent seasonal copulatory success was analyzed for fights between marked male northern elephant seals (Mirounga angustirostris). RHP asymmetries were measured as differences in estimated mass and prior residency asymmetries were measured as differences in beach tenure prior to the fight. The principal results were: (a) Neither differences in mass nor differences in beach tenure had any effect on fight outcome as separate factors. (b) Mass and tenure differences had an interactive effect on fight outcome; fight winners were either heavier males present for shorter periods (intruders) or lighter males present for longer periods (prior residents). (c) Winners of fights copulated more often than losers after a fight throughout the breeding season; this difference was smallest for low-ranking males, larger for high-ranking males in short fights, and greatest for high-ranking males in long fights. (d) Prior resident males who won long fights obtained significantly more copulations after a fight than the males they defeated, but this was not true for intruder males who won long fights. These results suggest that male northern elephant seals will incur greater contest costs (i.e., fight for longer periods and/or against heavier males) for higher reproductive payoffs. They also imply that, at least for males in long fights, differences in prior residence represent payoff asymmetries, with higher reproductive payoffs for winning prior residents than for winning intruders.     

Females prefer males with superior fighting abilities but avoid sexually harassing winners when eavesdropping on male fights

Selection imposed by male competition (intrasexual selection) and female choice (intersexual selection) can be con- or discordant. Specifically, females may or may not prefer mating with dominant males, and direct costs of interacting with dominant (and possibly more harassing) males have been suggested to explain avoidance of dominant males. Here, we exemplify that inter- and intrasexual selection may normally act in the same direction, but can be temporarily conflicting when social information becomes available. Using video playback techniques, we presented females of the Mexican livebearing fish Poecilia mexicana with two size-matched males and established association preferences. Half of the females could then observe the same two males fight and establish dominance, while control females saw both males side by side, but physically separated, and female preferences were subsequently re-evaluated. Females in the control group showed a significant preference for future winners in the subsequent testing, confirming an innate or acquired preference for male traits that are indicative of physical superiority, even when body size as a choice criterion is excluded. When allowed to eavesdrop on male fights, however, females did not show a preference for observed winners and even decreased time spent with them relative to the control treatment in which no fight was shown. A subsequent experiment found contest winners to show elevated levels of sexual behavior, so we argue that the temporary offset of the intrinsic female preference for dominant males after having observed a fight is indeed driven by direct costs females expect from more harassing contest winners.     

Weapon size versus body size as a predictor of winning in fights between shore crabs, Carcinus maenas (L.)

Relative body size (carapace width) and weapon size (chela length) were used as indicators of resource holding potential (RHP) in the agonistic behaviour of male shore crabs, Carcinus maenas (L.). Weapon size was found to be a more reliable predictor of the outcome of pairwise fights than body size. Crabs with longer chelae than their opponents were more likely to win fights than crabs with relatively larger bodies. Body size had less influence on the outcome of fights. Relative body and weapon size did not influence initiation of contests but did affect the likelihood of winning; however, this was significant only for weapon size. Winning crabs had heavier claws with greater surface area than losing crabs. There was no relationship between relative size and fight duration. The frequency of cheliped display increased with chela length and win- ners performed significantly more displays than losers. Received: 5 February 1997 / Accepted after revision: 20 May 1997     

Immune function, dominance and mating success in drumming male wolf spiders Hygrolycosa rubrofasciata

In this study, we provide a piece of experimental evidence that immune function is related to dominance and mating success in wild caught male wolf spiders, Hygrolycosa rubrofasciata. In the mating season, H. rubrofasciata males are actively searching for receptive females, and while searching males often engage in agonistic behavior (i.e., agonistic drumming signals, chases, and fights) with each other. The present results demonstrate that dominant males had higher lytic activities in their hemolymph than subordinates. Lytic activity estimates the concentration of antimicrobial peptides with lysozyme-like activity in hemolymph, which have been shown to play an important role in defense against bacteria, viruses, and fungi. Dominants also had higher courtship drumming rates than subordinates. Moreover, winners in mating competition had higher lytic activities than losers, but this was measured nonindependently of dominance status. Among males with mating failure, there was a moderate negative correlation between encapsulation rate and courtship drumming rate, suggesting that low quality males might not be able to bear the immunological costs of courtship behavior. These results suggest that females might gain immunological benefits by preferring vigorous males.     

Bright turquoise as an intraspecific signal in the chameleon grasshopper (Kosciuscola tristis)

Bright colours often communicate important information between conspecifics. In sexually dichromatic species where males exhibit bright colours, two hypotheses are often invoked to explain the function of the colour. First, if a male’s bright colour contains information about his quality, females may prefer brighter males. Equally, male colour may reliably provide other males with information about fighting ability or resource holding potential. In such circumstances, brighter males may win altercations and/or males may use rival colour to assess their likelihood of winning an interaction. In the chameleon grasshopper (Kosciuscola tristis), males but not females turn bright turquoise when their body temperature exceeds 25 °C. In this study, we tested whether the turquoise phase of colour change has a signaling role in inter- and intrasexual contexts. We predicted that females would prefer bright turquoise males over dull males, but found no evidence from several choice experiments to support this hypothesis. We also predicted that brighter males would win more fights than duller males. Whilst we did not find that brighter males won more fights in staged experiments, we found that the brightness of males who chose to enter fights was significantly correlated with their opponents’ brightness. Our results suggest that the brightness of males’ turquoise phase may provide competitors with important information about their rival’s fighting ability.     

