Matrilineal kin clusters and their effect on reproductive success in the field vole Microtus agrestis

Kin dusters of philopatric females are thought to form the basicsocial organization of breeding females in various vole species.The spatial association between young breeding females and theirmothers was studied in five 2500–m² enclosures in naturalhabitats during mid- and late breeding season in summer 1992.The mothers (founder females) were introduced into the enclosuresin die beginning of the study in June, whereas the young breedingfemales were born within the enclosures. Matrilineal kinshipwas determined by using radionudides. In July-August, 58% ofyoung breeding females were philopatric, but 82% of the mother-daughterassociations broke up until August-September. Of the young femalesthat started to breed in August-September, only 12% were philopatricThus the kin clusters found were short Hved and few in numberduring the late breeding season. The reduction in the incidenceof the kin clusters coincided with an increase of populationdensity. The mean reproductive success of females breeding inkin clusters was better than that of those breeding separately.In addition, among the nonphilopatric females there was a positive,significant correlation between reproductive success and thedistance to the nearest unrelated founder female. Such correlationwas not found among the philopatric females.     

Neonatal manipulation of oxytocin influences female reproductive behavior and success

During early neonatal development, oxytocin (OT) may influence the expression of adult behavior and physiology. Here we test the prediction that early postnatal exposure to OT or an oxytocin antagonist (OTA) can affect the subsequent expression of sexual receptivity and reproductive success of females. To test this hypothesis, female prairie voles (Microtus ochrogaster) received one of four treatments within 24 h of birth. Three groups received an intraperitoneal injection of OT, OTA, or isotonic saline. A fourth group was handled, but not injected. Around 75 days of age, females were paired with sexually experienced males for 72 h and sexual activity was recorded. Treatment had no effect on the probability of mating. Injection, regardless of treatment, reduced latency to mate compared with handled controls. OT and OTA treatment decreased mating bout frequency compared to saline and handled controls, while OTA treatment increased reproductive success, probability of successfully producing a litter. The results suggest that neonatally OT, both endogenous and exogenous, can affect the expression of adult female reproductive activity and that blocking the effects of endogenous OT during neonatal development can affect female reproductive success. Finally, the results suggest that a number of aspects of reproduction are regulated by OT during the postnatal period, but that the mechanism of action may differ depending upon the reproductive activity.     

Female philopatry and male-biased dispersal in a direct-developing salamander, Plethodon cinereus

The local resource competition hypothesis and the local mate competition hypothesis were developed based on avian and mammalian systems to explain sex-biased dispersal. Most avian species show a female bias in dispersal, ostensibly due to resource defence, and most mammals show a male bias, ostensibly due to male-male competition. These findings confound phylogeny with mating strategy; little is known about sex-biased dispersal in other taxa. Resource defence and male-male competition are both intense in Plethodon cinereus, a direct-developing salamander, so we tested whether sex-biased dispersal in this amphibian is consistent with the local resource competition hypothesis (female-biased) or the local mate competition hypothesis (male-biased). Using fine-scale genetic spatial autocorrelation analyses, we found that females were philopatric, showing significant positive genetic structure in the shortest distance classes, with stronger patterns apparent when only territorial females were tested. Males showed no spatial genetic structure over the shortest distances. Mark-recapture observations of P. cinereus over 5 years were consistent with the genetic data: males dispersed farther than females during natal dispersal and 44% of females were recaptured within 1 m of their juvenile locations. We conclude that, in this population of a direct-developing amphibian, females are philopatric and dispersal is male-biased, consistent with the local mate competition hypothesis.     

Mate competition and reproductive correlates of female dispersal in a polygynous primate species (Rhinopithecus roxellana)

Different mating systems in group-living animals have characteristic behavioral correlates that are primarily related to mate competition. Mate competition may push individuals to selectively make dispersal decisions for the purpose of maximizing of opportunities for reproduction. The Sichuan snub-nosed monkey (Rhinopithecus roxellana) is a polygynous primate species endemic to China. We provide the first data on female dispersal in a free-ranging group of R. roxellana in the Zhouzhi National Nature Reserve, Qinling Mountains, China. Both adult and subadult female dispersal occurred. Immigration/emigration rates of adult females are higher than those of subadult females. Mate competition is one apparent driving force behind adult female dispersal, and inbreeding avoidance is the possible proximate factor influencing subadult female dispersal. Adult female R. roxellana employ various reproductive strategies related to dispersal, which may increase their reproductive success.     

