Spatial organization and mating system of Microtus townsendii

Summary Space use by individual Townsend's voles, Microtus townsendii, was investigated in spring and summer by means of radiotelemetry and intensive live trapping in undisturbed grasslands near Vancouver, British Columbia. Home ranges of males were larger than those of females; females had significantly larger ranges in spring than in summer. Most males and females maintained territories free of individuals of the same sex in spring. Male-female pairs had their exclusive territories closely overlapping each other. The 1:1 operational sex ratio and the spatial association of pairs of males and females suggest that the voles were monogamous in the spring of 1988 and that 50% of the males were monogamous in the spring of 1989. In summer, there was more intrasexual overlap between home ranges of males and females and female ranges were considerably smaller than those of males. Females were more philopatric than males and females thought to be members of the same family group lived adjacent to each other or had overlapping home ranges. Males overlapped with more than one female in summer, but most females still overlapped with only one male, which suggests that the mating system is polygynous in summer. Thirty-five percent of the philopatric females became pregnant for the first time when the male spatially associated with their mother in the spring was still alive and thus could potentially have mated with their fathers. Male and female territoriality in spring is the proximate mechanism for the limitation of breeding density by spacing behaviour.[/p] Offprint requests to: C.J. Krebs     

Male preference for large females and assortative mating for body size in the sand lizard (Lacerta agilis)

Summary Male sand lizards (Lacerta agilis) are polygynous and guard individual females for several hours to days after copulation. Even though the copulation itself only lasts 2–4 min, the total time that a male invests per female is considerably more and may constitute a substantial investment during a mating season. In such situations, when male copulation frequency is constrained, or when variation in female fecundity is high, mate choice by males may be adaptive. Large body size in female sand lizards is correlated with higher fecundity. In choice experiments performed in the laboratory, male sand lizards preferred to court large females rather than small females. In addition, when there was little difference in size between the females in the experiment, the males visited the two females more often before they started to court the preferred female. The results from a field study during 1984 and 1987–1990 showed that females are non-aggressive, have small neighboring home ranges (c. 100 m²) and may share burrows and sites for thermoregulation. This means that females can be found close together and thus gives males the opportunity to choose a mate. Assortative mating with respect to size was observed in a natural population, as well as a limited number of direct choices of females by males. These results support the results of the choice experiment.     

Home range overlap of mothers and their offspring in the sleepy lizard, Tiliqua rugosa

This paper reports a field investigation of interactions between juveniles and their mothers in the Australian sleepy lizard, Tiliqua rugosa. In their first spring season, juvenile lizards maintain home ranges largely within the home range of their mother. Juvenile home ranges are significantly smaller than those of adult males and females, and juveniles move significantly less often and significantly shorter distances than adults. While siblings were never found together in the spring, they showed a significant tendency to be closer to each other than if they were randomly located in their home ranges. Juveniles and mothers were never found together, nor was there any evidence for any positive (or negative) spatial association. Nevertheless, the extended tolerance of home range overlap represents a greater degree of mother-offspring association than has been previously reported for other lizards. Despite this, the level of parental care can only be described as minimal. Received: 20 June 1997 / Accepted after revision: 29 December 1997     

Intrasexual selection in Mirza coquereli : evidence for scramble competition polygyny in a solitary primate

The primates of Madagascar (Lemuriformes) deviate from fundamental predictions of sexual selection theory in that polygynous species lack sexual dimorphism, have even adult sex ratios and often live in female-dominated societies. It has been hypothesized that intrasexual selection in these species is either reduced or primarily focused on traits related to scramble competition. The goal of this study was to examine these hypotheses by studying the mating system of a solitary nocturnal species, Mirzacoquereli. During a 4-year field study in western Madagascar, I captured and followed 88 individually marked animals. I found that adult males were significantly larger than females, providing the first evidence for sexual size dimorphism in lemurs. In addition, the adult sex ratio was biased in favour of females in 3 out of 4 years. There was no significant sex difference in canine size, however. Males showed pronounced seasonal variation in testis size with a 5-fold increase before and during the short annual mating season. During the mating season, males had more injuries than females and more than quadrupled their home ranges, overlapping with those of more than ten females, but also with about the same number of rivals. Only about one social interaction per 10 h of observation was recorded, but none of them were matings. Together, these results indicate that these solitary lemurs are clearly subject to intrasexual selection and that male-male competition is primarily, but not exclusively, of the scramble type. In addition, they suggest that the above-mentioned idiosyncracies may be limited to group-living lemurs, that social systems of solitary primates are more diverse than previously thought, and that the temporal distribution of receptive females is responsible for this particular male mating strategy. Received: 11 January 1997 / Accepted after revision: 18 April 1997     

