Adaptive significance of maternal induction of density-dependent phenotypes

Density has been demonstrated to impact life history traits such as growth, fecundity and survival. Some authors have proposed that morphological and behavioral traits have evolved in response to density conditions. To escape the adverse effect of density, individuals can either adapt to crowding or avoid crowding by dispersing. The aim of this work is to study the interplay between local adaptation and dispersal in four populations of the common lizard, Lacerta vivipara, where densities of both the maternal and juvenile environment have been experimentally manipulated. Density was decreased in the spring by removing a quarter of the population at two sites and was un-manipulated in two other sites. One month later, we caught some pregnant females and kept them in the laboratory until parturition. To manipulate density of postnatal neonates and juveniles, we divided each clutch into two, and released half of the juveniles either in a reduced density site or in a control one. We then recaptured individuals a year after release and recorded their size and weight. When density was reduced, females increased their clutch size, but produced offspring of lower body condition than in the control sites. The conspicuous ventral color of females was likewise increased when density was reduced. However, offspring growth rate, local survival and dispersal were not influenced by maternal density. Juvenile females released in the reduced-density site had lower survival rate than those released in the control density site. Contrary to expectations, offspring dispersal was significantly higher at the reduced compared to control density sites. There was no interaction between maternal density habitat and the juvenile release habitat indicating that maternal effects did not influence juvenile life history traits in a different way according to the level of density. Moreover, clutch size and offspring size had no effect on juvenile growth or survival.     

Factors influencing and consequences of breeding dispersal and habitat choice in female grey seals (<Emphasis Type="Italic">Halichoerus grypus</Emphasis>) on Sable Island,Nova Scotia

Selection of breeding location can influence reproductive success and fitness. Breeding dispersal links habitat use and reproduction. This study investigated factors affecting breeding dispersal and its reproductive consequences in grey seals (Halichoerus grypus) on Sable Island, Nova Scotia. Breeding dispersal distance was determined in 692 individually marked, known-age female grey seals observed from 2004 to 2014. We used generalized linear mixed-effects models to test hypotheses concerning environmental and demographic factors influencing breeding dispersal distance and the consequences of dispersal distance on offspring weaning mass. Grey seal females rarely exhibited fidelity to previous breeding sites. Median dispersal distance between years was 5.1 km. Only 2.9% of females returned to a previous breeding site. Breeding dispersal distance was affected by parity and density, but effects were small and are presumably of no biological significance. Variation in dispersal distance among adult females was large. Dispersal distance had no significant influence on offspring weaning mass; however, as previously found, pup sex and maternal age did. Although breeding location was not important, heavier pups were born in habitats with no tidal or storm-surge influence indicating that breeding habitat type did influence offspring size at weaning. The lack of site fidelity in grey seals on Sable Island is associated with an unpredictable and changing landscape (sand dunes) that could make it difficult for females to locate previous breeding locations. Although breeding location within habitat type had small consequences on offspring weaning mass, we detected no evidence that breeding site selection within the habitat had consequences to females.     

Interacting maternal and spatial cues influence natal – dispersal out of social groups

Proximate cues for animal dispersal are complex and varied. Multiple cues may provide information about different aspects of habitat quality, and these aspects may interact with each other, as well as with population density in different ways. We examined how individuals incorporate multiple cues in their decisions to emigrate and immigrate in the colonial orb‐weaving spider, Cyrtophora citricola. We manipulated maternal feeding as a cue for prey abundance and measured the size of the maternal web, which provides a limited space for philopatric offspring and a second potential dispersal cue. In addition, we recorded all immigration events to determine dispersal distances and the cues juveniles may use in settlement. Dispersal increased when mothers were poorly fed, web sizes were small and clutch sizes were large. In addition to these overall effects, maternal feeding also interacted with web size, indicating that offspring from well‐fed mothers were more tolerant of high sibling densities. We also detected a threshold for the effect of clutch size on dispersal for the first egg sac: below 20 offspring, there was no effect of clutch size, but dispersal increased with clutch size for larger clutches. Dispersal distances were often short, and immigrants preferred sheltered trees and those occupied by adult females. Dispersal not only depended on multiple cues, but these cues interacted, and the importance of web size suggested that saturation of the natal web might force dispersal, at least for spiders with poorly‐fed mothers. How one aspect of habitat quality influences dispersal can therefore depend on the state of other aspects of habitat quality. In particular, some natal resources, such as a nest or territory, may become saturated and limit group size, but this limit will also depend on other factors, such as prey availability.     