Offspring growth, survival and reproductive success in the bank vole: a litter size manipulation experiment

To estimate the optimality of brood size, it is essential to study the effects of brood size manipulation on offspring survival and reproductive success. Moreover, testing the generality of the hypothesis of reproductive costs requires experimental data from a diversity of organisms. Here I present data on the growth, survival and reproductive success of bank vole Clethrionomys glareolus individuals from manipulated litters. Furthermore, the survival of mothers whose litter size was manipulated was studied. At weaning, the mean weight of pups from enlarged litters was lower and from reduced litters higher compared to control litters. After winter, at the start of the breeding season, individuals from enlarged litters, especially males, were still lighter than individuals from the other two treatments. Litter enlargements did not increase the number of reproducing female offspring per mother, nor did the litter sizes of female offspring differ between treatments. There were no differences between treatments in winter survival of offspring after weaning, but among female offspring, weaning weight explained the survival probabilities over winter. A higher weight of females at winter determined the probability of starting to reproduce in spring. The survival of mothers did not seem to be influenced by litter manipulation performed the previous year. According to the results, mothers nursing enlarged or reduced litters do not gain any fitness benefits in terms of number of offspring surviving to breeding. The results are consistent with the majority of experiments conducted in birds, which have found costs of enlarged brood appearing as offspring trade-offs rather than parent trade-offs. Received: 14 December 1997 / Accepted: 1 March 1998     

Role of male dispersal in the evolution of female altruistic behavior in viscous populations

A computer simulation was conducted to examine the effect of differential dispersal of sexes on the evolution of altruism in viscous populations. First, a basic model, which was regarded as a purely viscous population model, was constructed. The model was assumed to be the same as the simulation model of Wilson etal. (1992), except that it assumed sexual reproduction and that only females show altruistic behavior toward females. For the basic model, altruism could not evolve when b/Nc, where b is the benefit of the altruism to the recipient, c is the cost to the altruist, and N is the number of interacting neighbors. The male dispersal model I assumed that females disperse to nine neighboring sites including the natal site, but males disperse to eight sites farther than females do. For this model, altruistic alleles could evolve when b/N was equal to c or b/N was slightly smaller than c only when the male dispersal distance was slightly larger than those of females. The male dispersal model II assumed that the male dispersal distance follows a normal probability distribution. The Vole model was based on actual data of the gray-sided vole, Clethironomys rufocanus bedfordiae, whose frequency distribution of dispersal distance was similar to a normal distribution. For these models, altruism could evolve under the condition that b/N was slightly smaller than c when the dispersal distances of males were larger than those of females. The results indicate that the differential dispersal of sexes, in which females are philopatric and males disperse farther than females, can somewhat increase the probability of spreading altruistic alleles in viscous populations.     

Understanding the population genetic structure of Gleditsia triacanthos L.: seed dispersal and variation in female reproductive success

Fine-scale genetic structuring is influenced by a variety of ecological factors and can directly affect the evolutionary dynamics of plant populations by influencing effective population size and patterns of viability selection. In many plant species, genetic structuring within populations may result from highly localized patterns of seed dispersal around maternal plants or by the correlated dispersal and recruitment of siblings from the same fruit. This fine-scale genetic structuring may be enhanced if female parents vary significantly in their reproductive success. To test these hypotheses, we used genetic data from 17 allozyme loci and a maximum-likelihood, ‘maternity-analysis’ model to estimate individual female fertilities for maternal trees across a large number of naturally established seedlings and saplings in two populations of Gleditsia triacanthos L. (Leguminosae). Maximum-likelihood fertility estimates showed that the three highest fertility females accounted for 58% of the 313 progeny at the first site and 46% of the 651 progeny at the second site, whereas 18 of 35 and 16 of 34 females, respectively, had fertility estimates that did not exceed 1%. Additional analyses of the second site found individual female fertility to vary significantly both within and among juvenile age classes. Female fertility at the first site was weakly correlated with maternal tree size and spatial location relative to the open, old-field portions of the population, where the great majority of seedlings and saplings were growing, but no such correlations were found at the second site. Estimates of realized seed dispersal distances indicated that dispersal was highly localized at the first site, but was nearly random at the second site, possibly reflecting differences between the two sites in the behaviour of animal dispersers. The combined estimates of seed dispersal patterns and fertility variation are sufficient to explain previously described patterns of significant fine-scale spatial genetic structure in these two populations. In general, our results demonstrate that effective seed dispersal distributions may vary significantly from population to population of a species due to the unpredictable behaviour of secondary dispersers. Consequently, the effects of seed dispersal on realized fine-scale genetic structure may also be relatively unpredictable.     

