Forced dispersal of juvenile guanacos (<Emphasis Type="Italic">Lama guanicoe</Emphasis>): causes,variation, and fates of individuals dispersing at different times

We examined adult-juvenile conflict in the guanaco (Lama guanicoe). During spring, territorial males become increasingly aggressive toward all juveniles born the previous year and begin expelling them from family groups. In an apparent effort to reduce aggression, juveniles display submissive crouches when being observed, approached, or attacked by the territorial male. Therefore, we assessed the influence of juvenile submissive behavior on the timing of dispersal and also examined if dispersal time was related to survival and reproductive performance as adults. We also evaluated hypotheses regarding the evolution of juvenile mammalian dispersal in the context of if and how each may favor the forced dispersal of juvenile guanacos by territorial males. Juveniles generally dispersed in late spring and early summer, and a nearly equal proportion of females (n=46; 48%) and males (n=49; 52%) dispersed. More-submissive animals generally dispersed later than less-submissive animals. Juvenile sex and dispersal time were not related to survival. In contrast, juvenile sex and dispersal time were related to reproductive performance. The probability of reproducing was highest when juveniles dispersed early and decreased with increasing time in family groups prior to dispersal. The largest proportion of juveniles was forced to disperse during a 2-week interval following the peak of the breeding season. Competition for food resources is likely very intense at this juncture and territorial males may force older juveniles to disperse in order to divert food resources to younger neonates. Additionally, juveniles may be forced to disperse after territorial males mate their mothers to prevent lost mating opportunities, because females leave territories when their offspring disperse and possibly prior to mating with males. We conclude that the forced dispersal of juvenile guanacos by territorial males is ultimately driven by competition for food resources on territories. The timing of dispersal, however, may be tempered by the chronology of matings between territorial males and particular adult females, and/or genetic relatedness between territorial males and juveniles.     

Dispersal: an alternative mating tactic conditional on sex ratio and body size

Summary Small male milkweed beetles are less successful at obtaining mates than are larger males. Larger males usually win fights and prevent smaller males from obtaining mates and from choosing larger more fecund females as mates. When sex ratios are male-biased, smaller males are particularly likely to experience these mating disadvantages. It follows that smaller males should be especially responsive to their local competitive environment and behave so as to minimize the mating disadvantages of their smaller size. This paper tests the hypothesis that smaller males disperse from host plant patches with male-biased sex ratios and remain in patches with female-biased sex ratios more readily than larger males.Results show both larger and smaller males disperse from patches with male-biased sex ratios more frequently than from patches with femalebiased sex ratios. As predicted, however, small males are more likely to disperse from patches with male-biased sex ratios and remain in patches with female-biased sex ratios than are larger males.The data also show that smaller males dispersing from patches with male-biased sex ratios obtain more matings than non-dispersing males.For milkweed beetles, moving between patches can be viewed as an alternative mating tactic conditional on male body size and local sex ratio.     

Disappearance and dispersal of Spermophilus elegans juveniles in relation to behavior

Summary Patterns of disappearance and dispersal of Spermophilus elegans juveniles during the first 6-weeks postemergence were compared for 1977 and 1978 and related to quantitative and qualitative changes in social interaction involving juveniles.Juvenile disappearance (emigration or mortality) and dispersal within the study site varied between the sexes within and between years. Female disappearance and dispersal were significantly greater in 1977, and male losses in 1978 significantly exceeded male losses in 1977. Greater female loss in 1977 resulted in total male — female losses being equivalent, whereas in 1978 juvenile loss was strongly biased toward males by the end of the 6-week period.Greater female loss in 1977 was attributed primarily to increased aggression between female juveniles in that year because of larger average litter size with more females per litter. Increased disappearance of males in 1978 showed no correlation with litter size or relative increase in number of males per litter. Male young interacted with individuals of several age/sex classes, and a possible behavioral influence on male disappearance was increased aggressiveness by yearling males toward juvenile males in 1978.Behavior appeared to act as a proximate factor in juvenile disappearance and dispersal, and the observed differences between how male and female juveniles interacted in 1977 and 1978 were hypothesized to reflect the operation of different selective pressures to increase individual male or female fitness.     