Population consequences of maternal effects: sex‐bias in egg‐laying order facilitates divergence in sexual dimorphism between bird populations

Abstract When costs and benefits of raising sons and daughters differ between environments, parents may be selected to modify their investment into male and female offspring. In two recently colonized environments, breeding female house finches (Carpodacus mexicanus) modified the sex and growth of their offspring in relation to the order in which eggs were laid in a clutch. Here we show that, in both populations, these maternal effects strongly biased frequency distribution of tarsus size of fully grown males and females and ultimately produced population divergence in this trait. Although in each population, male and female offspring show a wide range of growth patterns, maternal modifications of sex‐ratio in relation to egg‐laying order resulted in under‐representation of the morphologies that were selected against and over‐representation of morphologies that were favoured by the local selection on juveniles. The result of these maternal adjustments was fast phenotypic change in sexual size dimorphism within and between populations. Maternal manipulations of offspring morphologies may be especially important at the initial stages of population establishment in the novel environments and may have facilitated recent colonization of much of North America by the house finch.     

蜡皮蜥的两性异形和繁殖输出

为研究蜡皮蜥(Leiolepis reevesii)两性异形和繁殖输出,于2002、2003年4月下旬从海南乐东一种群捕获423头蜡皮蜥。经检测得到繁殖雌体的最小体长为89．0mm,据此判定≥89．0mm的个体为性成熟。研究结果表明：①蜡皮蜥具有两性异形,雄性大于雌性且具有较大的头部。成体雄性头长和头宽随体长的增长速率大于雌性,幼体头长和头宽随体长的增长速率无显著的两性差异。以性别和年龄(成、幼体)为因子的双因子ANOVA比较两性头长和头宽与体长的回归剩余值发现,雄性头部大于雌性,幼体头部相对大于成体。②饲养于实验室的母体中有42头于2002、2003年5月22日～7月16日产出正常卵,这些繁殖雌体具有年产多窝卵的潜力。窝卵数和卵重的变异系数分别为0．18和0．13,前者变异度大于后者。窝卵数、窝卵重和卵重均与母体体长无关。卵重与相对生育力之间无显著的负相关性,表明蜡皮蜥缺乏卵数量与卵大小之间的权衡。相对窝卵重与母体体长呈显著的负相关,表明较小的母体具有相对较大的繁殖输出。因雌体繁殖会滞缓其生长,小母体具有相对较大的繁殖输出,至少部分地解释了雌性蜡皮蜥的成体为什么个体较小。     

INTERACTIVE EFFECTS OF OFFSPRING SIZE AND TIMING OF REPRODUCTION ON OFFSPRING REPRODUCTION: EXPERIMENTAL,MATERNAL, AND QUANTITATIVE GENETIC ASPECTS

We demonstrate that egg size in side-blotched lizards is heritable (parent-offspring regressions) and thus will respond to natural selection. Because our estimate of heritability is derived from free-ranging lizards, it is useful for predicting evolutionary response to selection in wild populations. Moreover, our estimate for the heritability of egg size is not likely to be confounded by nongenetic maternal effects that might arise from egg size per se because we estimate a significant parent-offspring correlation for egg size in the face of dramatic experimental manipulation of yolk volume of the egg. Furthermore, we also demonstrate a significant correlation between egg size of the female parent and clutch size of her offspring. Because this correlation is not related to experimentally induced maternal effects, we suggest that it is indicative of a genetic correlation between egg size and clutch size. We synthesize our results from genetic analyses of the trade-off between egg size and clutch size with previously published experiments that document the mechanistic basis of this trade-off. Experimental manipulation of yolk volume has no effect on offspring reproductive traits such as egg size, clutch size, size at maturity, or oviposition date. However, egg size was related to offspring survival during adult phases of the life history. We partitioned survival of offspring during the adult phase of the life history into (1) survival of offspring from winter emergence to the production of the first clutch (i.e., the vitellogenic phase of the first clutch), and (2) survival of the offspring from the production of the first clutch to the end of the reproductive season. Offspring from the first clutch of the reproductive season in the previous year had higher survival during vitellogenesis of their first clutch if these offspring came from small eggs. We did not observe selection during these prelaying phases of adulthood for offspring from later clutches. However, we did find that later clutch offspring from large eggs had the highest survival over the first season of reproduction. The differences in selection on adult survival arising from maternal effects would reinforce previously documented selection that favors the production of small offspring early in the season and large offspring later in the season—a seasonal shift in maternal provisioning. We also report on a significant parent-offspring correlation in lay date and thus significant heritable variation in lay date. We can rule out the possibility of yolk volume as a confounding maternal effect—experimental manipulation of yolk volume has no effect on lay date of offspring. However, we cannot distinguish between genetic effects (i.e., heritable) and nongenetic maternal effects acting on lay date that arise from the maternal trait lay date per se (or other unidentified maternal traits). Nevertheless, we demonstrate how the timing of female reproduction (e.g., date of oviposition and date of hatching) affect reproductive attributes of offspring. Notably, we find that date of hatching has effects on body size at maturity and fecundity of offspring from later clutches. We did not detect comparable effects of lay date on offspring from the first clutch.     

