Scramble Competition Polygyny Research Articles

Ecology and social behavior of the tamarisk gerbil Meriones tamariscinus: insights from long-term research in the wild and semi-natural environments

Abstract The present review provides a compilation of the published data on the ecology and social behavior of tamarisk gerbils. Both field studies and direct observations under semi-natural conditions provide evidence that the tamarisk gerbil is a nocturnal herbivorous rodent that lives in highly seasonal habitats and displays seasonal fluctuations in reproduction and spatial organization. A typical feature of the tamarisk gerbils’ spatial organization is higher mobility of males during the breeding season (as compared with the nonbreeding period) and formation of temporary aggregations of males competing for access to receptive females; the composition of these aggregations was variable and depended on the reproductive condition of the females. Females tend to occupy exclusive home ranges irrespective of their reproductive condition. The mating system of the species can be defined as scramble competition polygyny with some features of polygynandry and promiscuity. The tamarisk gerbil has distinct features of a solitary species and its social structure is primarily based on aggressive interactions or mutual avoidance of conspecifics resulting in a dominance hierarchy among males and site-dependent dominance among females during the breeding season. By the end of the breeding season, males become less mobile and occupy nearly exclusive home ranges, consistent with solitary living. The main features of the spatial and social organization of this species, which distinguish it from other solitary rodents, are the higher mobility of males and the formation of temporary multimale–multifemale aggregations during the breeding season. Overall, the data presented expand our understanding of socioecology of gerbils.

Multi-modal locomotor costs favor smaller males in a sexually dimorphic leaf-mimicking insect

BackgroundIn most arthropods, adult females are larger than males, and male competition is a race to quickly locate and mate with scattered females (scramble competition polygyny). Variation in body size among males may confer advantages that depend on context. Smaller males may be favored due to more efficient locomotion leading to higher mobility during mate searching. Alternatively, larger males may benefit from increased speed and higher survivorship. While the relationship between male body size and mobility has been investigated in several systems, how different aspects of male body morphology specifically affect their locomotor performance in different contexts is often unclear.ResultsUsing a combination of empirical measures of flight performance and modelling of body aerodynamics, we show that large body size impairs flight performance in male leaf insects (Phyllium philippinicum), a species where relatively small and skinny males fly through the canopy in search of large sedentary females. Smaller males were more agile in the air and ascended more rapidly during flight. Our models further predicted that variation in body shape would affect body lift and drag but suggested that flight costs may not explain the evolution of strong sexual dimorphism in body shape in this species. Finally, empirical measurements of substrate adhesion and subsequent modelling of landing impact forces suggested that smaller males had a lower risk of detaching from the substrates on which they walk and land.ConclusionsBy showing that male body size impairs their flight and substrate adhesion performance, we provide support to the hypothesis that smaller scrambling males benefit from an increased locomotor performance and shed light on the evolution of sexual dimorphism in scramble competition mating systems.

Biparental mealybugs may be more promiscuous than we thought.

Knowledge on the reproductive biology of target insect pest is essential for the effective implementation of pheromone-based pest management tactics. In mealybugs, the second largest family of scale insects, the existence of female multiple mating was recently suggested. In this study, we aimed at testing how general is this behavior in mealybugs, by investigating polygyny and polyandry in two cosmopolitan pest mealybugs, Planococcus citri and Pseudococcus calceolariae. Males of these species were able to mate an average of 11.9 and 13.3 females, respectively, during their lifespan. The number of fertilized females per male decreased with male age/mating history for both mealybugs. We found no differences in female fecundity and fertility, when fertilized by males with different mating history. When we used male age as a proxy of mating history, we observed a significant negative effect on female fecundity. The females of both species remained receptive after first copula and eventually mated multiple times. The percentage of remated females of P. citri decreased linearly with time since first copula, with some maintaining receptivity up to 24 h. Males showed no preference between virgin and mated females, in static-air olfactometer tests. We found no benefit of female multiple mating in relation to fecundity. In biparental mealybugs, the mating system of males is possibly scramble competition polygyny; and that of females is possibly polyandry, with female receptivity restricted to a relatively short period. We discuss the practical implications of the results for pest management.

Reproductive effort and success of males in scramble-competition polygyny: Evidence for trade-offs between foraging and mate search.

