Male Reproductive Strategies Research Articles

Male consort behavior is driven by female mate choice and receptive synchrony in Tibetan macaques (Macaca thibetana)

In group-living animals, males’ fighting abilities were usually mediated via dominance rank to sort males into a queue to access fertile females, especially in non-human primates. However, the correlation between male reproductive success and consortship, an alternative high-ranking male reproductive strategy, varies across species, obscuring the adaptive significance of male consort behavior. The priority of access (PoA) model is commonly used to predict male mating/reproductive success based on male dominance rank and female reproductive synchrony. In this study, we aimed to assess the applicability of the PoA model in predicting the distribution of male mating success and consortship, while also identifying the main factors influencing these outcomes in a group of free-ranging Tibetan macaques (Macaca thibetana) at Huangshan, China. Our results showed that (1) the male mating was significantly skewed towards the highest-ranking males, deviating from the expectations set by the PoA model. This departure was attributed to both female mate choice and male consort behavior. Notably, the influence of male rank on mating success decreased as the number of receptive females increased. Additionally, female mate choice became less significant when consorted by males compared to situations when they were not consorted; (2) the observed consort behavior among males was lower than what the PoA model predicted for available receptive days, while high-ranking males exhibited a gradual increase in their consort behavior with the rising number of receptive females. Our results suggests that, in Tibetan macaques, male consort behavior imposed limitations on female mate choice, and mitigated the impact of male dominance rank on male mating success as the number of receptive females grew. We propose that male consort behavior could be driven by female counter-monopoly strategies, and the simultaneous occurrence of consort behavior and opportunistic mating may represent the most effective mating tactic in this promiscuous mating system.

Growth increase and gonadal dysfunction of the lined seahorse triggered by zinc exposure

Seahorses are characterized by unique characteristics such as a male pregnancy reproductive strategy and grasping preferences, which make these species vulnerable to various environmental risks. Zinc (Zn) is one of the most frequently occurring toxic elements in coastal waters; however, little is known about the effect of Zn exposure on seahorses. In the present study, line seahorses (Hippocampus erectus) were exposed to waterborne Zn (0.2 and 1.0 mg/L) and the impact on growth and gonadal development was investigated. Zn exposure induced growth improvement, but also led to gonadal dysfunction in the lined seahorse. Female seahorses exhibited high testosterone levels, immature follicles, and weight increase after Zn exposure, which is the typical characteristics of a polycystic ovary syndrome (PCOS)-like phenotype. Transcriptomic data suggested that the Zn-induced growth promotion resulted from the dysregulated expression of fat accumulation genes. Further investigation of gene expression profiles in the brain, ovaries, and testes indicated that Zn affected the expression of genes involved in growth, immunity, tissue remodeling, and gonadal development by regulating serum steroid hormone levels and androgen receptor expression. This study not only clarifies the complex impact of Zn on seahorses using physiological, histological, and molecular evidence but can also provide new insights into the mechanism underlying PCOS in reproductive-aged women. Moreover, this work demonstrates the risk of the common practice of Zn supplementation in the aquaculture industry as the consequent growth yield may not represent a healthy condition.

A new approach for solving global optimization and engineering problems based on modified sea horse optimizer

Abstract Sea horse optimizer (SHO) is a noteworthy metaheuristic algorithm that emulates various intelligent behaviors exhibited by sea horses, encompassing feeding patterns, male reproductive strategies, and intricate movement patterns. To mimic the nuanced locomotion of sea horses, SHO integrates the logarithmic helical equation and Levy flight, effectively incorporating both random movements with substantial step sizes and refined local exploitation. Additionally, the utilization of Brownian motion facilitates a more comprehensive exploration of the search space. This study introduces a robust and high-performance variant of the SHO algorithm named modified sea horse optimizer (mSHO). The enhancement primarily focuses on bolstering SHO’s exploitation capabilities by replacing its original method with an innovative local search strategy encompassing three distinct steps: a neighborhood-based local search, a global non-neighbor-based search, and a method involving circumnavigation of the existing search region. These techniques improve mSHO algorithm’s search capabilities, allowing it to navigate the search space and converge toward optimal solutions efficiently. To evaluate the efficacy of the mSHO algorithm, comprehensive assessments are conducted across both the CEC2020 benchmark functions and nine distinct engineering problems. A meticulous comparison is drawn against nine metaheuristic algorithms to validate the achieved outcomes. Statistical tests, including Wilcoxon’s rank-sum and Friedman’s tests, are aptly applied to discern noteworthy differences among the compared algorithms. Empirical findings consistently underscore the exceptional performance of mSHO across diverse benchmark functions, reinforcing its prowess in solving complex optimization problems. Furthermore, the robustness of mSHO endures even as the dimensions of optimization challenges expand, signifying its unwavering efficacy in navigating complex search spaces. The comprehensive results distinctly establish the supremacy and efficiency of the mSHO method as an exemplary tool for tackling an array of optimization quandaries. The results show that the proposed mSHO algorithm has a total rank of 1 for CEC2020 test functions. In contrast, the mSHO achieved the best value for the engineering problems, recording a value of 0.012 665, 2993.634, 0.01 266, 1.724 967, 263.8915, 0.032 255, 58 507.14, 1.339 956, and 0.23 524 for the pressure vessel design, speed reducer design, tension/compression spring, welded beam design, three-bar truss engineering design, industrial refrigeration system, multi-product batch plant, cantilever beam problem, and multiple disc clutch brake problems, respectively. Source codes of mSHO are publicly available at https://www.mathworks.com/matlabcentral/fileexchange/135882-improved-sea-horse-algorithm.