Persistent maternal effects on juvenile survival in North American red squirrels

Maternal effects can have lasting fitness consequences for offspring, but these effects are often difficult to disentangle from associated responses in offspring traits. We studied persistent maternal effects on offspring survival in North American red squirrels (Tamiasciurus hudsonicus) by manipulating maternal nutrition without altering the post-emergent nutritional environment experienced by offspring. This was accomplished by providing supplemental food to reproductive females over winter and during reproduction, but removing the supplemental food from the system prior to juvenile emergence. We then monitored juvenile dispersal, settlement and survival from birth to 1 year of age. Juveniles from supplemented mothers experienced persistent and magnifying survival advantages over juveniles from control mothers long after supplemental food was removed. These maternal effects on survival persisted, despite no observable effect on traits normally associated with high offspring quality, such as body size, dispersal distance or territory quality. However, supplemented mothers did provide their juveniles an early start by breeding an average of 18 days earlier than control mothers, which may explain the persistent survival advantages their juveniles experienced.     

Larger female fish contribute disproportionately more to self-replenishment

While chance events, oceanography and selective pressures inject stochasticity into the replenishment of marine populations with dispersing life stages, some determinism may arise as a result of characteristics of breeding individuals. It is well known that larger females have higher fecundity, and recent laboratory studies have shown that maternal traits such as age and size can be positively associated with offspring growth, size and survival. Whether such fecundity and maternal effects translate into higher recruitment in marine populations remains largely unanswered. We studied a population of Amphiprion chrysopterus (orange-fin anemonefish) in Moorea, French Polynesia, to test whether maternal size influenced the degree of self-recruitment on the island through body size-fecundity and/or additional size-related maternal effects of offspring. We non-lethally sampled 378 adult and young juveniles at Moorea, and, through parentage analysis, identified the mothers of 27 self-recruits (SRs) out of 101 recruits sampled. We also identified the sites occupied by each mother of an SR and, taking into account variation in maternal size among sites, we found that females that produced SRs were significantly larger than those that did not (approx. 7% greater total length, approx. 20% greater biomass). Our analyses further reveal that the contribution of larger females to self-recruitment was significantly greater than expected on the basis of the relationship between body size and fecundity, indicating that there were important maternal effects of female size on traits of their offspring. These results show, for the first time in a natural population, that larger female fish contribute more to local replenishment (self-recruitment) and, more importantly, that size-specific fecundity alone could not explain the disparity.     

Stress and body condition as prenatal and postnatal determinants of dispersal in the common lizard (Lacerta vivipara)

Dispersal is a complex phenomenon affected by multiple factors. Among the factors that influence dispersal in the common lizard (Lacerta vivipara), poor maternal body condition and stress are known to decrease dispersal propensity of juveniles. But the effect of individual factors on dispersal could change when several of them act concurrently or at different developmental stages. Prenatal factors can affect clutch and/or juvenile characteristics that later affect dispersal. Postnatal influences are mainly exerted on juvenile dispersal behavior. We investigated the role of body condition and stress on dispersal at a prenatal and a postnatal stage. Stress was mimicked by experimentally increasing corticosterone levels in pregnant females and recently born juveniles. We considered (1). the influence of maternal body condition and prenatal corticosterone treatment on clutch, juvenile characteristics and on dispersal behavior and (2). the influence of juvenile body condition and postnatal corticosterone treatment on juvenile dispersal behavior. There was an interaction between maternal condition and prenatal corticosterone treatment on juvenile dispersal. Dispersal decreased with maternal corticosterone increase only in juveniles from the more corpulent females, while it increased with juvenile body condition. Good maternal body condition affected clutch and juvenile characteristics favoring dispersal, while elevation of corticosterone level (stress) exerted the opposite effect. Juvenile body condition favored dispersal, while there was no effect of postnatal corticosterone treatment on juvenile dispersal propensity.     