Dispersal and social organization in the Neotropical Grey sac-winged bat Balantiopteryx plicata

Recent evidence suggests that tropical bats may frequently depart from the predominant mammalian male-biased dispersal pattern. So far, two emballonurid bat species that are closely related to our study species (Grey sac-winged bat, Balantiopteryx plicata) have been found to exhibit exceptional female-biased dispersal that is in accordance with father–daughter inbreeding avoidance. In contrast, using a combination of long-term behavioral observations of banded bats and DNA sequencing of the mitochondrial d-loop, our results suggest that B. plicata is the first Neotropical emballonurid with female philopatry and frequent male dispersal. However, just like in the other emballonurids, the age of females at first conception fell below the tenure of males. Thus, philopatric B. plicata females might face a father–daughter inbreeding risk if mating with males from their roosts. Such risk could be circumvented if mating occurs outside the nursing roost, e.g., in male mating aggregations. In contrast to other Neotropical emballonurid bats, the Grey sac-winged bat forms colonies with a greatly male-biased sex ratio (only 5 and 21 % females). Males of such colonies showed high roost fidelity and the sex ratio did not change throughout the year and for up to many years, suggesting an important role in mating. We conclude that studying the diverse mating and dispersal patterns of Neotropical emballonurids shows great potential to enlarge our understanding on how the proposed ultimate causes (i.e., avoidance of inbreeding and local competition between kin) affect the evolution of sex-biased dispersal.     

Laboratory and field evidence of sex-biased movement in the invasive round goby

Activity levels are modulated by trade-offs between reducing predation risk and the need to move in order to find food or mates. Because these trade-offs affect males and females differently, many species show sex-specific movement, dispersal patterns, and spatial navigation capacities, with the sex that gains the most from territory ownership often dispersing less. Unlike mammals and birds, sex differences in movement among fishes remain poorly studied, and the connections between tests of movement propensity in the laboratory and in the field are rarely made. Here, we examine the differences in movement between male and female round goby (Neogobius melanostomus) in both laboratory and field settings. This fish species is invasive in North America and currently undergoing further range expansions. In the laboratory, round goby males were more active and explored a novel environment more readily than did females. A large-scale mark–recapture study in Lake Ontario over two years revealed that males moved more than females between years, but there were no within-year sex differences. Thus, round goby display male-biased movement patterns, providing a comparison point to dispersal patterns in other taxa. Understanding sex-specific movement of round goby in the field will also help predict dispersal and population dynamics, both in areas where round goby have already become established and where they are continuing to invade.     

Sex-specific fitness consequences of dispersal in Siberian jays

In most birds, natal dispersal is female-biased, but the selective pressures leading to this pattern have rarely been explored with comprehensive data on lifetime reproductive success. In territorial birds, the benefit of philopatry should be higher for males than for females when males establish territories for which knowledge about the local environment is important. As females may use male characteristics for mate choice, and hence indirectly for territory choice, the benefit from the direct knowledge of the local environment may be lower for females than males. We tested this hypothesis using data from a long-term study of group living corvids, the Siberian jays (Perisoreus infaustus). In this species, the socially dominant offspring delay dispersal while the sub-dominant offspring leave the family group directly after reaching independence. Our results show that natal dispersal distance (a proxy for local knowledge) was related to sex and dispersal timing (a proxy for “quality”): Females and early dispersers traveled further on average than males and delayed dispersers. Furthermore, dispersal distance and timing were negatively related to the number of recruits produced over an individual’s lifetime in males, but not in females. Hence, the results support the hypothesis that the female-biased natal dispersal found in this and other bird species may come about through higher lifetime reproductive success of philopatric males than females.     