Determinants of reproductive success in female Columbian ground squirrels

Summary We studied the reproductive success of female Columbian ground squirrels (Spermophilus columbianus) in southwestern Alberta for nine years. We defined reproductive success as the number of offspring surviving their first hibernation, classified as yearlings. The number of weaned juveniles explained one third of the variance in number of yearlings at emergence from their first hibernation the following spring, and much of the variance in individual reproductive success originated after weaning. Weight of adult females at emergence from hibernation was correlated with annual reproductive success. The mother's survival beyond weaning and the subsequent winter's snow accumulation had positive effects on annual reproductive success, whereas population density and summer temperature had negative effects. We found no effects on annual reproductive success of date of litter emergence, weight at emergence as a yearling, presence or absence of adult kin, distance from the natal site, location within the study area, winter temperature or summer precipitation. Age of first breeding did not affect lifetime reproductive success, which ranged from 0 to 19 yearlings produced over a lifetime. The greatest source of variation in lifetime reproductive success for females surviving to breeding age was offspring survival, followed by reproductive lifespan.     

Effective gene dispersal and female reproductive success in Mediterranean maritime pine (Pinus pinaster Aiton)

Understanding population-scale processes that affect allele frequency changes across generations is a long-standing interest in genetic, ecological and evolutionary research. In particular, individual differences in female reproductive success and the spatial scale of gene flow considerably affect evolutionary change and patterns of local selection. In this study, a recently developed maximum-likelihood (ML) method based on established offspring, the Seedling Neighbourhood Model , was applied and exponentially shaped dispersal kernels were fitted to both genetic and ecological data in a widespread Mediterranean pine, Pinus pinaster Aiton. The distribution of female reproductive success in P. pinaster was very skewed (about 10% of trees mothered 50% of offspring) and significant positive female selection gradients for diameter (γ = 0.7293) and cone crop (γ = 0.4524) were found. The selective advantage of offspring mothered by bigger trees could be due to better-quality seeds. These seeds may show more resilience to severe summer droughts and microsite variation related to water and nutrient availability. Both approaches, ecological and of parentage, consistently showed a long-distance dispersal component in saplings that was not found in dispersal kernels based on seed shadows, highlighting the importance of Janzen-Connell effects and microenvironmental variation for survival at early stages of establishment in this Mediterranean key forest tree.     

Seasonal variation in reproductive success and post-nuptial moult of blue tits in southern Europe: an experimental study

Post-nuptial moult and reproductive success were studied in relation to timing of breeding in blue tits, Parus caeruleus, breeding in southern Europe. A group of experimentally delayed pairs was created by removing first clutches, thereby inducing late repeat clutches. Reproductive success and post-nuptial moult of delayed pairs were compared with both control pairs that bred early and unmanipulated late-breeding pairs. Delayed pairs fledged fewer young and with a lower body mass than control pairs. However, the number of fledged young and fledgling mass did not differ between delayed and late-breeding pairs. These results were more consistent with the date hypothesis, and it is concluded that the timing of breeding and reproductive success may be causally related in the blue tit. This study reveals a harmful effect of relaying on female body mass at the end of the nestling period. Therefore, females apparently pay the costs of relaying, since a reduction in body mass during the nestling period may be accompanied by a lowered survival probability. Delayed and late-breeding males often began moulting while still feeding young, but neither control males nor females from the three study groups did so. These results support the view that timing-related energy constraints on breeding may be important causes of a seasonal decline in reproductive success at different latitudes. Received: 15 March 1999 / Accepted: 19 July 1999     

Seasonal changes in components of male and female reproductive success in Raphanus sativus L. (Brassicaceae)

Summary To document seasonal changes in the reproductive behavior of the perfect-flowered, self-incompatible mustard, Raphanus sativus L., we monitored individual survival, flower and fruit production among 58 individuals in a California population over six census dates (cohorts). Population size declined dramatically and mean individual levels of fruit set changed significantly between cohorts. The frequency distribution of flower and fruit production became increasingly skewed over the first four cohorts. The phenotypic maleness of individuals, a standardized measure of phenotypic gender, oscillated during the reproductive season, peaking in the third and fourth cohorts. We calculated a simple estimate of expected male reproductive success of each plant (the number of fruits sired on conspecifics); this estimate was a function of an individual's flower production and the fruit production of its potential mates in our sampled population. Mean expected male success did not differ significantly among cohorts; expected male success per flower did, however, change significantly among cohorts. Among individuals within each cohort, maternal fruit production and expected male success were both positively correlated with flower production throughout the season. Spearman rank correlation coefficients indicate that the strength of these associations, however, changed during the season. Linear regressions of transformed variables indicated that the shape of several fitness functions also changed over time. In addition, the amount of variation in maternal or expected paternal success explained by flower production declined over the first four cohorts. If typical of wild populations, these temporal changes in these functions suggest that measurements of the intensity of phenotypic selection on flower production will depend on when and how fitness is measured in natural populations.     