浙江丽水中国石龙子的食性、两性异形和雌性繁殖

丽水分布的中国石龙子（Ｅｕｍｅｃｅｓ ｃｈｉｎｅｎｓｉｓ）摄入的食物均为无脊椎动物,分别属于环节、软体和节肢动物,涉及３０余科,成体和幼体的食物生态位宽度分别为７．２６和６．６９,成体的幼体的食物生态们重叠度为０．５９。性成熟雄性个体大于雌体。成雄和幼体的头长和头宽随体长ＳＶＬ的增长速率大于成雌,成雄头长随ＳＶＬ的增长速度显著大于幼体,成雌和幼体的头长随ＳＶＬ的增长速率无显著差异。成雄头部大于成雌     

Parentage analysis detects cryptic precapture dispersal in a philopatric rodent

Locating birthplaces using genetic parentage determination can increase the precision and accuracy with which animal dispersal patterns are established. We re-analyse patterns of movement away from the birthplace as a function of time, sex and population density for a sample of 303 banner-tailed kangaroo rats, Dipodomys spectabilis. We located birth sites using a combination of likelihood-based parentage analysis with live-trapping of mothers during the breeding season. The results demonstrate that natal-breeding site distances are density dependent in this species; in particular, both sexes emigrate earlier in the year, and females disperse farther than males, at low population densities. Banner-tailed kangaroo rats were chosen as a study system because live-trapping easily detects maternal and offspring locations; nevertheless, parentage analysis reveals that some offspring evade early detection and move substantial distances before their first capture. In a few cases, the approach even detects dispersal out of the natal 'deme' prior to first capture. Parentage analysis confirms the extreme philopatry of both sexes but indicates that prior estimates of median dispersal distance were too low. For D. spectabilis, more accurate location of individual birthplaces clarifies patterns of sex bias and density dependence in dispersal, and may resolve apparent discrepancies between direct and indirect estimates of dispersal distance. For species in which mothers can be more reliably trapped than juveniles, using offspring genotypes to locate parents is a novel way that genetic techniques can contribute to the analysis of animal dispersal.     

Differential effects of offspring and maternal inbreeding on egg laying and offspring performance in the burying beetle Nicrophorus vespilloides

We investigate the effect of offspring and maternal inbreeding on maternal and offspring traits associated with early offspring fitness in the burying beetle Nicrophorus vespilloides. We conducted two experiments. In the first experiment, we manipulated maternal inbreeding only (keeping offspring outbred) by generating mothers that were outbred, moderately inbred or highly inbred. Meanwhile, in the second experiment, we manipulated offspring inbreeding only (keeping females outbred) by generating offspring that were outbred, moderately inbred or highly inbred. In both experiments, we monitored subsequent effects on breeding success (number of larvae), maternal traits (clutch size, delay until laying, laying skew, laying spread and egg size) and offspring traits (hatching success, larval survival, duration of larval development and average larval mass). Maternal inbreeding reduced breeding success, and this effect was mediated through lower hatching success and greater larval mortality. Furthermore, inbred mothers produced clutches where egg laying was less skewed towards the early part of laying than outbred females. This reduction in the skew in egg laying is beneficial for larval survival, suggesting that inbred females adjusted their laying patterns facultatively, thereby partially compensating for the detrimental effects of maternal inbreeding on offspring. Finally, we found evidence of a nonlinear effect of offspring inbreeding coefficient on number of larvae dispersing. Offspring inbreeding affected larval survival and larval development time but also unexpectedly affected maternal traits (clutch size and delay until laying), suggesting that females adjust clutch size and the delay until laying in response to being related to their mate.     

