Male mating effort in a polygynous ungulate

Survival and mating success are key fitness components of mammalian males. Because energy is limited, life history theory predicts trade-offs between current and future reproduction. While many studies have examined fitness trade-off in females, we know little about trade-offs faced by males. In polygynous species, male mating success is largely dependent upon intrasexual competition. Consequently, males have greater uncertainty over the benefits of a given allocation than over its costs, and the correlation between mating effort and success is likely much weaker in males than in females. We analyzed 14 years of data on the mating effort and survival of marked bighorn rams to investigate fitness trade-offs. Dominant rams defended single estrous ewes (“tending”) while subordinates attempted to copulate after separating the tending pair (“coursing”). We estimated the participation in tending and coursing for each ram and the effort in searching for breeding opportunities by each ram each year. We compared these three behavioral indices of male mating effort to demographic parameters, individual characteristics, and both yearly and long-term survival. Mating effort during the rut was unrelated to ram overwinter survival, but longevity was positively correlated with mating effort between 2 and 5 years of age. Persistent variation among rams is likely to explain this pattern, suggesting that in natural populations a few high quality males enjoy both high mating success and high survival.An erratum to this article can be found at     

Mating group composition influences somatic costs and activity in rutting dominant male reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>)

In polygynous species, males devote considerable effort to reproduction during the rut. Both the number of females in the mating group and the ratio of sexually mature males to sexually mature females [adult sex ratio (ASR)] are expected to affect the amount of effort a male devotes to reproductive activities. We predicted the reproductive effort of dominant male reindeer, measured as relative mass loss, proportions of active reproductive behaviors, and frequencies of agonistic behaviors would (1) increase with an increasing number of females in the mating group and eventually level off, and (2) exhibit a dome shape with respect to ASR in the mating group. We tested these predictions using 12 years of data collected from semi-domesticated reindeer in northern Finland. We found a positive relationship between relative mass loss and the mean number of females in the mating group for mature, but not young males. The relationship between the proportion of active reproductive behaviors performed by mature males and the mean number of females in the group was quadratic while agonistic behaviors of mature males increased with the increasing female group size. We also found that active reproductive behaviors decreased with a rising mating group ASR for mature males; whereas, young males performed more agonistic behaviors as group ASR increased. Our results point to age-specific patterns of mass loss and activity during the mating season. They also indicate that both the number of females and ASR in the mating group are important factors in determining the level of reproductive effort of dominant male reindeer.     

Age- and state-dependent reproductive effort in male mountain goats, Oreamnos americanus

In mammalian polygynous mating systems, male reproductive effort consists mainly of male–male competition and courting of females, which entail substantial somatic costs. Males are thus expected to adjust their reproductive effort according to their age and condition. In this study, we examined how activity budgets of male mountain goats (Oreamnos americanus), a polygynous ungulate, varied with age in a marked population over two periods: (1) summers 1995–2006 and (2) ruts 2004–2006. We then assessed if the proportions of time spent in male–male competition and courtship behaviors were influenced by age-specific body mass and social rank during the rut. Males spent most of their time foraging and resting during summer, and rested more and foraged less with increasing age. During the rut, pronounced shifts in activity budgets occurred as juveniles (1–2 years) increased time spent foraging, whereas adults (≥3 years) increased standing and time spent in social interactions at the expense of foraging. At old age, reproductive effort either stabilized or decreased slightly, providing weak support for the ‘mating strategy–effort’ hypothesis, predicting that courtship behaviors should peak in prime-aged males. Age-specific body mass did not affect time spent in male–male competition, but was positively related with time spent in courtship behaviors, providing support for the ‘individual quality’ hypothesis, predicting that males with more resources at the start of the rut should spend more time in mating-related activities. Age-specific social rank did not affect reproductive effort. Surviving to prime age while increasing mass each year should thus allow male ungulates to gain greater ability to court estrus females.     

Sexual size dimorphism in fallow deer (Dama dama): do larger, heavier males gain greater mating success?

Sexual size dimorphism may evolve as a result of both natural and sexual selection. In polygynous mammals, the main factor resulting in the evolution of large body size in males is the advantage conferred during competition for mates. In this study, we examined whether sexual selection acts on body size in mature fallow bucks (Dama dama) by examining how the following traits are inter-related: age, body (skeletal) size, body mass, prerut dominance rank, rut dominance rank and mating success. This is the first study to examine how all these factors are together related to the mating success of a large sexually dimorphic and polygynous mammal. We found that male mating success was directly related to body size, but not to body mass. However body mass was related to prerut dominance rank which was in turn strongly related to rut dominance rank, and thus there was an indirect relationship between mating success and body mass. Rut dominance rank was the variable most strongly related to mating success. Mating success among mature males was unrelated to age. We conclude that larger mature fallow bucks have advantages over other males when competing for matings, and sexual selection therefore continues to act on sexual size dimorphism in this species. Heavier fallow bucks also have advantages, but these are mediated through the dominance ranks attained by males before the rut.     