Dimerization of melanocortin 4 receptor controls puberty onset and body size polymorphism.

Xiphophorus fish exhibit a clear phenotypic polymorphism in puberty onset and reproductive strategies of males. In X. nigrensis and X. multilineatus, puberty onset is genetically determined and linked to a melanocortin 4 receptor (Mc4r) polymorphism of wild-type and mutant alleles on the sex chromosomes. We hypothesized that Mc4r mutant alleles act on wild-type alleles by a dominant negative effect through receptor dimerization, leading to differential intracellular signaling and effector gene activation. Depending on signaling strength, the onset of puberty either occurs early or is delayed. Here, we show by Förster Resonance Energy Transfer (FRET) that wild-type Xiphophorus Mc4r monomers can form homodimers, but also heterodimers with mutant receptors resulting in compromised signaling which explains the reduced Mc4r signaling in large males. Thus, hetero- vs. homo- dimerization seems to be the key molecular mechanism for the polymorphism in puberty onset and body size in male fish.

Reproductive endocrinology, morphological traits, and sexual selection in a population of wild South African giraffes

Sex steroids are pervasive in mammals and evolutionarily conserved, but differences in the nuances of endocrine profiles characterize distinct species. Two sex steroids, testosterone and progesterone, feature prominently in the life history of mammalian taxa, but neither one has been analyzed from wild giraffes. Our study was designed to address this gap in knowledge by examining how these sex steroids are related to biological features and giraffe life history. We conducted the research at Rooipoort Nature Reserve a 44,000 ha private nature reserve in South Africa on a population of South African giraffes (Giraffa camelopardalis giraffa). Eleven adult giraffe cows and seven adult bulls were immobilized and various biological samples and morphological measurements were obtained. We analyzed both testosterone and progesterone using Ultra High Performance Liquid Chromotography – Tandem Mass Spectrometry (UHPLC-MS/MS). We found that non-pregnant females had lower progesterone concentrations than pregnant females and that those in the last trimester of pregnancy showed a slight drop in progesterone, as well as an increase in testosterone. Among males, chronological age was correlated with testicle size, testosterone concentrations, and ossicone volume. We propose that the progesterone decline functions partly to accelerate resumption of ovulation because giraffes become pregnant while lactating, and that the testosterone elevation provides an endocrine mileu for female defense of neonatal calves, given that lion predation is a major threat to calf survival. We suggest that male reproductive strategies are mediated by the age-related impacts of testosterone on growth in both body mass and ossicone volume as a consequence of sexual selection. We conclude by noting that the robust and solid ossicones of male giraffes function in mate competition as wedges that are used to topple opponents by raising their legs and placing them in an off-balanced position that can seriously injure them when falling to the ground.

Does good food make male mice sexy? The influence of diet on male sexually selected traits

Diet may have an important influence on life history and behavioral traits involved in sexual selection. Males of high condition should be better able to bear the costs of high trait expression and hence express more elaborate sexual signals. Here, we follow Mus musculus domesticus raised in semi-natural conditions across four generations of standard (SQ) versus high quality (HQ) food, containing a higher energy level and manufactured to boost body condition during reproduction. We investigate multivariate effects on life history, pre- and post-copulatory traits of males. In addition, we investigate the effects of a food switch in the third generation on trait expression. From the F3 generation on, HQ males were more dominant. This resulted in HQ males being preferred in mate choice even though no post-copulatory traits were affected, suggesting a condition-dependent allocation to different traits. Sons of food switched males showed reduced pre- but not post-copulatory trait expression, irrespective of the direction of the food switch, indicating a mismatch rather than adaptive plastic adjustments in the first generation. We conclude that males balance investment into different traits based on the diet to achieve similar fitness. Furthermore, we show that adaptive shifts in male reproductive strategies need several generations to emerge while first responses to changing environments reflect a constrained trait expression.