Role of breeding and natal movements in lifetime dispersal of a forest‐dwelling rodent

The lifetime movements of an individual determine the gene flow and invasion potential of the species. However, sex dependence of dispersal and selective pressures driving dispersal have gained much more attention than dispersal at different life and age stages. Natal dispersal is more common than dispersal between breeding attempts, but breeding dispersal may be promoted by resource availability and competition. Here, we utilize mark–recapture data on the nest‐box population of Siberian flying squirrels to analyze lifetime dispersal patterns. Natal dispersal means the distance between the natal nest and the nest used the following year, whereas breeding movements refer to the nest site changes between breeding attempts. The movement distances observed here were comparable to distances reported earlier from radio‐telemetry studies. Breeding movements did not contribute to lifetime dispersal distance and were not related to variation in food abundance or habitat patch size. Breeding movements of males were negatively, albeit not strongly, related to male population size. In females, breeding movement activity was low and was not related to previous breeding success or to competition between females for territories. Natal philopatry was linked to apparent death of a mother; that is, we did not find evidence for mothers bequeathing territories for offspring, like observed in some other rodent species. Our results give an example of a species in which breeding movements are not driven by environmental variability or nest site quality. Different evolutionary forces often operate in natal and breeding movements, and our study supports the view that juveniles are responsible for redistributing individuals within and between populations. This emphasizes the importance of knowledge on natal dispersal, if we want to understand consequences of movement ecology of the species at the population level.     

The population determines whether and how life-history traits vary between reproductive events in an insect with maternal care

The last reproductive event of a female is often associated with major changes in terms of both maternal and offspring life-history traits. However, the nature of these changes and the importance of population-specific environmental constraints in shaping their expression are difficult to predict and, as a consequence, poorly understood. Here, we investigated whether and how life-history traits vary between reproductive events and whether this variation is population-dependent in the European earwig Forficula auricularia. In this insect species, females produce up to two clutches during their lifetime and express extensive forms of maternal care. We conducted a common garden experiment, in which we measured 11 life-history traits of the first and second clutches of 132 females originating from three populations. Our results showed that clutch size was higher and the level of care expressed towards juveniles lower in second as compared to the first clutches in all three populations. In contrast, we found a population-specific effect on whether and how the reproductive event shaped juvenile quality and a trade-off between egg developmental time and female weight at hatching. Overall, these findings emphasise that the last reproductive event of a female entails both positive and negative effects on various life-history traits of the female herself and her clutch of juveniles. Moreover, our study stresses the importance of population idiosyncrasies on the expression and nature of such cohort-specific effects.     

Factors influencing offspring traits in the oviparous multi-clutched lizard,<Emphasis Type="Italic">Calotes versicolor</Emphasis> (Agamidae)

The determinants of offspring size and number in the tropical oviparous multi-clutched lizard,Calotes versicolor, were examined using both univariate and multivariate (path) analyses. InC. versicolor maternal snout-vent length (SVL) and body condition influence clutch mass and clutch size but have no significant influence on offspring size. The positive effect of maternal SVL and body condition on offspring number is counterbalanced by a negative effect of breeding time on egg mass. In fact, breeding time directly influences the offspring body mass and condition through variation in the egg mass. There is a trade-off between offspring mass and condition with offspring number, and breeding time influences both. Offspring hatched from the eggs of early (May–June) or mid (July–August) breeding periods invariably show lower mass and condition than those hatched from the eggs of late breeding season (September–October). Yet, there is no variation in offspring SVL among early, mid and late clutches. Thus, inC. versicolor offspring SVL is optimized while body mass and condition are not optimized.     

Differences in size between first and replacement clutches match the seasonal decline in single clutches in Tree Swallows Tachycineta bicolor

The seasonal decline in clutch size in birds can be a response to the environmentally conditioned decrease in prospects for offspring or a consequence of a lower physical ability of late‐breeding females. To find out which of the explanations apply in Tree Swallows Tachycineta bicolor, we assessed whether replacement clutch size in this species is affected by an individual female's ability to lay a certain number of eggs. To do this, we measured the decline in clutch size as a function of laying date between first and replacement clutches in individuals that re‐nested following natural failure, and compared this with the rate of decline in clutch size with laying date for Tree Swallows that laid only a single clutch in that season. Additionally, we assessed whether the clutch size and the rate of its seasonal decline varied across years. We accounted for the truncated and under‐dispersed nature of clutch size data by using a Bayesian approach in the analysis. We found little variation in the rate of clutch size decline across years at our breeding site. Accounting for this seasonal decline in clutch size, mean clutch size was similar between single‐time breeding females and those that laid replacement clutches, implying that the number of eggs laid on the second attempt by female Tree Swallows is determined by laying date, rather than by the female's physical ability to produce a clutch of a certain size.     