Lek promiscuity in a semelparous mammal,Antechinus stuartii (Marsupialia: Dasyuridae)?

Summary All male Antechinus stuartii die following a brief rut towards the end of their first year of life. Outside the rutting period, both males and females forage in clearly-defined invidual home ranges, but neither sex is territorial. As temperatures drop during the winter, both sexes may leave their foraging range each day to spend the latter part of the night and most of the daylight hours in communal nests which may be used by fifteen animals simultaneously, and over forty individuals during one winter. These temporary movements away from the foraging range cause an increase in the daily home range revealed by radiotelemetry. When the mating season commences, males abandon their foraging range and aggregate in a few of the communal nest trees, where they spend most of the night. There is some movement between aggregations, but the movement appears to be rapid and direct. Females continue to use their foraging area, and nest solitarily. However, they make excursions to the male aggregations, and spend time in the nests with the males. By the third week of the mating season, immediately prior to male mortality, males move frequently from one aggregation to another, but are still confined to the aggregations and direct movements between those nests. By this time some females have ceased visiting the aggregations and nest solitarily within their foraging range. These data suggest that the mating system of A. stuartii is lek promiscuity, and support the hypothesis that the abrupt mortality of males is a cost of endocrine changes which facilitate gluconeogenic mobilisation of body protein, allowing the males to sustain their vigil at the leks.     

Mating effort and female receptivity: how do male guppies decide when to invest in sex?

Males vary in the degree to which they invest in mating. Several factors can explain this variation, including differences in males’ individual condition and the fact that males allocate their energy depending on the context they face in each mating attempt. Particularly, female quality affects male reproductive success. Here, we studied whether male guppies (Poecilia reticulata) strategically allocated more mating effort, in terms of mating behaviour and male–male competition, when they were matched with a receptive (R) female than a non-receptive one. In accordance with our prediction, we found that males increased their mating behaviour when they were with a receptive female. Even though male guppies can inseminate non-receptive females, we only found high levels of courtship between males that were with a receptive female rather than a non-receptive one. Although there was little affect of female receptivity on male–male competition, we found that males chased and interrupted courtships more with receptive females than with non-receptive females regardless of odour. Finally, we also studied whether the sexual pheromone produced by receptive female guppies is a cue that males use in order to increase their mating effort. We found that males were more attracted to a female when they perceived the sexual pheromone, but only increased their mating and aggressive behaviours when females showed receptive behaviour. This strategic increase in mating effort could result in higher male reproductive success because mating attempts towards receptive females are likely to be less costly and males could have a greater probability of fertilisation.     

Multiple signals in the palmate newt: ornaments help when courting

There is increasing evidence that female mate choice is often based on the assessment of multiple male traits, involving both morphology and behavior. We investigated female mate choice for multiple male traits in the palmate newt, Lissotriton helveticus, including male tail filament length, hind foot web size, crest development, body size, ventral coloration, and courtship display activity. Observations of courtship display in the field revealed that females spent more time in front of males with longer tail filaments. Laboratory experiments revealed a more detailed relationship between filament length and courtship display. We found that females took more sperm masses from males with both longer filaments and greater display activity. Experimental shortening of the tail filament length substantially decreased the number of male sperm masses transferred. However, when we experimentally reversed relative filament length between two males in mating trials, male mating success was explained by courtship activity and not by filament length. Our results show that female palmate newts value multiple traits during mate choice, including both morphological ornaments and reproductive behaviors in males. Our results further suggest that, when filament length is below a certain threshold, females may value the information content of courtship activity over that of filament length.     

The significance of hotspots to lekking topi antelopes (<Emphasis Type="Italic">Damaliscus lunatus</Emphasis>)

Controversy has surrounded the question of why lek-breeding has evolved in certain ungulate species. Can the behavior be explained simply by males mapping onto a female distribution that is determined by factors unrelated to mating? Or are leks created because estrous females distinguish between males and favor males who cluster? Here I address these questions by looking at spatial distribution in lekking topi antelopes (Damaliscus lunatus). Contrary to the predictions of a model assuming male clustering in the zone of maximum female range overlap, territories were highly clustered also within this zone, and lek size correlated positively with population density. In support of models derived from the ideal free distribution of males onto female dispersion, leks were in areas with high female density during the rut. However, models not taking into account both individual variation in male quality and female mate preferences failed to explain the extreme male clumping also within high density areas, which was revealed by a strongly male-biased sex-ratio on leks. Additional support for the female preference-based model came from the finding that estrous females concentrated onto leks. Female preference for clustered males may develop if males initially follow an ideal free distribution of unequal competitors with high quality males slightly clustered at density hotspots; positive feedback between female benefits of preference for clustered males and male benefits of clustering could lead to contraction of the territorial network and lek behavior. Thus only the female preference-based model correctly predicted a negative correlation between male mating rate and resource density.     