Relatedness and avoidance of inbreeding in the lizard, Tiliqua rugosa

Two potential mechanisms for reducing the level of inbreeding, sex-biased dispersal and kin avoidance, were examined in the Australian sleepy lizard, Tiliqua rugosa. The home range centres, and the genotypes at four polymorphic microsatellite DNA loci were determined for adult lizards in a 70-ha study area near Mount Mary, South Australia. From estimates of genetic relatedness, females were as closely related to other females as they were to males, both within the whole study area, and within home ranges. Similarly, males were as closely related to other males as they were to females. This suggests that dispersal in the population is not sex-biased. Sleepy lizards form monogamous pairs during the spring. Partners were less closely related to each other than to other potential partners in the home range area. This suggests active choice of unrelated partners. The mechanism for recognising related from unrelated individuals is unknown, but the behaviour could reduce inbreeding. Received: 7 November 1998 / Accepted: 30 May 1999     

Social monogamy and extra-pair fertilization in an Australian lizard, Tiliqua rugosa

This study investigates social monogamy in the Australian sleepy lizard, Tiliqua rugosa. At a 70-ha site near Mount Mary, South Australia, we radio tracked 55 adult female and 39 adult male lizards during their spring activity periods. Each lizard was observed in 1–5 years. Females were observed with a single male partner on an average of 10.8 days per year, although in 17.3% of cases, females were observed on 2 or fewer days with a male. The most intense pairing period each year was 15 September–15 November when females were with male partners on an average of 36% of observation days. Partnerships lasted an average of 43.3 days each year. After mating in early November, the pairs separated. Observations of females pairing with other males were rare. Most males (82%) were also consistently monogamous, although 7 were observed pairing with 2 females within one season. To investigate paternity, we allowed 21 gravid females to give birth to 42 offspring in the laboratory. We determined genotypes at five polymorphic microsatellite DNA loci for the females, their male partners and their offspring. Four litters (19%) and 6 of the offspring from those litters (14.3%) showed evidence of extra-pair fertilization (EPF). Although the sample sizes are small, females of polygynous males were more likely to experience EPF. Received: 22 February 1998 / Accepted after revision: 23 May 1998     

Female distribution affects mate searching and sexual selection in male northern water snakes (Nerodia sipedon)

Mating systems and sexual selection are assumed to be affected by the distribution of critical resources. We use observations of 312 mating aggregations to compare mate-searching success of male northern water snakes (Nerodia sipedon) in two marshes in which differences in mating substrate availability resulted in more than fourfold differences in female dispersion. Reproductive males had significantly larger home ranges where females were dispersed than where females were clumped. The number of females encountered by males increased significantly with male home range size where females were dispersed, and decreased significantly where females were clumped. Where females were clumped, males were more likely to encounter other males when they located females. We found no evidence in either population that mate searching was energetically expensive or that males with relatively more energy had larger home ranges. However, males with greater fat reserves at the start of the season participated in more mating aggregations when females were dispersed, suggesting that fat reserves could affect a male’s willingness to attempt mating or to persist in aggregations. When females were dispersed there was weak stabilizing selection acting to maintain male body size (β=–0.14), but strong directional selection favoring larger (β=0.50) and fatter (β=0.37) males. Over 7 years, the intensity of selection favoring larger males varied substantially (β=0.14–1.15), but that variation was not related to variation in the operational sex ratio. We found no evidence of directional selection on either body size (β=0.05) or fat reserves (β=0.10) of males when females were spatially clumped. Overall, the distribution of females had a pronounced effect on male behavior, on the factors that affected male success in locating females, and probably on the extent of sperm competition once females had been located. Received: 23 November 1998 / Received in revised form: 9 August 1999 / Accepted: 18 August 1999     

Large males dominate: ecology,social organization,and mating system of wild cavies,the ancestors of the guinea pig