Patterns of male reproductive allocation provide insight into life-history characteristics. The trade-offs associated with resource and female group defence are well-defined. However, less is understood about trade-offs in species that practise scramble-competition polygyny, where successful strategies may favour competitive mate-searching rather than contest competition and fighting. White-tailed deer (Odocoileus virginianus) practise scramble-competition polygyny where solitary males search for and assess receptivity of females scattered across the landscape. Physically mature males are expected to do most of the breeding because of the high energetic costs of reproduction and high social status. However, young males may collectively sire one-third of offspring. To gain a better understanding of trade-offs associated with scramble-competition polygyny, we quantified metrics associated with reproductive effort and success. We quantified changes in body mass of harvested males, energetic costs of locomotion based on movements of GPS radiocollared males and timing of reproduction via temporal genetic parentage assignments. Young males (1.5 and 2.5years old) sired offspring, but their mating success was mainly limited to peak rut, when most females were in oestrus. Furthermore, multiple paternity was common, indicating opportunistic reproduction. Reproductive effort, indexed by body mass loss, was highest in prime-age males (5.5-6.5years old). Surprisingly, young and postprime males also exhibited significant body mass loss, indicative of investment in reproductive effort. Movement rates increased twofold to fourfold during rut as a function of mate search activities, but cost of locomotion would cause only about one-third of observed body mass loss. Because males are capital breeders, we infer most of body mass loss is due to reduced foraging. In scramble-competition polygyny, the repeated location of potential mates and assessment of their oestrous status appear to be important constituents of male mating strategies. Therefore, mating success may be influenced by time management and spatial memory, and not based solely on social dominance. Thus, reproductive effort should be greater for individuals capable of reducing time foraging. For those that cannot, opportunistic mating opportunities may arise when operative adult sex ratios are low. Our analyses reveal valuable insight into the trade-offs associated with scramble-competition polygyny.

Male Body Size Predicts Reproductive Success But Not Within-Clutch Paternity Patterns in Gopher Tortoises (Gopherus polyphemus).

In many vertebrates, body size is an important driver of variation in male reproductive success. Larger, more fit individuals are more likely to dominate mating opportunities, skewing siring success and resulting in lower effective population sizes and genetic diversity. The mating system of the gopher tortoise (Gopherus polyphemus) has been characterized as both female-defense and scramble-competition polygyny. Mating systems are typically not fixed and can be influenced by factors such as population density, demographic structure, and environmental conditions; however, most populations will have a predominant strategy that results from local conditions. We assessed how male body size influences patterns of paternity and reproductive success in a natural population of gopher tortoises in Florida, United States. Using microsatellites, we assigned parentage of 220 hatchlings from 31 nests collected during 2 reproductive seasons. Larger males were significantly more likely to sire offspring and sired more offspring than smaller males; however, the likelihood of a clutch being multiply sired was unrelated to male body size. We also found evidence of mate fidelity across years. Although paternity patterns in this high-density population are more consistent with defense polygyny, female monopoly by males was incomplete, with both large and small males contributing to multiply sired clutches. Additional behavioral data are needed to clarify the role of female mate selection in paternity outcomes. The context-dependence of mating systems underscores the need to compare parentage patterns across populations and to recognize the potential for more than 1 strategy to be employed within a single population.

Males, but not females, perform strategic mate searching movements between host plants in a leaf beetle with scramble competition polygyny.

Mate searching is assumed to be performed mostly by males, but when females benefit from multiple mating or are under risk of failing to mate, they may also perform mate searching. This is especially important in scramble competition polygynies, in which mate searching is the main mechanism of mate competition. Typically, more mobile individuals are expected to achieve higher mating success because mobility increases their probability of finding mates. If we assume individual movements are mainly explained by mate searching in scramble competition polygynies, we can investigate searching strategies by asking when individuals should leave their location and where they should go. We hypothesize that individuals will leave their locations when mating opportunities are scarce and will seek spatially close sites with better mating opportunities. We tested these hypotheses for males and females of Leptinotarsa undecimlineata, a leaf beetle with scramble competition polygyny in which both sexes are promiscuous. Individuals mate and feed exclusively on Solanum plants, and thus, individual movements can be described as switches between plants. Females were less likely than males to leave isolated plants, and both males and females moved preferentially to neighboring plants. Males were more likely to leave when the local number of females was low, and the number of males was high. They moved to plants with more females, a behavior consistent with a mate searching strategy. Females were more likely to move to plants with fewer males and many females, a behavior consistent with male harassment avoidance. Strategic movement is widely considered in foraging context, but seldom in a mate searching context. Considering that selection to minimize searching costs, maximize mating success, and minimize harassment may be ubiquitous in nature, we argue that strategic movements by mate searching individuals are likely to occur in many species.