Social behavior of free-ranging juvenile sooty mangabeys (Cercocebus torquatus atys)

Sooty mangabeys are terrestrial monkeys exhibiting female philopatry and male dispersal. Studies in captivity as well as in the wild have found that adult females form linear dominance hierarchies. However, while captive studies found no evidence for a matrilineal social system, a previous study in Taï National Park, Ivory Coast, suggested that relatedness could influence both dominance rank and affiliation pattern among adult females. Here I test whether the dominance rank, coalitionary behavior, and affiliative behavior of juveniles in a group of free-ranging mangabeys in the Taï National Park are in accordance with a matrilineal, individual, or age-related dominance system. I found that juvenile females' dominance ranks remained stable over time and were highly correlated with the dominance ranks of their mothers, whereas juvenile males' dominance ranks were initially correlated with the ranks of their mothers, but showed greater instability with increasing age. Moreover, coalitions occurred mainly between juveniles and animals that were close in rank, including their mothers and siblings. Finally, juvenile females associated and groomed preferentially with close-ranking juvenile and adult females. Juvenile males showed similar preferences in affiliation with adult females, but when associating with juvenile males, they preferred peers. The observed social behavior of free-ranging juvenile sooty mangabeys resembled the social behavior described for juveniles of many matrilineal primate species.     

Pre-reproductive survival in a tropical bird and its implications for avian life histories

The factors that affect survival until reproduction are essential to understanding the organization of life histories within and among species. Theory predicts, for example, that survival until reproduction influences the optimum level of reproductive investment by parents, which might partly explain prolonged parental care in species with high first-year survival. Tests and refinements of life-history theory have been hampered, however, by a lack of field-based estimates of pre-reproductive survival, especially for tropical species, which have been the subject of many comparative analyses. Tropical species are predicted to have higher first-year survival and delayed reproduction compared to Northern Hemisphere species. We estimated survival until reproduction, age at first reproduction, and sources of variation in juvenile survival in a Neotropical passerine, the Western Slaty-Antshrike (Thamnophilus atrinucha), in central Panama. We observed that fledged antshrikes had 76% survival through the dependent period and 48% survival to the age of 1 year; survival rate was lowest during the first week after leaving the nest. Timing of fledging within the breeding season, fledgling mass, and age at dispersal influenced survival, while sex of offspring and year did not. Individuals did not breed until two years of age, and post-fledging pre-reproductive survival was 41% of annual adult survival. High survival until reproduction in antshrikes balanced their low annual productivity, resulting in a stable population. Survival during the post-fledging period of dependence and the first year of independence in the Western Slaty-Antshrike exceeded estimates for Northern Hemisphere species. This difference appears to be associated with the extended post-fledging parental care, delayed dispersal, low costs of dispersal, and the less seasonal environment of antshrikes.     

Offspring sex ratio in relation to parental structural size and body condition in the long-lived wandering albatross (<Emphasis Type="Italic">Diomedea exulans</Emphasis>)

Sex ratio theory is one of the most controversial topics in evolutionary ecology. Many deviations from an equal production of males and females are reported in the literature, but few patterns appear to hold across species or populations. There is clearly a need to identify fitness effects of sex ratio variation. We studied this aspect in a population of a long-lived seabird, the wandering albatross (Diomedea exulans), using molecular sex-identification techniques. We report that parental traits affect both (1) fledgling traits in a sex-dependent way and (2) chick sex: Sons are overproduced when likely to be large at fledging and, to a lesser extent, daughters are overproduced when likely to be in good body condition at fledging. Because for the same population, a previous study reported that post-fledging survival was positively affected by size in males and by body condition in females, our results suggest that wandering albatrosses manipulate offspring sex to increase post-fledging survival.     