Ecological factors differently affect male and female animals and thereby importantly influence their life history and reproductive strategies. Caviomorph rodents are found in a wide range of habitats in South America and different social and mating systems have evolved in closely related species. This permits to study the impact of ecological factors on social evolution. In this study, we investigated the social organization and the mating system of the wild cavy (Cavia aperea), the ancestor of the domestic guinea pig, in its natural habitat in Uruguay. Based on our laboratory investigations, we expected a polygynous system with large males controlling access to females. Results from radiotelemetry and direct observations showed that females occupied small stable home ranges which were largely overlapped by that of one large male, resulting in a social organization of small harems. In some cases, small satellite males were associated with harems and intermediate-sized roaming males were occasionally observed on the study site. However, microsatellite analyses revealed that offspring were exclusively sired by large males of the same or neighboring harems, with a moderate degree of multiple paternity (13–27%). Thus, the mating system of C. aperea can be described as polygynous and contrasts with the promiscuous organization described for other species of cavies (Cavia magna, Galea musteloides and Microcavia australis) living under different ecological conditions. Our findings stress the strong impact of environmental factors on social evolution in Caviomorphs as resource distribution determines female space use and, thereby, the ability of males to monopolize females. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Social organization and nest co-occupancy in Peromyscus californicus,a monogamous rodent

Summary Home ranges, social organization, and nest co-occupancy of Peromyscus californicus were studied using radiotelemetry at the Hastings Natural History Reservation, California. Mated pairs were ascertained by the transfer of fluorescent pigments from lactating females to putative fathers. Mated pairs had largely overlapping home ranges that were not statistically distinguishable, whereas adjacent adults had mostly exclusive, statistically distinguishable home ranges. There was no difference in the mean home range of males and females, but mated females tended to have smaller ranges than their mate. Home range size was extremely variable (range: 150–3788 m²) and averaged 1161 m² across all individuals. Male home range size was inversely correlated with population density, suggestive of a social influence on home range. Putative fathers spent comparable amounts of time to females in the nest — presumably caring for the young — which supports previous laboratory reports of paternal care in this species. All data collected in this study are consistent with previous suggestions that P. californicus live in semi-permanent family groups and are monogamous. Offprint requests to: D.O. Ribble     

Dynamics of space use and male vigour amongst wood mice,Apodemus sylvaticus,in the cereal ecosystem

The spatial organisation of male and female wood mice,Apodemus sylvaticus, was investigated in a large-scale radio-tracking study on arable farmland near Oxford, United Kingdom, during the breeding season. Both sexes had significantly larger home ranges in the breeding season than at other times, and the breeding season home ranges of male (X = 1.44 ha) were significantly larger than those of females (X = 0.49 ha). Home range overlap was significantly greater between males, and between males and females, than it was between females. Overlap between males tended to be greatest in heavily utilised areas. Except during sexual consortship, there was minimal evidence of dynamic interaction among individuals. Home range sizes of breeding males varied widely, as did their body weights. There was no relationship between male body weight and home range size or any other movement parameter. However, males with the largest home ranges had the highest scores on all other movement parameters, indicating that they expended more energy in movement. These more vigorous males had access to the home ranges of more females than did males with small home ranges.     

Food or sex—males and females in a sex role reversed pipefish have different interests

In a sex role reversed pipefish, Syngnathus typhle, we found that basic life history allocations were directly influenced by sexual selection. We investigated time allocation to foraging and mating, respectively, in a choice experiment, giving males and females, of small or large body size, a choice between food and a potential partner. We found that males were more interested in foraging than mating, i.e., were more frequently observed in front of the food than in front of the partner, whereas females were more interested in the potential partner. This reflects sexual selection operating differently on the two sexes, as males and females are relatively similar in other life history traits, such as growth, mortality, age of maturity, dispersal, and parental expenditure. Moreover, large individuals allocated more time to mating activities, small to feeding. Individuals more interested in mating compared to food were subsequently more critical when given a choice between a large (high-quality) and a small (low-quality) partner, whereas individuals more interested in food were not selective. These findings are consistent with our predictions: sex-role reversed males can be relatively sure of achieving one or more matings, and should allocate more time to feeding and, hence, to parental investment, growth and/or future reproduction. Females, on the other hand, have more uncertain mating prospects and should allocate time to imminent reproductive activities, thereby foregoing other life history traits such as growth and future egg production. By this, they also sacrifice future fecundity and attractiveness.     