Male reproductive strategies are influenced by the maternal sociosexual environment

Male reproductive strategies are influenced by the maternal sociosexual environment

The Sense in Senseless Violence: Male Reproductive Strategy and the Modern Sexual Marketplace as Contributors to Violent Extremism

The Sense in Senseless Violence: Male Reproductive Strategy and the Modern Sexual Marketplace as Contributors to Violent Extremism

The use of song playback experiments to study the social mechanisms of bird settlement decisions

Acoustic playback techniques involve broadcasting of various sound stimuli to simulate the presence of a certain species. In this article, we applied a variation of the method using automatic sound-reproducing stations (ASRS), which imitate the daily vocal activity of birds that is close to natural. We investigated whether acoustic playback experiments help to examine the role of social cues for bird settlement decisions, as well as to detect the presence of rare and low-density species. Using this method, we were able to increase the local density of the rustic bunting Ocyris rusticus Pallas, 1776 (Emberizidae) – species, which is currently listed in the Red Book of the Russian Federation (2001). The findings of our study show that rustic buntings preferred to settle on plots where the song imitating the presence of conspecifics was broadcasted by ASRSs. In addition, the playback method was used to examine the social structure of the population of the wood warbler Phylloscopus sibilatrix Bechstein, 1793 (Sylviidae), a small species that lives here on the northern periphery of the range. The wood warbler songs were broadcasted by ASRSs before the arrivals of the first conspecifics to the breeding area. Songs with different acoustic activity (2 or 6 trills per min) were used to create a difference between the social environments on plots. The gathered data showed a display of two different behavioral responses of individuals (males) to different song types broadcasted on plots. Based on this observation, the main hypothesis is about the existence of both conservative and opportunistic male reproductive strategies of the wood warbler. The results demonstrate the possibility of using acoustic playback techniques to study the role of acoustic signals in territorial behavior and social structure of a bird population. Understanding the ways birds make settlement decisions is essential for the development of conservation practices.

Drivers and consequences of female reproductive competition in an egalitarian, sexually monomorphic primate

Even after the 150th anniversary of sexual selection theory, the drivers and mechanisms of female sexual selection remain poorly studied. To understand demographic circumstances favoring female-female competition, trade-offs with kin selection and interactions with male reproductive strategies, we investigated female evictions in redfronted lemurs (Eulemur rufifrons). Based on 24 years of demographic data of known individuals, we show that female redfronted lemurs target close female kin for forcible, permanent, and presumably lethal eviction, even though groups contain multiple unrelated males whose voluntary emigration actually mitigated the probability of future female evictions. Female eviction and male emigration were predicted by group size, but male emigration was primarily driven by a proportional increase of male rivals. Female evictions were more likely than male emigrations when there were more juvenile females in a group, but the identity of evicted females was not predicted by any intrinsic traits. While birth rates were reduced by the number of juvenile females, they were higher when there were more adult females in a group and in years with more rainfall. Early infant survival was reduced with increasing numbers of juvenile females, but variation in female lifetime reproductive success was not related to any of the predictors examined here. Thus, there seems to be a limit on female group size in this lemur species. More generally, our study demonstrates a balanced interplay between female reproductive competition, competition over group membership between both sexes, and kin selection, contributing new insights into the causes and consequences of female competition in animal societies.Significance statementThe evolutionary causes of female competition in vertebrate societies remain poorly known. Evictions represent an extreme form of female competition because even close kin are evicted when same-sized unrelated males are theoretically also available as victims. We studied drivers and consequences of evictions in redfronted lemurs (Eulemur rufifrons) using 24 years of demographic data from multiple groups. We show that while voluntary male emigration mitigates the probability of future female evictions, females nonetheless appear to accept the fitness costs of evicting female kin. While group size seems to be the main driver of departures by either sex, the number of juvenile females present in groups is the key variable triggering eviction events as well as physiological responses that could be interpreted as female reproductive restraint. Our study therefore revealed that competition does trump cooperation under some circumstances in the intricate interplay between sexual selection and kin selection on females.