Operational sex ratio and alternative reproductive behaviours in the European bitterling,<Emphasis Type="Italic"> Rhodeus sericeus</Emphasis>

We investigated the effects of male population density and male-biased operational sex ratio (OSR) with constant and limited resource density on male mating tactics shown by a freshwater fish, the European bitterling, Rhodeus sericeus. This species spawns inside living unionid mussels. Large males defended territories and were aggressive towards conspecifics under equal sex ratios. They also monopolised pair spawnings with females, releasing 98% of all sperm clouds during mating. However, the mating tactic changed at high male density where large males ceased to be territorial and instead competed with groups of smaller males to release sperm when females spawned. Large, medium and small males now obtained 61%, 33%, and 6% of sperm releases respectively, thereby reducing the opportunity for sexual selection by half. Females spawned at equal rates in the two densities of males, despite lower courtship at high density. These results run counter to the usual expectation that an increasingly male-biased OSR should lead to higher variance in male mating success. Instead, the use of alternative reproductive behaviours by males can lead to lower resource competition and mating variance at high male densities.     

Dispersal behaviour and colony structure in a colonial spider

Group living in spiders is characterised by two principle modes, the cooperative social mode and the colonial non-cooperative mode. Kin-relationships due to reduced dispersal determine population genetic structure in social spiders, but the dispersal mechanisms underlying group structure remain poorly understood in colonial spiders. Assuming similar ecological benefits of group living, we address the question whether reduced dispersal shapes population structure in a colonial spider, Cyrtophora citricola (Araneidae). We analysed dispersal by studying settling decisions under semi-natural conditions in experimental trees with and without colonies, and in natural populations, we estimated dispersal and colony structure using population genetic analyses. The propensity to disperse decreased with increasing age in experimental colonies. Adult females did not disperse in the experiment. Sub-adult female spiders preferred trees with a colony to trees without a colony. Dispersal in third instar juveniles was influenced significantly by wind but not by the presence of a colony. Thus, we showed that being in a colony did not inhibit juvenile dispersal, but pre-mating females were philopatric. Genetic differentiation among colonies in natural populations was heterogeneous, colonies being either little or highly differentiated. The heterogeneous structure is likely caused by colony founding by one or a few females followed by dispersal among perennial colonies. Gene flow, however, was slightly male-biased. The experimental and indirect, genetic approaches combined showed that dispersal and the breeding system of C. citricola resemble that of solitary spiders, with juvenile dispersal occurring in both sexes, while the colonial distribution is maintained by female philopatry.     

Alternative reproductive tactics and the impact of local competition on sex ratios in the ant Hypoponera opacior

The ant Hypoponera opacior exhibits alternative reproductive morphs of males and females associated with distinct sexual behaviours. Our long-term study reports strong seasonality in sexual production with a mating season in early and one in late summer. Winged (alate) reproductives emerge in June, swarm during the monsoon season and establish new colonies independently. In contrast, wingless worker-like (ergatoid) reproductives that appear in late August mate within their natal or adjacent nests and either do not disperse or establish new nests close by. These divergent dispersal patterns allowed us to analyse the impact of local factors on investment strategies by comparing sex allocation between and within the two reproductive events. The optimal sex ratio for ergatoid reproductives should be influenced both by competition for matings between brothers (local mate competition) and rivalry among young locally dispersing queens for workers, nest sites or food (local resource competition). The greater importance of local resource competition was demonstrated both by a male-biased sex ratio for wingless sexuals and a stronger increase in the number of males with total sexual production than for the number of queens. Microsatellite analysis revealed that inter-nest variation in relatedness asymmetry cannot explain split sex ratios in the August generation. Instead, nests with related ergatoid males raised a male-biased sex ratio contrary to the expectations under local mate competition. In conclusion, male bias in wingless H. opacior indicates that local mate competition is less strong than local resource competition among ergatoid queens over the help of workers during nest foundation.     