Paternity and male mating strategies of a ground squirrel ( Ictidomys parvidens ) with an extended mating season

Abstract Animal mating systems are driven by the temporal and spatial distribution of sexually receptive females. In mammals, ground-dwelling squirrels represent an ideal clade for testing predictions regarding the effects of these parameters on male reproductive strategies. While the majority of ground squirrel species have a short, highly synchronous annual breeding season that occurs immediately after females emerge from hibernation, the Mexican or Rio Grande ground squirrel ( Ictidomys parvidens ) differs markedly in having an extended mating season (2 months) and a long delay between emergence from hibernation and female receptivity (1–2 months). Both traits are expected to favor polygyny by increasing the chances that a male can secure matings with multiple females (e.g., females that come into estrus on different days). To test this prediction, we used microsatellite markers to characterize the mating system of a population of Rio Grande ground squirrels from Carlsbad, New Mexico. Our analyses indicated a high frequency of multiple paternity of litters in this population. Paternity was not related to spatial overlap between known mothers and assigned fathers, suggesting that territory defense is unlikely to be an effective male reproductive strategy in the study population. Dominance interactions among males were frequent, with heavier males typically winning dyadic interactions. Surprisingly, however, males with lower dominance scores appeared to have higher reproductive success, as did males that were active over a greater extent of the study site. Collectively, these results suggest that the mating system of the Rio Grande ground squirrel is best described as scramble competition polygyny, with the primary male reproductive strategy consisting of searching for estrous females. Similar patterns of male–male competition have been reported for a few other ground squirrel species, providing potentially important opportunities for comparative studies of the factors favoring this form of male reproductive strategy.

Mobility and mating frequency in the scramble competition polygyny of a chrysomelid beetle

In scramble competition polygyny, male and female mobility may be under strong selection as a result of fitness effects of searching for reproductive resources such as mates, oviposition sites, or resources for egg production. We analyzed the relationship between mating frequency, mobility, and body size in males and females of the chrysomelid beetle Leptinotarsa undecimlineata. We obtained a detailed data set of movement and mating frequency of an entire population (1037 adults) over a full reproductive season using individual tagging and direct behavioral observations. Unlike previous studies, we found a negative relationship between mobility and mating success for both sexes. Size was positively correlated to mating frequency in females, but negatively in males. High male mobility may be the result, and not a cause, of low mating success in scramble mating polygynies where rejected or displaced males switch plants more often searching for mating opportunities. More mobile females may be looking for competition-free oviposition substrate and thus experiencing fewer sexual encounters.

Male mating tactics in spider monkeys: sneaking to compete

I investigated the mating system and male mating tactics for a population of wild spider monkeys (Ateles belzebuth chamek), to identify the behaviors males used to achieve and maintain access to sexually receptive females, and to examine if some males used more tactics than other males and/or had differential access to females. Results show that the mating system mostly involved scramble competition polygyny and that males used a range of mating tactics and behaviors, previously unreported for spider monkeys. The most unusual feature of spider monkey mating behavior was the secretive nature of copulations-nearly all copulations were clandestine, but a few were in the presence of other group members. Fifteen sexually mature males were observed to copulate 43 times. These data provide the first opportunity to evaluate how female availability influences male-male competition. First, the operational sex ratio was highly skewed toward males because usually only one female was receptive in each community per month. Second, females only mated with a few males in their community in any one mating period, but some females mated over the course of multiple consecutive mating periods, eventually mating with most or all of the males in their community. Across all communities, 9 (21%) of the 43 copulations involved a single male-female partner, 20 (47%) involved four males mating with the same female, and males mated with from one to four different females. Fourteen of the 16 total adult males and 1 subadult male (10 total) copulated. One or two males in each community were successful in monopolizing access to receptive females, and these males did not usually have the highest rates of copulation. In this system, clandestine copulations are one behavioral solution to the complex problem of gaining mating exclusivity and, probably, exercising mate choice.