Genomic basis of Y-linked dwarfism in cichlids pursuing alternative reproductive tactics.

Sexually antagonistic selection, which favours different optima in males and females, is predicted to play an important role in the evolution of sex chromosomes. Body size is a sexually antagonistic trait in the shell-brooding cichlid fish Lamprologous callipterus, as "bourgeois" males must be large enough to carry empty snail shells to build nests whereas females must be small enough to fit into shells for breeding. In this species, there is also a second male morph: smaller "dwarf" males employ an alternative reproductive strategy by wriggling past spawning females into shells to fertilize eggs. L.callipterus male morphology is passed strictly from father to son, suggesting Y-linkage. However, sex chromosomes had not been previously identified in this species, and the genomic basis of size dimorphism was unknown. Here we used whole-genome sequencing to identify a 2.4-Mb sex-linked region on scaffold_23 with reduced coverage and single nucleotide polymorphism density in both male morphs compared to females. Within this sex region, distinct Y-haplotypes delineate the two male morphs, and candidate genes for body size (GHRHR, a known dwarfism gene) and sex determination (ADCYAP1R1) are in high linkage disequilibrium. Because differences in body size between females and males are under strong selection in L.callipterus, we hypothesize that sexual antagonism over body size initiated early events in sex chromosome evolution, followed by Y divergence to give rise to bourgeois and dwarf male reproductive strategies. Our results are consistent with the hypothesis that sexually antagonistic traits should be linked to young sex chromosomes.

Sandy beach dwelling crickets (Orthoptera: Trigonidiidae) have no preference for substrates that match their body color

ABSTRACTHabitat choice is an important behavior for maximizing fitness. One of the factors involved in habitat choice is preference for habitat substrates. In the context of predatory avoidance, it is adaptive to prefer substrates that are matched to own body color. The ground cricket Dianemobius csikii (Bolívar, 1901) occurs on sandy beaches. The body color of D. csikii resembles the sand of their habitat, which could be a form of crypsis. If D. csikii is not able to choose a suitable habitat background that matches their body color, the crickets incur a high risk of predation. The risk could result in population decline of the cricket. To examine whether D. csikii shows preference for substrates that match its body color, we undertook a two‐choice test of sand or soil substrates after controlling the type of flooring substrates used during rearing. The crickets did not show preference for sand. There was no significant effect of flooring substrates during the nymph stage on the preference. These results suggest that when habitats are fragmented, crickets do not actively choose a cryptic background. There were significant differences in the substrate preference between sexes. These results reflect differences in the reproductive strategies of males and females.

Diverse grouping and mating strategies in the Critically Endangered Hainan gibbon (Nomascus hainanus)

Understanding flexibility in the social structure and mating strategies of the world’s last remaining population (35 individuals) of wild Hainan gibbons (Nomascus hainanus) is critical for developing effective management plans to aid in their population recovery. Three of the five remaining Hainan gibbon groups (A, B, and C) currently live in a social unit characterized by two or three adult males, two reproducing adult females, and offspring. A fourth group (D) contains one adult male, two adult females, and offspring, and Group E contains a single adult male–adult female pair with a young infant. In this study, we describe observations of copulations between multiple resident males and one of the two resident females in Group C. Group C is best described as a small multi-male/multi-female group. We found that this breeding female (F2) solicited copulations from two resident adult males (M1 and M2) on the same day, and also mated with each of these two males on different days. Resident males were not observed to interrupt the mating pair. Although factors such as a biased adult sex ratio, severe population disruption, and habitat degradation can help explain variation in group composition and mating strategies in Hainan gibbons, we argue that there exists considerable mating system variability across gibbon species, and that this variability offers important insights into male and female Hainan gibbon group structure and reproductive strategies.

Immigrant males’ knowledge influences baboon troop movements to reduce home range overlap and mating competition

Abstract Mechanistic models suggest that individuals’ memories could shape home range patterns and dynamics, and how neighbors share space. In social species, such dynamics of home range overlap may be affected by the pre-dispersal memories of immigrants. We tested this “immigrant knowledge hypothesis” in a wild population of chacma baboons (Papio ursinus). We predicted that overlap dynamics with a given neighboring troop’s home range should reflect males’ adaptive interests in overlap when the alpha male had immigrated from this neighboring troop but less so when the alpha male originated from elsewhere. We used data collected between 2005 and 2013 on two neighboring troops in Namibia, comprising GPS records of daily ranges, male natal origins, daily females’ reproductive status, and a satellite index of vegetation growth. We found support for our prediction in line with male reproductive strategies but not in line with foraging conditions. In periods with a higher relative number of fertile females over adult males in the focal troop, male baboons would benefit from reducing overlap with their neighbors to mitigate the costs of between-troop mating competition. This was indeed observed but only when the alpha male of the focal troop was an immigrant from that neighboring troop, and not with alpha males of other origins, presumably due to their different knowledge of the neighboring troop. Our findings highlight the role of reproductive competition in the range dynamics of social groups, and suggest that spatial segregation between groups could increase through the combination of dispersal and memory.