Inbreeding depression in an insect with maternal care: influences of family interactions,life stage and offspring sex

Although inbreeding is commonly known to depress individual fitness, the severity of inbreeding depression varies considerably across species. Among the factors contributing to this variation, family interactions, life stage and sex of offspring have been proposed, but their joint influence on inbreeding depression remains poorly understood. Here, we demonstrate that these three factors jointly shape inbreeding depression in the European earwig, Forficula auricularia. Using a series of cross‐breeding, split‐clutch and brood size manipulation experiments conducted over two generations, we first showed that sib mating (leading to inbred offspring) did not influence the reproductive success of earwig parents. Second, the presence of tending mothers and the strength of sibling competition (i.e. brood size) did not influence the expression of inbreeding depression in the inbred offspring. By contrast, our results revealed that inbreeding dramatically depressed the reproductive success of inbred adult male offspring, but only had little effect on the reproductive success of inbred adult female offspring. Overall, this study demonstrates limited effects of family interactions on inbreeding depression in this species and emphasizes the importance of disentangling effects of sib mating early and late during development to better understand the evolution of mating systems and population dynamics.     

Relationship between female mating strategy, litter success and offspring dispersal

The relationship between mating systems and dispersal has generally been studied at the population and species levels. It has hardly ever been investigated at the individual level, by studying the variations of mating and dispersal strategies between individuals. We investigated this relationship in a natural population of the common lizard ( Lacerta vivipara ). Assuming that dispersal has a genetic basis, juvenile dispersal would be expected to be more family-dependent in monoandrous litters than in polyandrous litters. The opposite pattern was observed. Thus, maternal effects and/or litter effects play a greater role than genetic determinism in shaping the dispersal phenotype of juveniles. Moreover, the relationship between female mating strategy and offspring dispersal depended on litter success, in a way consistent with an influence of mother-offspring competition. Such a link between mating and dispersal strategies of individuals may have major repercussions for the way we consider the roles of these processes in population functioning.     

Sex biased natal dispersal in a closed, saturated population of Seychelles warblers Acrocephalus sechellensis

The distances that individuals disperse, from their natal site to the site of first breeding and between breeding sites, have important consequences for the dynamics and genetic structure of a population. Nearly all previous studies on dispersal have the problem that, because the study area encompassed only a part of the population, emigration may have been confounded with mortality. As a result long-distance dispersers may have been overlooked and dispersal data biased towards short distances. By studying a virtually closed population of Seychelles warblers Acrocephalus sechellensis we obtained almost unbiased results on several aspects of dispersal. As in the majority of other avian species, natal dispersal distance was female biased in the Seychelles warbler. Female offspring also forayed further from the natal territory in search of breeding vacancies than male offspring. The sex bias in natal dispersal distance did, however, depend on local breeding density. In males, dispersal distance decreased as the number of territories bordering the natal territory increased, while in females, dispersal distance did not vary with local density. Dispersal by breeders was rare and, unlike in most species, distances did not differ between the sexes. We argue that our results favour the idea that the sex bias in natal dispersal distance in the Seychelles warbler is due to inbreeding avoidance and not resource competition or intrasexual competition for mates.     

Heritability of dispersal rate and other life history traits in the Glanville fritillary butterfly

Knowing the variances and heritabilities (h(2)) of life history traits in populations living under natural conditions is necessary for a mechanistic understanding of respective evolutionary processes. I estimated heritabilities of several life history traits, including dispersal rate, body mass, age at first reproduction, egg mass, clutch size and lifetime reproductive success, in the Glanville fritillary butterfly (Melitaea cinxia) using parent-offspring regression. Experiments were conducted under field conditions in a large population cage (32 x 26 m). Heritability estimates ranged from zero to almost one and several were significantly different from zero. Body size for both sexes, female age at first reproduction and egg weight were all moderately to highly heritable, whereas heritabilities were low or non-existent in clutch size and lifetime egg production. Heritability estimates for dispersal rate varied between the sexes, so that dispersal was heritable from mother to her female offspring only. This finding is consistent with previous results showing that the F1 female but not male offspring of females that naturally established new populations in the field are significantly more dispersive than butterflies in old populations.