Mating in the red-sided garter snake,Thamnophis sirtalis parietalis: differential effects on male and female sexual behavior

Summary Female red-sided garter snakes, Thamnophis sirtalis parietalis, become unattractive to most males after mating in the field and in the laboratory. Male red-sided garter snakes vary in their latencies to court attractive females following copulation, with courtship resuming in minutes to hours. Unsuccessful males in mating balls disperse from mating pairs, but are not residually inhibited from courting attractive females. These patterns of behavior indicate that males have evolved mechanisms to maximize opportunities for copulation with several females, while females mate only once per season.     

Seasonal variation in male-female competition,cooperation and selfish hoarding in a monogamous songbird

Both cooperation and conflict between the sexes are commonplace in monogamous mating systems. However, little is known about how cooperation and competition varies seasonally in monogamous species that maintain permanent territories. We presented territorial pairs of male and female New Zealand robins (Petroica australis) with a large supply of insect prey at monthly intervals for 2 years. Behavioural observations after food presentation were then made to quantify seasonal and sexual differences in aggressive interactions over prey, prey acquisition rates, mate provisioning, offspring provisioning, selfish food hoarding and cache retrieval. Data were used to evaluate sex-specific behavioural strategies of mediating competition for food. Results showed that males aggressively excluded females from experimental food sources year-round. Females only accessed food sources when males left them unattended. Consequently, females acquired fewer prey than males. After controlling for differences in prey acquisition, both sexes consumed similar amounts of prey in the non-breeding season. Even though males aggressively excluded females from accessing food sources directly, males fed large amounts of prey to females during the breeding season. Both sexes provisioned young at similar rates. Males cached less prey than females in the breeding season but more prey than females in the non-breeding season. Females showed similar caching intensities year-round. Although males tried to defend their hoards, females frequently retrieved male-made caches. Overall, results showed that although New Zealand robins cooperate to raise offspring during the breeding season, conflict between the sexes occurs year-round. Males and females display different behavioural strategies to gain access to experimental food sources, which appear to lessen male–female competition for food and evenly distribute food resources between the sexes.     

The cause of size-assortative mating in the leaf beetle Trirhabda canadensis (Coleoptera : Chrysomelidae)

Summary Examples of positive assortative mating by body size are abundant but its causes remain controversial. I show that size-assortative mating occurs in the chrysomelid beetle Trirhabda canadensis and I test a series of alternative hypotheses to explain how this mating pattern comes about. Results suggest that assortative mating in this beetle is due to the greater ease with which size-matched pairs can achieve intromission, and not due to size-biased skews in the availability of mates or mate choice favoring large individuals. There was no correlation between male and female elytron length (a measure of body size) at the initiation of courtship, but pairs assorted positively by size at the onset of intromission. Moreover, in the laboratory, there was a negative correlation between male and female size for pairs engaged in courtship that terminated without mating. Assortative mating was not associated with a large-male mating advantage and there was no evidence of female choice of large males. Nor was there unequivocal evidence for male choice of large females; although mating females were slightly larger and considerably heavier than solitary females, males did not differ in the frequency with which they rejected large and small females. Assortative mating in T. canadensis appeared to be caused by the lower ability of mismatched pairs to achieve intromission after an encounter, both when males were larger and when they were smaller than the female.     

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.     

Variation in homeward orientation performance in the sleepy lizard (Tiliqua rugosa): effects of sex and reproductive period

Sleepy lizards are monogamous skinks which show high pair fidelity. This study reveals inter- and intrasexual differences in homeward orientation performance in this lizard. Male and female lizards were displaced during three phases of the spring activity period, the pre-pairing, pairing/mating, and post-pairing periods. All groups (with the exception of post-pairing males) were significantly oriented homewards, but males were significantly better oriented towards home than females during the pairing period. Furthermore, males were significantly better homeward oriented during the pre-pairing and pairing periods than in the post-pairing period. Similar results were observed for rate of movement away from the release site. In sleepy lizards, sex-based differences in homing behaviour are unlikely to be attributable to differences in the area of familiarity, or availability of orientation mechanisms. However differences in homing motivation may explain these differences. Males may miss mating if absent from the home range during the pre-pairing and pairing periods, while females may still be able to obtain a mating even when absent. Females however may be more motivated than males to return to the familiar home range during the post-pairing period to ensure efficient feeding during internal embryo development. Received: 16 February 1998 / Accepted after revision: 28 March 1998     

Home range and social behavior of three species of European Pitymys (Mammalia,Rodentia)