The context of sexual coercion in orang-utans: when do male and female mating interests collide?

Sexual coercion is widespread in the animal kingdom. Its direct forms, including harassment and forced copulation, have largely been investigated as an expression of (alternative) male reproductive strategies, rather than the result of a sexual conflict between the sexes. Likewise, the frequent occurrence of forced copulations in orang-utans (Pongo spp.) has been attributed to male strategies and more recently also to concealed female fecundity. So far, however, the immediate contextual variables leading to forced copulations have rarely been examined. We compared two orang-utan populations, Suaq (Pongo abelii, Sumatra) and Tuanan (Pongo pygmaeus, Borneo), both characterized by an individual-based fission–fusion lifestyle, whereas their socioecology differs. We assessed how the occurrence of female-resisted and voluntary copulations was affected by female reproductive state, male morph (unflanged or flanged), measures of male–male competition, male–female relationship and ecological factors. Besides female reproductive state and male morph, predictors of female resistance were related to male–male competition. First, female resistance was more likely towards subordinate males who were displaced from proximity to the female by another male during that association. Second, the presence of additional flanged males increased the probability of female resistance. Third, the latency to both the arrival of another male and to the end of the association after sexual interactions was shorter if there was female resistance. We conclude that sexual coercion in orang-utans is highly dependent on the vicinity of more dominant males and can only be understood in the light of sexual conflict: While males force copulations when at risk of losing access to a female and thus follow a ‘now-or-never’ strategy, female resistance follows a ‘not-you-now’ pattern, which is ultimately consistent with an infanticide avoidance strategy.

Costs of male infanticide for female capuchins: When does an adaptive male reproductive strategy become costly for females and detrimental to population viability?

Infanticide in white-faced capuchins (Cebus capucinus imitator) typically occurs in association with alpha male replacements (AMRs). Although infanticide is likely adaptive for males, it imposes costs on females that are difficult to quantify without long-term demographic data. Here we investigate effects of AMRs and infanticide on female reproductive success and how these costs affect capuchin groups. We investigate (1) effects of AMR frequency on the production of surviving infants; (2) energetic and (3) temporal "opportunity costs" of infant loss; and (4) how AMR frequency impacts capuchin group sizes. We censused six groups (7-33 years/group, 74 adult females). We modeled surviving infant production in relation to AMR. We estimated a female's energy requirements for lost infants and the temporal cost relative to the median reproductive window. We simulated how varying AMR rates would affect future capuchin group sizes. Females exposed to more frequent AMR tended to produce fewer surviving offspring. We estimate the average lost infant requires approximately 33% additional energy intake for its mother and represents 10% of the average reproductive opportunity window available to females. Simulated populations remain viable at the observed rate of AMR occurrence but decrease in size at even slightly higher rates. While infanticide is adaptive for males, for females it affects lifetime reproductive success and imposes energetic and opportunity costs. Although capuchin populations have evolved with AMRs and infanticide, small increases in AMR frequency may lead to population decline/extinction. Infanticide likely plays a large role in population maintenance for capuchins.

No risk to scrambling? Mating tactic does not affect the frequency of leg autotomy in a New Zealand harvestman