A paternity advantage for speedy males? Sperm precedence patterns and female re-mating frequencies in a sexually cannibalistic praying mantid

Scramble competition polygyny is expected when females and/or resources are widely dispersed and not easily monopolized by males, or when there is an abundance of mates during an extremely restricted reproductive period. Additional factors such as first male sperm precedence or low female re-mating rate might further explain the propensity of males to engage in scramble competition. The sexually cannibalistic praying mantid Pseudomantis albofimbriata exhibits a polygynous mating system, where females exist in low-density populations and male competition manifests as the race to find females rather than as direct physical fighting. Here, we aim to determine whether there is a paternity advantage for the first-male to mate and/or a low frequency of female re-mating. First, we determined sperm precedence patterns in P. albofimbriata using the sterile male technique. Second, we tested the likelihood of female re-mating in P. albofimbriata by comparing the close-range approach behaviour and frequency of successful mating attempts for males when paired with virgin as opposed to recently mated females, and by comparing the frequency of long-distance male attraction between virgin and mated females. We found no paternity advantage for the first male to mate, rather a second male advantage. Although mated females were not rejected by males when approached from close-range, they were chemically unattractive to males searching from a distance. Since initial mate attraction in many praying mantids, including P. albofimbriata, is mediated via long-distance chemical communication, we believe the latter result is more ecologically relevant and therefore more important. These results suggest that the relatively low frequency of female re-mating observed in P. albofimbriata may be an additional factor driving scramble competition in this system.

Copulation, mating system and sexual dimorphism in an Australian millipede, Cladethosoma clarum

Investigations of millipede mating dynamics have focussed primarily on Old World or North American species. In contrast, data for the diverse and abundant polydesmid millipedes of Australia are lacking. Here I examine copulation, behaviour and sexual dimorphism in an Australian millipede, Cladethosoma clarum. In this species, copula position is best described as ‘in parallel’. Both females and males mated with two or more individuals of the opposite sex, and average copulation duration was 4.7 min. Behavioural observations revealed differences between the sexes in patterns of behaviour. Specifically, females devoted most of their time to feeding, whereas males were more mobile and were observed walking more often than females. The sexes also exhibited marked sexual dimorphism in that females exhibited greater body width and body mass compared with males, whereas males had longer legs relative to females. In millipedes, longer legs allow an individual to walk faster. Consequently, male C. clarum appear to be capable of relatively rapid locomotion. As Australian millipede densities are spatially unpredictable, the increased mobility of males likely reflects an active mate-location strategy. Furthermore, the occurrence of multiple mating by females suggests that male leg length may have evolved as an adaptation for rapid mate location under scramble competition polygyny.

Patterns of Dispersion and Burrow Use Support Scramble Competition Polygyny in Gopherus polyphemus

Gopher tortoises spend most of their time in burrows from which they emerge to forage and perform behaviors such as courtship and mating. Previous literature is divided regarding the mating system of this species; some assert that gopher tortoises conform to female defense polygyny, and others assert that scramble competition polygyny is more likely. Here, telemetry data were used to record the frequency with which pairs of tortoises shared burrows and the frequency with which they apparently chased each other from burrows. Additionally, telemetry locations were used to estimate patterns of dispersion of individuals. If gopher tortoises conformed to female defense polygyny, then males should have: (1) moved frequently to share burrows with females, (2) rarely shared burrows with males, (3) infrequently displaced females from burrows, and (4) frequently displaced males from burrows. Similarly, females should have: (1) infrequently moved to share burrows with either sex, and (2) infrequently chased either sex. Also, females should have shown an aggregated dispersion relative to other females. On the contrary, we found that males moved equally frequently to share burrows with adults of both sexes and chased females from burrows more frequently than they chased other males. Females moved more frequently to share burrows with males than with females and chased males more often than they chased other females. Females did not have an aggregated pattern of dispersion relative to other females. These data were most consistent with scramble competition polygyny.