Females reward courtship by competing males in a cannibalistic spider

Despite widespread recognition that intersexual interactions shape reproductive strategies, studies of male competition do not typically include effects imposed by females. In cannibalistic redback spiders, escalated fighting between rival suitors is predicted, as males are unlikely to mate with more than one female, and strong first-male sperm precedence favours mating with virgins. In staged competitions for matings between size mismatched rivals, smaller males adopted an alternative sneaking strategy. However, despite initial agonistic interactions, larger males did not pursue or incapacitate smaller males. When inter-male competition occurred, females struck at males frequently, although strikes were rarely seen when males courted in the absence of a rival. After minimal fighting, larger males engaged in significant courtship (3 h) rather than killing inferior rivals. Prolonged courtship was favoured by female behaviour, as males that attempted rapid copulation (smaller, sneaking males) were cannibalised before mating was completed. This premature cannibalism significantly decreases paternity in redback spiders. Thus, significant features of male competitive behaviour (i.e. prolonged courtship by larger males) may be predicted with consideration of the female’s response to male reproductive strategies. Although the effect of females may be more subtle in systems without the extreme reversed size-dimorphism of redbacks, these results suggest that female interests should be explicitly considered when studying inter-male interactions.     

Prior encounters with the opposite sex affect male and female mating behavior in a wolf spider (Araneae,Lycosidae)

Mate availability can vary widely in nature depending upon population density and sex ratio and can affect the ability of individuals to be selective in mate choice. We tested the effects of prior encounters with the opposite sex (i.e., exposure to the opposite sex either with or without mating) on subsequent mating behavior in two experiments that manipulated mate availability for both males and females in the wolf spider, Hogna helluo. The probability of mating in the experimental trial depended upon whether the prior encounter involved mating or not, and males and females responded in opposite directions. Exposure without mating resulted in a higher subsequent frequency of mating for females and a lower subsequent frequency of mating for males, while prior mating experience resulted in a lower frequency of female remating and a higher frequency of male remating. Prior exposure without mating did not affect female aggression. However, mated females engaged in precopulatory cannibalism more frequently than virgins. Mated males escaped postcopulatory cannibalism more frequently than virgins. Our results show that males respond to exposure without mating in the expected manner. However, prior mating (1 week earlier) had unexpected effects on males, which may be due to mated males being of higher quality. There were little or no effects of the size of the prior exposure individual or mate on subsequent mating behaviors. Further research is needed to determine why different species use different degrees of prior information in mate choice.     

"Cryptic" direct benefits of mate choice: choosy females experience reduced predation risk while in precopula

Despite the central role that female mate choice plays in the production of biological diversity, controversy remains concerning its evolution and maintenance. This is particularly true in systems where females are choosy but do not receive obvious direct benefits such as nuptial gifts that increase a female's survival and fecundity. In the absence of such direct benefits, indirect benefits (i.e., the production of superior offspring) are often invoked to explain the evolution of mate choice. However, females may receive less obvious, or "cryptic," direct benefits, particularly in species with prolonged pre-mating interactions (e.g., precopulatory mate guarding). We assessed the “cryptic” direct benefits of female choice for large male size in two species of freshwater amphipods that do not receive obvious direct benefits. Females paired with large males experienced decreased predation from fish. However, we found that the size of a female’s mate did not affect her predation risk against predatory dragonflies or the harassment she received by single males while paired. Our results demonstrate that even when females receive no traditional direct benefits, female choice for large male size can still provide important direct benefits. Such “cryptic” direct benefits may be common, especially in species with prolonged mating interactions, and are likely important for fully understanding the evolution of mate choice.     

Mate choice plasticity in the field cricket <Emphasis Type="Italic">Teleogryllus oceanicus</Emphasis>: effects of social experience in multiple modalities

Social experience can elicit phenotypically plastic changes in mate choice, but little is known about the degree to which social information from one modality can influence mating decisions based on information from a different modality. I used the field cricket Teleogryllus oceanicus to test whether experience of chemical cues mimicking a high density of sexually mature males causes changes in mate choice based on acoustic signals. T. oceanicus males produce long-range calling songs to attract females for mating, but they also produce waxy, non-volatile hydrocarbons on their cuticle (CHCs) which, when deposited on a substrate, can be detected by females and may provide demographic information. I manipulated female experience of substrate-bound male CHCs and then performed acoustic mate choice trials. When CHCs were present on the substrate during trials, females showed greater motivation to respond to male calling song. This effect diminished with repeated exposure to male songs, demonstrating that the importance of olfactory cues in altering acoustic mate choice decreased with increasing exposure to acoustic signals. However, the temporal nature of CHC experience mattered: previous experience of CHCs did not alter subsequent female choice for male calling song traits. Exposure to male song increased the threshold of mate acceptance over time, and individuals varied considerably in overall levels of responsiveness. Taken together, the results demonstrate that mate choice is dependent on social context mediated by multiple modalities in T. oceanicus, but they do not support the idea that prior experience of social cues in one modality necessarily influences later mating decisions based on other signalling modalities.     