Sex allocation and local mate competition in Old World non-pollinating fig wasps

The populations of many species are structured such that mating is not random and occurs between members of local patches. When patches are founded by a single female and all matings occur between siblings, brothers may compete with each other for matings with their sisters. This local mate competition (LMC) selects for a female-biased sex ratio, especially in species where females have control over offspring sex, as in the parasitic Hymenoptera. Two factors are predicted to decrease the degree of female bias: (1) an increase in the number of foundress females in the patch and (2) an increase in the fraction of individuals mating after dispersal from the natal patch. Pollinating fig wasps are well known as classic examples of species where all matings occur in the local patch. We studied non-pollinating fig wasps, which are more diverse than the pollinating fig wasps and also provide natural experimental groups of species with different male morphologies that are linked to different mating structures. In this group of wasps, species with wingless males mate in the local patch (i.e. the fig fruit) while winged male species mate after dispersal. Species with both kinds of male have a mixture of local and non-local mating. Data from 44 species show that sex ratios (defined as the proportion of males) are in accordance with theoretical predictions: wingless male species<wing-dimorphic male species<winged male species. These results are also supported by a formal comparative analysis that controls for phylogeny. The foundress number is difficult to estimate directly for non-pollinating fig wasps but a robust indirect method leads to the prediction that foundress number, and hence sex ratio, should increase with the proportion of patches occupied in a crop. This result is supported strongly across 19 species with wingless males, but not across 8 species with winged males. The mean sex ratios for species with winged males are not significantly different from 0.5, and the absence of the correlation observed across species with wingless males may reflect weak selection to adjust the sex ratio in species whose population mating structure tends not to be subdivided. The same relationship is also predicted to occur within species if individual females adjust their sex ratios facultatively. This final prediction was not supported by data from a wingless male species, a male wing-dimorphic species or a winged male species. Received: 27 July 1998 / Received in revised form: 11 January 1999 / Accepted: 16 January 1999     

Tests of the harassment-reduction function and frequency-dependent maintenance of a female-specific color polymorphism in a damselfly

Color polymorphisms have provided classical examples of how frequency-dependent selection maintains genetic variation in natural populations. Here we tested for the first time, the hypothesized adaptive function of a female-specific color polymorphism in odonates to lower male harassment towards females generally. Under conditions controlling for sex ratio, population density and morph frequency, we also tested two major frequency-dependent selection hypotheses for the maintenance of the polymorphism. Using groups of captive Enallagma hageni, whose females are either green or a male-like blue, we varied morph frequency at two sex ratios. We quantified sexual harassment towards females by visual observations, and by the presence of dust on females that was transferred from dusted males. Per capita harassment rate for the female-monomorphic treatments did not differ from that of the female-polymorphic treatments. At a male-biased sex ratio, per capita harassment rate towards blue, but not green females increased with morph frequency, providing partial support for frequency-dependent selection resulting from male learning of female morphs. Even at high frequency, green females were not harassed more than blue, contrary to the prediction that males should always recognize green females as mates. Moreover, frequency-dependent harassment towards blue females was not detectable using harassment measured with dust evidence, which greatly underestimated the incidence of sexual harassment. Our findings identified problems with the use of insectaries and the dusting technique to quantify male sexual harassment towards females, as well as with a past insectary experiment on Ischnura elegans that failed to demonstrate frequency-dependent harassment.     

Sex ratios,mating behavior and sexual size dimorphism of the northern water snake,Nerodia sipedon

Competition among males to mate is generally associated with male-biased size dimorphism. In this study we examine mating behavior in the northern water snake (Nerodia sipedon), a species in which males are much smaller than females despite substantial competition among males to mate. Competition among males was a consequence of a male-biased operational sex ratio due to slightly higher female mortality from a birth sex ratio of 1 : 1, and, in 1 year, more synchronous and longer mating activity by males. Approximately one-third of both males and females appeared not to mate in a given year. Larger males were generally more likely to attempt mating, but size did not explain the variance in the number of aggregations in which individual males participated. Within aggregations, males that were successful at achieving intromission were larger than unsuccessful males in 1 of 2 years. Variation in condition (mass relative to length) and relative tail length were not generally useful predictors of either mating effort or success in males. Because large size was often advantageous to males, sexual size dimorphism appeared not to be a consequence of sexual selection favoring smaller males. Because sexual dimorphism was evident at birth, and both males and females matured sexually at about 4 years, sexual dimorphism was not simply a consequence of one sex growing at the maximum rate for longer. Female fecundity increased with size, and sex differences in size-fecundity relations may underly the pattern of sexual size dimorphism. However, because multiple mating by females is common, sperm competition is likely to be important in determining male reproductive success. Therefore, allocation of energy to sperm rather than growth may also prove to be an important influence on male growth rates and sexual size dimorphism.     