Sex in the dark: determinants and consequences of mixed male mating tactics in Microcebus murinus, a small solitary nocturnal primate

In mammals with solitary females, the potential for males to monopolize matings is relatively low, and scramble competition polygyny is presumed to be the predominant mating system. However, combinations of male traits and mating tactics within this type of polygyny have been described. The main aim of our study was to identify the relative importance of, and interactions among, potential determinants of contrasting male reproductive tactics, and to determine their consequences for male reproductive success in a small solitary nocturnal Malagasy primate, the gray mouse lemur (Microcebus murinus). We studied their mating behavior over three consecutive annual mating seasons. In addition, we determined the genetic relationships among more than 300 study animals to quantify the reproductive success of individual males. We found that, with a given relatively low overall monopolization potential, successful male mouse lemurs roamed extensively in search of mates, had superior finding ability and mated as early as possible. However, contest competition was important too, as temporary monopolization was also possible. Males exhibited different mating tactics, and heavier males had a higher reproductive success, although most litters had mixed paternities. Switching between tactics depended on short-term local variation in monopolization potential determined by a pronounced dynamic in fertilization probability, number of alternative mating opportunities, and the operational sex ratio. This study also revealed that the dynamics of these determinants, as well as the mutual interactions between them, necessitate a detailed knowledge of the mating behavior of a species to infer the impact of determinants of alternative mating tactics.

Mating Opportunities of Female Gopher Tortoises, Gopherus polyphemus, in Relation to Spatial Isolation of Females and Their Burrows

Gopher Tortoises (Gopherus polyphemus) are thought to exhibit harem defense polygyny because adult females that are relatively aggregated are thought to receive increased rates of mating visits by large, dominant males relative to females that are more isolated. To examine this expected pattern, we regressed rates of social behaviors on (1) degree of isolation of burrows occupied by female tortoises and (2) degree of isolation of female tortoises. The rate at which other tortoises visited a female was negatively associated with degree of isolation of burrows and females. However, we found no relationship between rates of behaviors associated with reproduction (interactions, courts, and mounts) and degree of isolation of burrows or females. Additionally, there was no association between degree of isolation of females and the number or body mass of males that mounted them. These results indicate that, for large, contiguous populations of Gopher Tortoises in high-quality habitat, such as our study site, distance from neighboring tortoises does not affect mating opportunities of reproductive females. Our findings suggest that patterns of reproduction of Gopher Tortoises conform more closely to expectations of scramble-competition polygyny than to harem defense polygyny.

The hidden matrilineal structure of a solitary lemur: implications for primate social evolution.

Kin selection affects many aspects of social behaviour, especially in gregarious animals in which relatives are permanently associated. In most group-living primates with complex social behaviour, females are philopatric and organized into matrilines. Models of primate social evolution assume that females in solitary primates are also organized into matrilines. We examined the genetic structure and the mating system of a population of Coquerel's dwarf lemur (Mirza coquereli), a solitary primate from Madagascar, to test this assumption. Our genetic and behavioural analyses revealed that this population of solitary individuals is indeed structured into matrilines, even though this pattern was not predicted by behavioural data. Specifically, females sharing a mitochondrial DNA haplotype were significantly clustered in space and the average genetic and geographical distances among them were negatively correlated. Not all females were philopatric, but there is no evidence for the successful settlement of dispersing females. Although not all adult males dispersed from their natal range, they were not significantly clustered in space and all of them roamed widely in search of oestrous females. As a result, paternity was widely spread among males and mixed paternities existed, indicating that scramble competition polygyny is the mating system of this species. Our data therefore revealed facultative dispersal in both sexes with a strong bias towards female philopatry in this primitive primate. We further conclude that complex kinship structures also exist in non-gregarious species, where their consequences for social behaviour are not obvious.

Mating success in the spittlebug Cercopis sanguinolenta (Scopoli, 1763) (Homoptera, Cercopidae): the role of body size and mobility

In insects, a sexual size dimorphism commonly occurs, with larger females. However, as a deviation from this general rule, larger males are found in some species. In these species often sexual selection for large males has been presumed. The spittlebug Cercopissanguinolenta exhibits a distinct sexual size dimorphism with larger males. Mating behaviour was studied in a field population in respect to mating success of males and females. The aim of this study was to examine the mechanisms that lead to the observed non-random mating pattern. The results showed a mating pattern without size-assortative mating. A correlation was found between mating success and body size in males. In females no such correlation was found. The mobility of males depends on their body size and mobility is high only when females are present. However, in an analysis of covariance it was found that male mating success is not correlated with mobility, when controlled for body size. The mating system of the spittlebug was classified as scramble competition polygyny.