Intense sexual selection on males may drive the evolution of exaggerated weaponry, typically used in contests for females or reproductive sites. In some species, males have discontinuous variation in weapon morphology that is accompanied by alternative reproductive tactics (ARTs). Major males with enlarged weapons usually exhibit a mating tactic based on female or resource defence, which makes them relatively sedentary. Minor males with reduced or absent weapons may exhibit a scramble competition mating tactic, which involves increased movement in search of females. Thus, the main costs paid by majors involve the expression/maintenance of exaggerated structures and potential injuries during contests. In turn, the main costs paid by minors are likely related to increased exposure to predation during mate search, yet this cost has rarely been considered for minors. Harvestmen are an arachnid group with diverse male weaponry, including many cases of male polymorphism associated with ARTs. Species of the suborder Eupnoi use leg autotomy as a common antipredator defence. Here we investigated whether leg autotomy (a proxy for predation risk) is more frequent in minors of the weapon polymorphic harvestman Forsteropsalis pureora , which has three male morphs with drastically different cheliceral size and morphology. Leg autotomy was very common, with 54% of wild-caught individuals missing at least one leg (mostly leg II), but we found no evidence for differential predation risk between male morphs during adulthood. In a predator simulation experiment, we found no difference in the likelihood or latency to autotomize a leg between male morphs or leg types (legs I, II, III or IV). However, males already missing legs were more reluctant to autotomize an additional leg. Our results suggest that while leg autotomy is a common antipredator strategy, with no difference between male tactics, costs are compounded as additional legs are autotomized and males may strategically decide not to autotomize in future predator encounters. • New Zealand harvestmen autotomize (lose) legs to escape predators. • This defence strategy is common in wild populations. • Sensory legs are lost more often than locomotory legs. • Harvestmen that have lost legs in the past take longer to drop an additional leg. • Male reproductive strategy does not affect autotomy risk.

Y chromosome evolution spurs behavioural diversity in male fish.

The Y chromosome of the freshwater fish Poecilia parae may have successively evolved five haplotypes that are maintained in the population for alternative male reproductive strategies.

Male reproductive strategies in two species of spider crabs, Leurocyclus tuberculosus and Libinia spinosa

Reproductive strategies vary according to season length, individual reproductive traits, and factors associated with the social context such as density and sex ratio of the individuals. The social context predicts the intensity of sexual selection due to its effect on mate choice and intra-sexual competition. Moreover, it exacerbates the costs derived from sexual conflict and allows for the existence of alternative reproductive tactics to counteract such costs. However, these mechanisms are still underexplored in many animal taxa. Majoidea “spider crabs” show a diversity of life history, and behavioral and morphological adaptations for reproduction, which have evolved as an outcome of competition to maximize reproductive success. We aimed to analyze in this study, male reproductive strategies through mating behavior under different social contexts, density of individuals, and males size differences in two species of “spider crabs”, Leurocyclus tuberculosus and Libinia spinosa. The experiments were as follows: one female and one male per aquarium (Experiment 1); a female and three males of similar sizes per aquarium (Experiment 2); and a female and three males of distinct sizes per aquarium (Experiment 3). The experiments showed that mating behavior and duration of mate guarding differ between species and among different social contexts. The presence of males with similar or different sizes showed a relevant different behavior, the female copulated with more than one male when males were similar in size (E2), and copulated only with the largest male when males were different in size (E3). In mate guarding, the shape and the use of the chelipeds had an important role: the male of Leurocyclus tuberculosus grasped the female with the chelipeds and did not fight with other males, but the male of Libinia spinosa surrounded the female with the chelipeds and fought with other males. The longest duration of mate guarding was during precopulatory guarding. Precopulatory guarding was present in all the experiments and its duration did not present significant differences among the experiments. However, in both species, the experiments differ in the duration of postcopulatory guarding until spawning. Gonopod morphology revealed important differences between species and are probably associated with sperm competition. The experiments under distinct social contexts linked to the use and shape of the chelipeds and the morphology of the gonopods, could allow to understand the functional significance of the different reproductive strategies.

Intersexual differences in density-dependent dispersal and their evolutionary drivers.

Dispersal is well recognized as a major driver of evolutionary processes in local populations. Nevertheless, dispersal abilities should also be perceived as a life-history trait, being subject to evolutionary changes in response to various drivers. Empirical studies investigating these drivers rarely consider that they may influence male and female dispersal differently. The purpose of our study was to document intersexual differences in density-dependent emigration from local habitat patches. As a model system, we used a metapopulation of Maculinea (Phengaris) teleius butterfly, in which densities of both sexes vary greatly throughout the flying season. Following intensive mark-release-recapture surveys, the parameters and predictors of dispersal were analysed with the Virtual Migration model and the multi-state recapture model. The emigration rate in males was substantially higher in the early season, especially at smaller habitat patches. With the proportion of females increasing with the season progression, males became reluctant to emigrate from their natal patches. In turn, higher female emigration in the later part of the season was most strongly associated with female tendency to reduce intraspecific competition experienced by their offspring. Our findings provide evidence for the impact of reproductive strategies on dispersal in both sexes. The difference in reproductive strategies of males and females explains sex-biased dispersal in different parts of the season, which carries important implications for metapopulation functioning.