Mate preferences by male guppies (Poecilia reticulata) in relation to the risk of sperm competition

We investigated male mate preferences in relation to the perceived risk of sperm competition in the guppy (Poecilia reticulata), a freshwater fish with a promiscuous mating system. Our laboratory experiments revealed that male mate choice behaviour is not influenced by the presence of rival males that are merely in close proximity to a potential mate, as there was no significant difference in the amount of time that males spent with females that were recently either alone or in close proximity to four rival males. Male mate choice behaviour was, however, strongly influenced by the presence of rival males in a second experiment, where those rivals were permitted to copulate with one of the females. In that situation, males spent significantly more time with, and directed significantly more sigmoid courtship displays toward, females that they had recently seen alone compared to females they had seen receiving forced copulations from up to four rival males. Our results therefore indicate that male guppies are sensitive to the risk of sperm competition and alter their mate choice behaviour in an adaptive fashion.Communicated by K. Lindström     

Sexual voyeurs and copiers: social copying and the audience effect on male mate choice in the guppy

In most mating systems, males and females are commonly within signalling and receiving distance of conspecifics during courtship and mating activities. Although it is well known that females who observe sexual interactions between conspecifics will use public information obtained from these interactions when making their own mating decisions, much less is known about whether males use this type of information in making mating decisions. We used the Trinidadian guppy (Poecilia reticulata) to test whether males use public information to (i) copy the apparent mate choice of another male and (ii) modify their mating preference for a given female in the presence of one or two sexual rivals (potential copiers). We show that males use public information to copy the mate choice of other males and that males alter their mating preferences in response to the presence of an audience of sexual rivals, but find no evidence of a stronger audience effect when the number of sexual rivals increases. Collectively, these results indicate that males pay attention to their immediate social environment in making mating decisions and suggest that they avoid having another male copy their mate choice by weakening or even reversing their initial mating preference in the presence of eavesdropping male sexual competitors. Our findings highlight the importance of social context and public information in male mate-choice decisions and have implications for the evolution of male mating preferences and of social information use in populations.     

Male moths undertake both pre- and in-copulation mate choice based on female age and weight

Variation in reproductive potential usually occurs among individuals of both sexes; for example, some individuals may carry more gametes or be able to continue to generate more gametes in their reproductive life than others. Therefore, to maximize their reproductive success, both sexes are expected to show adaptations for mate choice. However, most authors concentrated on how females choose their mates with the belief that females invest more in reproduction than males, and males are generally eager to pair with any female. In this paper, we report our work on a polygamous moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), with special emphasis on male pre- and in-copulation mate choice in relation to the reproductive quality and re-copulation potential of females. We show that in E. kuehniella, younger and heavier females have significantly higher reproductive value than older and lighter ones, and male sperm supply significantly decreases over successive copulations. Males exercise pre-copulation mate choice by selecting females with higher reproductive potential for copulation and in-copulation mate choice by allocating more sperm to females with higher reproductive potential. However, high-quality females are more likely to re-copulate than low-quality ones, and allocation of more sperm to females by males does not increase female reproductive outputs. It is suggested that the allocation of more sperm to high-quality females functions to increase sperm competitiveness against rivals.     

Do female association preferences predict the likelihood of reproduction?

Sexual selection acting on male traits through female mate choice is commonly inferred from female association preferences in dichotomous mate choice experiments. However, there are surprisingly few empirical demonstrations that such association preferences predict the likelihood of females reproducing with a particular male. This information is essential to confirm association preferences as good predictors of mate choice. We used green swordtails (Xiphophorus helleri) to test whether association preferences predict the likelihood of a female reproducing with a male. Females were tested for a preference for long- or short-sworded males in a standard dichotomous choice experiment and then allowed free access to either their preferred or non-preferred male. If females subsequently failed to produce fry, they were provided a second unfamiliar male with similar sword length to the first male. Females were more likely to reproduce with preferred than non-preferred males, but for those that reproduced, neither the status (preferred/non-preferred) nor the sword length (long/short) of the male had an effect on brood size or relative investment in growth by the female. There was no overall preference based on sword length in this study, but male sword length did affect likelihood of reproduction, with females more likely to reproduce with long- than short-sworded males (independent of preference for such males in earlier choice tests). These results suggest that female association preferences are good indicators of female mate choice but that ornament characteristics of the male are also important.