Costs and benefits of female-biased natal philopatry in the common goldeneye

Sex-biased natal dispersal in long-lived species may resultin interactions between parents and mature young of the philopatricsex. To investigate the evolutionary basis of natal philopatryin a noncooperative species, the common goldeneye Bucephalaclangula, we studied possible costs and benefits of simultaneousbreeding of females and philopatric daughters. We did not find any fitness consequences of a daughter's breeding on their mother'sbreeding in terms of nest-site selection, body weight, clutchsize, hatching date, or hatching success. Our results, therefore,did not support the assumption of the local resource competitionhypothesis, that the natally philopatric sex should be morecostly to a breeding parent. As possible benefits for daughters returning to their natal area, we tested inheritance of nestsites from mothers and explored whether daughters utilize thepresence of their mother by parasitically sneaking into hermother's nest. Daughters' nest-site selection was not associatedwith the presence of their mothers. A comparison between daughtersand control females revealed that daughters chose their nestsite closer to their natal nest than expected by nest-siteavailability alone. Daughters could not expect to inherit anest site from their mother, and we did not find other indicationsof cooperation between relatives either. The mother's clutchsize did not increase in the year breeding with the daughter, indicating daughters do not parasitize their mother's nest.We suggest that benefits such as decreased nest predation riskassociated with nesting close to the natal nest site may beimportant in the natal philopatric behavior of the species.     

Height and reproductive success in a cohort of british men

Two recent studies have shown a relationship between male height and number of offspring in contemporary developed-world populations. One of them argues as a result that directional selection for male tallness is both positive and unconstrained. This paper uses data from a large and socially representative national cohort of men who were born in Britain in March 1958. Taller men were less likely to be childless than shorter ones. They did not have a greater mean number of children. If anything, the pattern was the reverse, since men from higher socioeconomic groups tended to be taller and also to have smaller families. However, clear evidence was found that men who were taller than average were more likely to find a long-term partner, and also more likely to have several different long-term partners. This confirms the finding that tall men are considered more attractive and suggests that, in a noncontracepting environment, they would have more children. There is also evidence of stabilizing selection, since extremely tall men had an excess of health problems and an increased likelihood of childlessness. The conclusion is that male tallness has been selected for in recent human evolution but has been constrained by developmental factors and stabilizing selection on the extremely tall. The National Child Development Study is carried out by the researchers of the Centre for Longitudinal Studies, Institute of Education, London. The data are housed at the Data Archive of the University of Essex, from where they were obtained under license for the present study. I am grateful to Paul Preece for his assistance with data processing, and to Benjamin Campbell and two anonymous referees for invaluable advice which has, I hope, improved the quality of this paper. Daniel Nettle is lecturer in biological psychology at the Open University. With a first degree in psychology and a Ph.D. in biological anthropology, his work has been concerned with the application of evolutionary models to such topics as language (e.g., Linguistic Diversity, Oxford University Press, 1999) and individual differences (Strong Imagination, Oxford University Press, 2001).     

Assessment of individual and conspecific reproductive success as determinants of breeding dispersal of female tree swallows: A capture–recapture approach

Breeding dispersal is a key process of population structure and dynamics and is often triggered by an individual's breeding failure. In both colonial and territorial birds, reproductive success of conspecifics (RSc) can also lead individuals to change breeding sites after a failure on a site. Yet, few studies have simultaneously investigated the independent contribution of individual reproductive success (RSi) and of RSc on dispersal decision. Here, we develop a modeling framework to disentangle the effects of RSi and RSc on demographic parameters, while accounting for imperfect individual detection and other confounding factors such as age or dispersal behavior in the previous year. Using a 10‐year capture–recapture dataset composed of 1,595 banded tree swallows, we assessed the effects of nonmanipulated RSi and RSc on female breeding dispersal in this semicolonial passerine. Dispersal was strongly driven by RSi, but not by RSc. Unsuccessful females were 9.5–2.5 times more likely to disperse than successful ones, depending if they had dispersed or not in the previous year, respectively. Unsuccessful females were also three times less likely to be detected than successful ones. Contrary to theoretical and empirical studies, RSc did not drive the decision to disperse but influenced the selection of the following breeding site once dispersal had been initiated. Because detection of individuals was driven by RSi, which was positively correlated to RSc, assuming a perfect detection as in previous studies may have lead us to conclude that RSc affected dispersal patterns, yet our approach corrected for this bias. Overall, our results suggest that the value and use of RSc as public information to guide dispersal decisions are likely dictated by multiple ecological determinants, such as landscape structure and extent, if this cue is indeed used.     