Possible causes and consequences of philopatry and breeding dispersal in red-backed shrikes <Emphasis Type="BoldItalic">Lanius collurio</Emphasis>

Studies of animal breeding dispersal have often focused on possible causes, whereas its adaptive significance has received less attention. Using an information-theoretic approach, we assessed predictions of four hypotheses relating to causes and consequences of breeding dispersal in a migratory passerine, the red-backed shrike Lanius collurio. As predicted by the reproductive performance hypothesis, probability of breeding dispersal in females (though not in males) decreased with increasing annual average number of fledglings produced in the past year, but there was no association with conspecific reproductive performance in either sex. The site choice hypothesis, stating that individuals disperse to improve breeding site quality, received support in males only, as dispersal probability was positively associated to a measure indicating low territory quality. The social constraints hypothesis, referring to dispersal in relation to intraspecific interactions, received little support in either sex. The predation risk hypothesis was hardly supported either. Consequences of dispersal were marginal in both sexes because neither fledgling production in females, nor territory quality in males improved after dispersal. In addition, males settled on territories closer to the forest edge than those occupied predispersal, which is opposite to the prediction of the predation risk hypothesis. We conclude that own reproductive success was the major factor determining dispersal behavior in females, whereas territory quality and possibly predation risk were most important in males. Overall, breeding dispersal appeared not to be adaptive in this dense population inhabiting an optimal habitat.     

Baboons adjust secondary sex ratio in response to predictors of sex-specific offspring survival

Summary Most theories of sex ratio adjustment assume that parents will adjust the sex ratio of births (secondary sex ratio) in a manner that maximizes offspring reproductive success (as long as this does not jeopardize parental reproductive success). Survival to maturity is typically the largest component of variance in offspring reproductive success. This should make environmental predictors of sex-specific offspring survival strong predictors of secondary sex ratio adjustment. We tested this survivorship maximization hypothesis for secondary sex ratio adjustment using data from a 17-year demographic study of 315 yellow baboon infants at Mikumi National Park, Tanzania. Sex differences were found in the degree to which several social and ecological conditions affected infant survival to 1 year. Female, but not male, infant survival was inversely correlated with birth order and the proportion of infant females in their birth cohort. Male, but not female, infant survival was inversely correlated with the amount of rainfall early in the year (January) and the proportion of infant males in their birth cohort; male survival also was positively correlated with maternal dominance rank. The consistency and timing of these effects across years suggested that such information was, in fact, available to females around the time of conception. Most importantly, social and ecological conditions that predicted improved survivorship of a given sex also were positively correlated with production of that sex. Early births were female-biased; and low January rainfall was correlated with a male-biased sex ratio, becoming increasingly female-biased as January rainfall increased. However, no sex ratio effects were correlated with maternal rank. Data supported the hypothesis that females adjusted secondary sex ratio in a manner that maximized sex-specific infant survival. This hypothesis also offered a plausible explanation for some of the contradictory data that have arisen from studies of maternal rank effects on sex ratio both within and between species.Correspondence to: S. K. Wasser     

Male and female developmental differences in chemosensory investigations by African elephants (Loxodonta africana) approaching waterholes