Summary Habits, home range, and social behavior of three species of voles of the genus Pitymys, P. multiplex, P. subterraneus, and P. savii, were studied in Tessin (Switzerland) by radioactive tagging. P. multiplex and P. savii are fossorial and often use the burrow systems of moles (Talpa); P. subterraneus moves around on the surface under dense vegetation. Males and females of P. multiplex were territorial and their home ranges averaged 342 and 229 m² respectively. The mating system appeared to be monogamous. P. savii male and female home ranges averaged 445 and 298 m² respectively. Groups of up to one to three adult females, one or more adult males, and several young, occupied exclusive communal territories. The mating system appeared to be flexible, including monogamy, polygyny, and polyandry. P. subterraneus females occupied exclusive territories (mean 256 m²) overlapped by the home ranges of one or more males (1025 m²). The mating system appeared to be promiscuous. Individuals of the three species occupied one to seven nests. In P. savii all family members utilized three or more communal nests at the same time or alternatively. Co-nesting between males and females of P. multiplex or P. subterraneus were observed less frequently.     

Mouse lemurs in space and time: a test of the socioecological model

Socioecological theory predicts that the distribution of fertile females in space and time is the major determinant of male spacing behavior and mating strategies. Using a small nocturnal Malagasy primate, the gray mouse lemur (Microcebus murinus), we determined the spatiotemporal distribution of estrous females during the brief annual mating season to examine the predictive power of the socioecological model for male mating strategies. Mouse lemurs are particularly interesting in this respect because this polygynous species is characterized by seasonal reproduction, seasonally reversed sexual dimorphism, and relatively large testes. All resident animals in our 8-ha study area, a total of 30 adult males and 27 adult females, were individually marked and regularly recaptured to determine female reproductive status and to obtain home range data. We found that the mating season is limited to 4 weeks following female emergence from hibernation. Only 3-9 females could have synchronized estruses during a given week, indicating a moderately high male monopolization potential. However, receptive females were not spatially clumped and male ranges overlapped with those of many other rivals. Therefore, we suggest that individual powerful males may be unable to defend exclusive permanent access to receptive females because of prohibitive costs of range defense resulting from the strongly male-biased operational sex ratio and the corresponding intruder pressure. Our general conclusions are (1) that the socioecological model provides a useful heuristic framework for the study of mating systems, but that (2) it does not specify the degree of spatiotemporal clumping of receptive females at which male mating strategies switch among mate guarding, spatial exclusion of rivals, and roaming, and that (3) the operational sex ratio can have profound effects on male mating strategies as well.     

Chemosensory mate recognition may facilitate prolonged mate guarding by male snow skinks, Niveoscincus microlepidotus

We provide evidence that male lizards can use chemosensory cues to identify individual females and probably therefore maintain long-term associations with these females in the wild. In the laboratory, males preferentially followed the scent trail of their vitellogenic female “partner” rather than that of another vitellogenic female. Our 5-year field study of the small viviparous scincid lizard (Niveoscincus microlepidotus) in alpine Tasmania showed that sexually mature males and females commonly formed “pairs” for long periods (on average 29 days). These pairs occurred primarily during the mating season, always involved one adult male and one adult female, and usually involved vitellogenic rather than gravid females. Our laboratory experiments suggest that a significant factor in maintaining those prolonged partnerships is male scent trailing of partners. Received: 28 October 1997 / Accepted after revision: 28 May 1998     

Sexual dimorphism and the mating ecology of polar bears (Ursus maritimus) at Svalbard

We assessed the role of size, mass, and age in mating and non-mating polar bears (Ursus maritimus) at Svalbard, Norway, during the spring breeding season. The ratio of male to female mass, in male-female pairs, ranged from 1.00 to 3.02 ([`(x)] = 1.99 \overline x = 1.99 ) indicating that mating males were larger than mating females but with substantial variation. Paired males were older than unpaired males and male mass was related to age. However, males paired with females were not significantly different in body mass from those males caught alone. Wounds and scars resulting from fights between males began at about 6 years of age and peaked at about 17 and 20 years of age, respectively. The frequency of broken canines in males, presumably due to increased male-male conflicts, increased with age but showed little increase in females. The wide range of male size in male-female pairs and the age-related signs of injury suggest that male polar bears engage in both scramble competition and contest competition for access to breeding females. The mating system of polar bears is variable but is best described as female defense polygyny or serial monogamy.