The social organization and breeding system of Brants' whistling rat (Parotomys brantsii)

Brants' whistling rat Parotomys brantsii is a gregarious rodent that undergoes marked population fluctuations every few years. Previous studies have variously described them as solitary or social. In this field study, based on direct observations, the social structure and breeding system of a population that almost quadrupled over 7 months was investigated. At the onset of the breeding season, the animals were solitary, each occupying a separate warren. By the end of the breeding season, there was multiple occupancy of some warrens. Each individual maintained a discrete territory within the warren, and exhibited no apparent social behaviour. Outside the breeding season the behaviour of males and females did not differ and both remained at their resident warrens. During the breeding season male P. brantsii visited more neighbouring warrens than did the females, and their visits were to unoccupied warrens or to those occupied by females. Females tended to remain within their own warren. Both sexes avoided warrens occupied by males and few male–male interactions were observed. Females were solicited at their warrens by males, who were frequently chased away. There was, however, evidence that reproductively receptive females were more tolerant of males, who may guard females following mating. This breeding system, based on competitive mate searching by males, and not overt aggression between males, parallels the scramble competition polygyny described for a number of other species, including North American ground squirrels.

The social organization and breeding system of Brants' whistling rat (Parotomys brantsii)

AbstractBrants' whistling rat Parotomys brantsii is a gregarious rodent that undergoes marked population fluctuations every few years. Previous studies have variously described them as solitary or social. In this field study, based on direct observations, the social structure and breeding system of a population that almost quadrupled over 7 months was investigated. At the onset of the breeding season, the animals were solitary, each occupying a separate warren. By the end of the breeding season, there was multiple occupancy of some warrens. Each individual maintained a discrete territory within the warren, and exhibited no apparent social behaviour. Outside the breeding season the behaviour of males and females did not differ and both remained at their resident warrens. During the breeding season male P. brantsii visited more neighbouring warrens than did the females, and their visits were to unoccupied warrens or to those occupied by females. Females tended to remain within their own warren. Both sexes avoided warrens occupied by males and few male–male interactions were observed. Females were solicited at their warrens by males, who were frequently chased away. There was, however, evidence that reproductively receptive females were more tolerant of males, who may guard females following mating. This breeding system, based on competitive mate searching by males, and not overt aggression between males, parallels the scramble competition polygyny described for a number of other species, including North American ground squirrels.

The race for females : The mating system of the red mason bee, Osmia rufa (L.) (Hymenoptera: Megachilidae)

The ecology of a species strongly influences the strategies with which males and females maximize their lifetime reproductive success. When males and females do not invest equally in offspring, the sex with the higher parental investment becomes a rare resource for the other. The spatial and temporal distribution of the limiting sex forms the basis of the mating system. In nest-constructing Aculeata such as the red mason bee, Osmia rufa, females perform intensive brood care, whereas males do not invest in their offspring but instead compete for access to mates. Receptive females of this species are widely distributed and do not assemble at certain places. Therefore, territorial behavior is not an advantageous mating tactic for males, which instead search for females within individual home ranges usually centered around food plants. The unmarked or defended home ranges of different males may completely overlap. Competitive searching leads to a random distribution of matings among males that is largely independent of body size. The mating system of O. rufa can be described as scramble competition polygyny.

Parameters of Mating Effort and Success in Male European Ground Squirrels, Spermophilus citellus

AbstractBehavioural and physiological changes during the reproductive period were documented in male European ground squirrels including: initiated conflict, scent marking, locomotion and range size, as well as testis size, scrotal pigmentation, testosterone levels and changes in body mass. Spring emergence dates in adult males varied but preceded those of females. Gonadal maturation was complete in adult males at emergence. Most yearlings did not mature and emerged with females. After a quiescent premating phase, male range, range overlap, conflict rates and locomotion increased at female emergence. The synchrony of females, their small ranges and short latencies to ovulation produced a highly male‐biased operational sex ratio (OSR) and potential for scramble competition polygyny. In contrast to other ground squirrel species, each female apparently mated with only a single male. Scramble competition did appear to occur, in that males tried to contact as many females as often as possible before copulation. Females placed high temporal prerequisites on males. Costs of scrambling were documented by body mass loss during mating, which covaried with the number of ‘acquired’ females. Testicular regression occurred earlier, and fattening, moult and onset of next hibernation were delayed in successful males. The determinant factor in these costs was found to be mass loss intensity during the main mating period. Surprisingly, this factor was best related to feeding changes and not to locomotion or aggression. Time budget constraints perhaps imposed by the female, with compounding influences of exercise and stress effects of conflict, then negatively affected ensuing seasonal activities. The individual changes may adversely affect individual survivability and/or future reproductive success.