No relationship between long-distance acoustic mate attraction signals and male fertility or female preference in spring field crickets

To father offspring, a male must succeed at two processes of sexual selection: (1) mate with a female and (2) fertilize her eggs. We investigated the relationships between pre- and post-copulatory male traits and female mating responses in wild-captured and laboratory-reared spring field crickets, Gryllus veletis. The phenotype-linked fertility hypothesis suggests that females may receive a direct benefit, enhanced fertilization efficiency, by mating with males that signal attractively. We measured fine-scale components of male acoustic mate attraction signals as well as how much time males spent signalling, measured female preference for males in mating trials and then quantified sperm number and viability. We found no relationship between male signalling traits and male fertility or female preference, providing no evidence for the phenotype-linked fertility hypothesis. We also found no difference in sperm metrics between wild-captured and laboratory-reared males. While female crickets may receive benefits by choosing males based on acoustic signal characteristics, whether the benefits are a result of genetic quality, seminal fluid contents or some other male trait remains unknown.     

Weapons or mating signals? Claw shape and mate choice in a fiddler crab

Males often possess traits, such as horns, claws, and tusks, which are used during male–male combat. Studies suggest that selection has fine tuned these weapons to improve their effectiveness and that the shape of these weapons provides cues for males to assess the strength of rivals. Similarly, females might also assess a male’s weaponry to determine his value as a mate. The largest weapon relative to body size is the major claw of male fiddler crabs (genus: Uca). Males use their claws both as a weapon and as a courtship signal, waving it to attract females. We used robotic crabs in two-choice experiments to test female mating preferences based on male claw shape in Uca mjoebergi. First, females did not prefer conspecific claws over any of three alternatively shaped stimuli matched for color and for the rate and pattern of waving. The alternative stimuli were two different heterospecific claws and a plain rectangular shape. Second, females significantly preferred the alternative stimuli when they were presented at a faster wave rate. We conclude that claw shape in U. mjoebergi has not evolved under sexual selection imposed by female mating preferences and is more likely to have evolved under selection for effectiveness during male–male competition.     

Males are faster foragers than females: intersexual differences of foraging behaviour in the Apennine chamois

Availability of food resources and individual characteristics can influence foraging behaviour, which can differ between males and females, leading to different patterns of food/habitat selection. In dimorphic species, females are usually more selective in food choice, show greater bite rates and spend more time foraging than males. We evaluated sexual differences in foraging behaviour in Apennine chamois Rupicapra pyrenaica ornata, during the warm season, before the rut. Both sexes selected nutritious vegetation patches and spent a comparable amount of time feeding. However, males had a significantly greater feeding intensity (bite rate) and a lower search effort for feeding (step rate), as well as they spent more time lying down than females. Females selected foraging sites closer to refuge areas than males. In chamois, sexual size dimorphism is seasonal, being negligible in winter–spring, but increasing to 30–40 % in autumn. Our results suggest that males enhance their energy and mass gain by increasing their food intake rate during the warm season, to face the costs of the mating season (November). Conversely, females seem to prioritize a fine-scale selection of vegetation and the protection of offspring. A great food intake rate of males in the warm season could have developed as a behavioural adaptation leading herbivores to the evolutionary transition from year-round monomorphism to permanent dimorphism, through seasonal dimorphism.     

Male genital length and mating status differentially affect mating behaviour in an earwig

Phenotypic variation in male genitalia may affect copulation behaviour, which can have important fitness consequences for males. Male genitalia commonly possess traits that increase male control over copulation, but in species where females control mating, a poor functional understanding often prevents insight into the processes responsible for such effects. Here, I investigate the effect of male genital length on copulation behaviour in the earwig Euborellia brunneri, where both sexes exhibit extremely elongated genitalia that correspond in shape. This model system is particularly suitable because pairs mate repeatedly and females can limit both the number and duration of copulations. I used both virgin and mated males and females in a double-mating design because longer male genitalia confer benefits in sperm competition. Consistent with a greater predicted male mating effort in mated females, the duration of individual copulations increased, but this traded off against mating frequency as cumulative mating duration remained unchanged. In contrast, male genital length increased both individual and cumulative mating duration, regardless of mating status. This difference suggests that, while males may modify copulation duration in response to mating status, females facultatively adjust mating frequency to prevent mating excessively or express preferences for increased male genital length. Notably, this study demonstrates that male genital phenotypes that are successful in sperm competition also enjoy female-mediated mating benefits.     