Effects of floral herbivory on foraging behaviour of bumblebees and female reproductive success in Pedicularis gruina (Orobanchaceae)

The effects of floral herbivores on floral traits may result in alterations in pollinator foraging behaviour and subsequently influence plant reproductive success. Fed-upon plants may have evolved mechanisms to compensate for herbivore-related decreased fecundity. We conducted a series of field experiments to determine the relative contribution of floral herbivores and pollinators to female reproductive success in an alpine herb, Pedicularis gruina, in two natural populations over two consecutive years. Experimental manipulations included bagging, hand supplemental, geitonogamous pollination, and simulated floral herbivory. Bumblebees not only avoided damaged flowers and plants but also decreased successive visits of flowers in damaged plants, and the latter may reduce the level of geitonogamy. Although seed set per fruit within damaged plants was higher than that in intact plants, total seed number in damaged plants was less than that in intact plants, since floral herbivory-mediated pollinator limitation led to a sharp reduction of fruit set. Overall, the results suggest that resource reallocation within inflorescences of damaged plants may partially compensate for a reduction in seed production. Additionally, a novel finding was the decrease in successive within-plant bumblebee visits following floral herbivory. This may increase seed quantity and quality of P. gruina since self-compatible species exhibit inbreeding depression. The patterns of compensation of herbivory and its consequences reported in this study give an insight into the combined effects of interactions between floral herbivory and pollination on plant reproductive fitness.     

Inter-locality variation in reproductive success of the citril finch Serinus citrinella

Habitat quality is generally thought to affect breeding success. We tested this effect comparing differences in clutch size and reproductive success between citril finch Serinus citrinella sub-populations closely located (<5 km) but differing in habitat quality, within the Port del Comte mountain, in the Catalonian Pre-Pyrenees. We found that birds in the low quality area (Bofia) showed significantly lower hatching, breeding and nesting success than finches in the high quality area (Vansa). These differences in reproductive success fit well with recently found differences in citril finch body mass, fat score, diet, survival rate and speed of moult between these two localities.     

Age variation in a fluctuating population of the common vole

We analysed variation in age in a fluctuating population of the common vole (Microtus arvalis) in southern Moravia, Czech Republic, to test the assumption of the senescence hypothesis that the age of voles increases with increasing population density. Between 1996 and 1998, we monitored the demographic changes by snap-trapping and live-trapping in a field population passing through the increase, peak and decline phase of the population cycle. We used the eye lens mass method to determine the age of snap-trapped animals and those that died in live-traps. The average age of winter males was clearly higher after the peak phase breeding season than before it. No such phase-dependent shift in age, however, was observed in the female component. Male age continued to increase from autumn to spring over the pre-peak winter, and the highest age was in spring of the peak phase year. However, after the peak phase breeding season the highest age was achieved in winter, with the decline phase males during the next spring tending to be younger. The average age of females in spring populations was always lower than in winter populations. The average age of voles from live-traps was always higher than voles from snap-traps, particularly in winter and spring populations, suggesting the presence of senescent animals. Although the density-dependent changes in age are consistent with those observed for other voles, they provide only weak evidence that population cycles in the common vole are accompanied by pronounced shifts in individual age, particularly in female voles.Due to an error in the citation line, this revised PDF (published in December 2003) deviates from the printed version, and is the correct and authoritative version of the paper.     

Sex-related spatial kin structure in a spring population of grey-sided voles Clethrionomys rufocanus as revealed by mitochondrial and microsatellite DNA analyses

Polymerase chain reaction-directed mitochondria (mt) and microsatellite DNA analyses were performed to examine the kin structure in a spring population of grey-sided voles Clethrionomys rufocanus in Hokkaido, Japan. The spatial distribution of 81 voles in a trapping grid (about 1 ha) was estimated by using the catch-mark-release method. DNA samples were extracted from the toes clipped for individual identification. Maternal lineages of voles were unequivocally determined by the mtDNA haplotypes, as identified by nucleotide sequencing of the control region. Relatedness between individuals was estimated based on the genotype and allele frequencies at several microsatellite loci. Although the distribution of voles was uniform within the grid, neighbouring females were frequently from the same maternal lineage. Relatedness values between females correlated negatively with geographical distances. Combination of the two molecular markers revealed four clusters of closely related, matrilineal females in the population, whereas no such cluster was apparent in males. The present study first demonstrated a sex-related spatial kin structure in a natural population of the grey-sided vole.