Although communication is vital for members of a social species, the sexes may differ in the type and degree of information sought. In many polygynous societies, males search for reproductively active females and compete intrasexually for access to females with older males often being most successful. In social mammals, females may mature sooner than males and thus at an earlier age behave more like adults. This maturation may include the assessment of potential mates directly or via indicative signals. In this study, we observed the behavior of African elephants (Loxodonta africana) during their approach to waterholes. Waterholes provide an opportunity for elephants to investigate conspecific chemical signals from feces and urine, and each other. We examined the presence of sexual dimorphism in behaviors of the trunk that are indicative of olfactory investigation. We predicted that upon approach to a waterhole, adult males would show greater interest than females in conspecific chemical signals. Females were predicted and observed to exhibit adult-like rates of investigatory behavior at a younger age than males because females mature sooner. Adult males took the longest to reach the waterhole in the last 100 m of approach; they also demonstrated greater investigatory rates to conspecific feces. Each sex showed adult-type investigative behaviors with the trunk before the age of reproduction. Rather than showing a common chronological developmental pattern across sex, the exhibition of investigatory, chemosensory behaviors reflected sex-specific changes in reproductive development, perhaps reflective of the relative strength of intra- and intersexual selection on communication patterns.     

Sexual conflict in the snake den

Red-sided garter snakes (Thamnophis sirtalis parietalis) court and mate in spring, soon after they emerge from large communal overwintering dens in south-central Manitoba. Because of a massive bias in the operational sex ratio, every female attracts intense courtship from dozens to hundreds of males. We suggest that this courtship constitutes significant ”harassment,” because it delays the females’ dispersal from the den and hence increases their vulnerability to predation. Small females may face the greatest costs, because they are less able to escape from amorous males (who court all females, even juvenile animals). Our measurements show that males are stronger and faster than females. Experimental trials confirm that the locomotor ability of females (especially small females) is greatly reduced by the weight of a courting male. Arena trials show that intense courtship stimulates females to attempt to escape. Remarkably, some females that are too small to produce offspring may nonetheless copulate. This precocious sexual receptivity may benefit juvenile females because copulation renders them unattractive to males, and thus allows them to escape more easily from the den. Female ”tactics” to escape male harassment may explain other puzzling aspects of garter snake biology including size-assortative mating, temporal patterns in dispersal from the den, avoidance of communal dens by young-of-the-year snakes, and female mimicry. Hence, sexual conflict may have influenced important features of the mating system and behavioral ecology of these animals. Received: 8 May 2000 / Revised: 28 July 2000 / Accepted: 30 July 2000     

Immigration pattern and success in red squirrels

Summary We studied the characteristics of immigrants and the effects of immigration on reproductive activity and spacing behaviour in red squirrels living in high-quality woodlands. Male immigration peaked in spring, female immigration in autumn. There was no sex bias in dispersal distance of local recruits or in the proportion of male/female immigrants, but more subadults than adults immigrated on the study plots. Hence, hypotheses explaining sex-biased dispersal were irrelevant in explaining immigration patterns in our study populations. Immigrant females were not in breeding condition, nor had they produced a litter prior to immigration. Hence breeding dispersal did not occur. Red squirrels are promiscuous, and females defend intrasexual territories while males have overlapping home ranges with a dominance hierarchy (Wauters et al. 1990; Wauters and Dhondt 1992). Site fidelity is very important to reproductive success and most parents still have a high residual reproductive value after having produced a litter. Under such circumstances, the resident fitness hypothesis (RFH; Anderson 1989) predicts that parents can benefit by forcing emigration of offspring if the latter are likely to find nearby vacancies. The settlement pattern of successful immigrants, which had a higher probability of becoming established when they had high body mass and when they were settling in plots with reduced intrasexual competition, agreed with the RFH and with the proximate dispersal mechanism suggested by Gliwicz (1992), that dispersal tendency in both sexes depends on the degree of intrasexual competition under local conditions. The fact that close inbreeding was never observed could indicate that random immigration of both sexes, within the social environment of a partly territorial, relatively long-lived species, has evolved not only to reduce competition for resources between parents and offspring but also as an inbreeding avoidance mechanism. Correspondence to: L. Wauters