Providing parental care entails variable mating opportunity costs for male Temminck’s stints

Males of many species theoretically face a fitness tradeoff between mating and parental effort, but quantification of this is rare. We estimated the magnitude of the mating opportunity cost paid by incubating male Temminck’s stints (Calidris temminckii), taking advantage of uniparental care provided by both sexes in this species. “Incubating males” provide all care for an early clutch, limiting subsequent mating possibilities. “Non-incubating” males include males that failed to obtain, lost to predation, or actively avoided incubating clutches. These males were free to pursue extrapair copulations and to mate with females laying later clutches, which females usually incubate themselves. Male incubation classes did not differ in measures of quality, and many individuals changed classes between years, suggesting the use of conditional reproductive tactics. However, specialist non-incubators may also exist. Using microsatellites to assign parentage, we compare males’ total fertilizations and the subset “free of care” fertilizations between incubation classes. Incubators were more likely to gain at least one fertilization per season and averaged one more per season than non-incubators. However, successful non-incubators were more likely to gain “free of care” fertilizations, averaging two more than successful incubators. The relative success of male incubation classes also changed with local sex ratios. With higher female proportions, non-incubators gained disproportionately more offspring, suggesting that the use of tactics should be partly determined by the availability of potentially incubating females. Overall, we estimate the opportunity cost of incubating to be 13–25 % of the potential annual reproductive output.     

The effect of male sodium diet and mating history on female reproduction in the puddling squinting bush brown<Emphasis Type="Italic"> Bicyclus anynana</Emphasis> (Lepidoptera)

The males of butterflies transfer a spermatophore to the female during mating that can contain nutrients enhancing the reproductive potential of their partners. The nutrients transferred by males can be derived from both larval and adult feeding. These nutrients may be depleted by multiple matings. An apparent difference in adult feeding behaviour between the sexes is puddling on mud, dung and carrion, which in most butterfly species is exclusively a male behaviour. A possible explanation for this division in feeding behaviour is that nutrients derived from puddling by males are transferred to the female during mating. Here, we test this hypothesis in the African fruit-feeding butterfly Bicyclus anynana. We varied the male nuptial gift by (1) feeding males either a diet with or without sodium, and (2) varying the number of previous successful copulations by remating males up to five times on consecutive days. The results show both a strong effect of order of mating on the mating duration, and an individual effect with some males typically copulating for a shorter time than others. The effects on female reproduction were, however, minimal. The total number of eggs per female and the sodium content of the eggs did not differ significantly between diets, nor were they affected by the mating histories of the males. Eggs showed a non-significant lower hatching for females partnered by a male who had already mated several times. There was an indication of an interaction with male diet: the sodium treatment showing a decline in egg hatchability with order number of male mating, whilst the control treatment showed a constant hatchability. The results are discussed in relation to determinants of male gift-giving strategy and to other potential explanations for a restriction of puddling to males in butterflies.Communicated by M.A. Elgar     

Repeatable female preferences, mating order and mating success in the poeciliid fish, Heterandria formosa

Sexual selection via female choice can afford preferred males comparably higher mating success than those males that lack preferred traits. In addition, many models of sexual selection assume that both male traits and female preferences are heritable. In this study we test whether females of the poeciliid fish, Heterandria formosa, have repeatable pre-copulatory preferences for larger males. We also test whether female pre-copulatory preferences are always reliable indicators of male mating success. When given a choice between a large and a small male, females prefer larger males, and the repeatability of this preference is high. Although there are no overall differences in male mating success between large and small males, large males have a higher mating success when they are the first to mate than when they are the second to mate. Likewise, preferred males also have higher mating success when they are the first to mate than when they mate second. Therefore, the repeatable female preferences observed in this study only predict male mating success when the preferred male mates first. These results illustrate that even significantly repeatable female preferences do not translate into male mating success, which is an assumption of many examinations of the importance of female choice in sexual selection.     

Male mating competitiveness and age-dependent relationship between testosterone and social rank in bighorn sheep

In males, the acquisition and development of behavioral and morphological secondary sexual traits typically depends on testosterone and correlates with mating success. Testosterone level could affect competition for mates and thus be a target of sexual selection. We sought to relate testosterone levels to male mating competitiveness, by teasing apart the relationships between testosterone, behavior, and growth before the mating period. We monitored 24 adult bighorn rams (Ovis canadensis) at Ram Mountain, Alberta, from 2008 to 2011. Using linear mixed models, we tested the relationships between testosterone metabolites in feces, social rank, and both growth and size of two sexually selected traits: horns and body mass. The correlation between testosterone and social rank varied with age. Testosterone and rank were weakly and negatively correlated for young rams, positively correlated for prime-aged rams, and negatively correlated for older rams. Although testosterone had an increasingly positive effect on total horn length until 8 years of age, we could not detect any effects on annual growth rate of horns or body mass. Testosterone may be related to male’s ability to compete for mates through its relationship with behaviors determining social rank, rather than by influencing the development of morphological traits. Differences in testosterone levels among competitors may be a proximate cause of variance in fitness.     

Correlates of male mating success in great bustard leks: the effects of age, weight, and display effort

We examined how mating success varied in relation to age, weight, body size, and display behavior among great bustard Otis tarda males. The estimated mating success was strongly skewed, with 45% of adult males being involved in copulation attempts and only 9.7% actually seen copulating successfully. Unlike most birds, body size continued increasing in great bustards several years after reaching sexual maturity. Age, weight, and display effort were all significant and independent predictors of male mating success. The higher display effort involved performing longer full-display bouts. Older males could detach from the male flock earlier in the season as well as on each day and spend longer seasonal and daily periods displaying as solitary birds, which contributed to increase their mating success. In contrast, males weighing more did not invest more in display, which suggests that they could be recognized as dominants by other males and selected by females through assessment of their plumage sexual traits. In contrast to most other bird species, the system described for great bustards resembles that found in some lek-mating ungulates, where social rank is a complex trait determined by both age and mass, and as in these mammals, it suggests that sexual selection continues to favor a high male weight in this extremely sexually dimorphic species.     

Phenotypic correlates and survival consequences of male mating success in lek-mating greater prairie-chickens (<Emphasis Type="Italic">Tympanuchus cupido</Emphasis>)

Female choice and male–male aggression are two modes of sexual selection that can lead to elaboration of male morphological and behavioral traits. In lek-mating species, male mating success is often strongly skewed, and it is puzzling why variation in male traits is still observed given directional female choice. If male traits correlated with reproductive success are honest signals of male quality, there may be survival costs associated with the expression of those traits. In this study, we examined whether morphological, behavioral, and territorial traits are correlated with male mating success and survival in the lek-mating greater prairie-chicken (Tympanuchus cupido). We introduce a novel application of multinomial discrete choice models for analysis of female mate choice behavior. We found that behavioral and territorial attributes showed 6.5 times more variability among males than morphological traits. Both display and aggressive behaviors were strong predictors of male mating success, suggesting that both female choice and male–male aggression were important in determining mating success among male greater prairie-chickens. Moreover, annual survival of male prairie-chickens was independent of mating success and male traits. Females appear to be choosing males based on behavioral traits where large variation exists between males (coefficient of variation >30%). Behavioral traits were the most important factor in determining mating success of male prairie-chickens, but the mechanism underlying this relationship is unknown. In the future, experimental manipulations of male hormones or parasite loads could bridge the proximate mechanisms and ultimate consequences of factors mediating male mating success in lek-mating grouse.     

Female encounter rate and the calling behavior and mating propensity of male Requena verticalis (Orthoptera: Tettigoniidae)

Mating effort, the energy exerted in finding and persuading a member of the opposite sex to mate, may be influenced by how frequently potential mates are encountered. Specifically, males that frequently encounter females may reduce calling effort and be less eager to mate than males that infrequently encounter females. An experiment was set up to test this hypothesis, using the tettigoniid Requena verticalis. We examined the song structure, calling activity and mating propensity of individual males exposed to one of five different encounter rates with virgin females. Song structure and calling effort were significantly altered by an encounter with a female. After an encounter, males significantly increased chirp rate and decreased variability in interchirp interval. Encounters also stimulated a male to call and to continue to call for up to two hours. The elapsed time since mating affected mating propensity but not calling activity. Mating propensity asymptotically increased to reach a maximum by day 17 since last mating. However, neither the frequency of encounters, nor the number of previous encounters experienced by a male, influenced calling activity or the propensity of a male to mate. The significance of changes in song structure and calling activity following an encounter, and of increasing male mating propensity over time, are discussed. Correspondence to: G.R. Allen     

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.