Growth regulation in brook charr Salvelinus fontinalis.

Fish growth can be modulated through genetic selection. However, it is not known whether growth regulatory mechanisms modulated by genetic selection can provide information about phenotypic growth variations among families or populations. Following a five-generation breeding program that selected for the absence of early sexual maturity and increased growth in brook charr we aimed to understand how the genetic selection process modifies the growth regulatory pathway of brook charr at the molecular level. To achieve this, we studied the regulation of growth traits at three different levels: 1) between lines-one under selection, the other not, 2) among-families expressing differences in average growth phenotypes, which we termed family performance, and 3) among individuals within families that expressed extreme growth phenotypes, which we termed slow- and fast-growing. At age 1+, individuals from four of the highest performing and four of the lowest performing families in terms of growth were sampled in both the control and selected lines. The gene expression levels of three reference and ten target genes were analyzed by real-time PCR. Results showed that better growth performance (in terms of weight and length at age) in the selected line was associated with an upregulation in the expression of genes involved in the growth hormone (GH)/insulin growth factor-1 (IGF-1) axis, including the igf-1 receptor in pituitary; the gh-1 receptor and igf-1 in liver; and ghr and igf-1r in white muscle. When looking at gene expression within families, family performance and individual phenotypes were associated with upregulations of the leptin receptor and neuropeptid Y-genes related to appetite regulation-in the slower-growing phenotypes. However, other genes related to appetite (ghrelin, somatostatin) or involved in muscle growth (myosin heavy chain, myogenin) were not differentially expressed. This study highlights how transcriptomics may improve our understanding of the roles of different key endocrine steps that regulate physiological performance. Large variations in growth still exist in the selected line, indicating that the full genetic selection potential has not been reached.

Resource Availability, Sex, and Individual Differences in Exploration Drive Individual Diet Specialization.

AbstractIndividual diet specialization (IDS) is widespread and can affect the ecological and evolutionary dynamics of populations in significant ways. Extrinsic factors (e.g., food abundance) and individual variation in energetic needs, morphology, or physiology have been suggested as drivers of IDS. Behavioral traits like exploration and boldness can also impact foraging decisions, although their effects on IDS have not yet been investigated. Specifically, variation among individuals in exploratory behavior and their position along the exploration/exploitation trade-off may affect their foraging behavior, acquisition of food items, and home range size, which may in turn influence the diversity of their diet. Here, we analyzed stable carbon and nitrogen isotopes in hair of wild eastern chipmunks, Tamias striatus, to investigate the influence of individual differences in exploration on IDS. We found that exploration profile, sex, and yearly fluctuations in food availability explained differences in the degree of dietary specialization and in plasticity in stable carbon and stable nitrogen over time. Thus, consistent individual differences in exploration can be an important driver of within-population niche specialization and could therefore affect within-species competition. Our results highlight the need for a more thorough investigation of the mechanisms underlying the link between individual behavioral differences and diet specialization in wild animal populations.

The Feast and the Famine: Spring Body Mass Variations and Life History Traits in a Pulse Resource Ecosystem.

AbstractPhenotypic plasticity is the most immediate mechanism of adaptative response to environmental change. Studying plastic changes in response to fluctuating environments provides insights into how such adjustments may impact life history traits. Here, we used a 14-year data set of repeated body mass measurements in male eastern chipmunks (Tamias striatus) to assess the extent of plastic changes for this trait in a resource pulse ecosystem. We first determined the magnitude of variation in body mass at the population level in response to the drastic change in food resource availability from American beech tree seeds (Fagus grandifolia). Males that emerged in the spring from winter torpor following a nonmast year had a lower body mass than males emerging after a mast year, but they tended to recover this loss by mid-June. We found significant among-individual variation in spring body mass plasticity (i.e., individual-by-environment interaction). We then investigated the relationships between individual spring body mass plasticity, longevity, and lifetime reproductive success. Interestingly, heavier males lived longer than lighter males, but more plastic males had a lower longevity and lower lifetime reproductive success than less plastic males. The report of such plastic response in a stochastic resource system provides valuable insights into the interplay between the costs and benefits of phenotypic plasticity as an adaptation to environmental fluctuations.

Thermal regime during parental sexual maturation, but not during offspring rearing, modulates DNA methylation in brook charr (Salvelinus fontinalis).

Epigenetic inheritance can result in plastic responses to changing environments being faithfully transmitted to offspring. However, it remains unclear how epigenetic mechanisms such as DNA methylation can contribute to multigenerational acclimation and adaptation to environmental stressors. Brook charr (Salvelinus fontinalis), an economically important salmonid, is highly sensitive to thermal stress and is of conservation concern in the context of climate change. We studied the effects of temperature during parental sexual maturation and offspring rearing on whole-genome DNA methylation in brook charr juveniles (fry). Parents were split between warm and cold temperatures during sexual maturation, mated in controlled breeding designs, then offspring from each family were split between warm (8°C) and cold (5°C) rearing environments. Using whole-genome bisulfite sequencing, we found 188 differentially methylated regions (DMRs) due to parental maturation temperature after controlling for family structure. By contrast, offspring rearing temperature had a negligible effect on offspring methylation. Stable intergenerational inheritance of DNA methylation and minimal plasticity in progeny could result in the transmission of acclimatory epigenetic states to offspring, priming them for a warming environment. Our findings have implications pertaining to the role of intergenerational epigenetic inheritance in response to ongoing climate change.

Telomere length positively correlates with pace-of-life in a sex- and cohort-specific way and elongates with age in a wild mammal.

Understanding ageing and the diversity of life histories is a cornerstone in biology. Telomeres, the protecting caps of chromosomes, are thought to be involved in ageing, cancer risks and life-history strategies. They shorten with cell division and age in somatic tissues of most species, possibly limiting lifespan. The resource allocation trade-off hypothesis predicts that short telomeres have thus coevolved with early reproduction, proactive behaviour and reduced lifespan, that is, a fast pace-of-life syndrome (POLS). Conversely, since short telomeres may also reduce the risks of cancer, the anticancer hypothesis advances that they should be associated with slow POLS. Conclusion on which hypothesis best supports the role of telomeres as mediators of life-history strategies is hampered by a lack of study on wild short-lived vertebrates, apart from birds. Using seven years of data on wild Eastern chipmunks Tamias striatus, we highlighted that telomeres elongate with age (n = 204 and n = 20) and do not limit lifespan in this species (n = 51). Furthermore, short telomeres correlated with a slow POLS in a sex-specific way (n = 37). Females with short telomeres had a delayed age at first breeding and a lower fecundity rate than females with long telomeres, while we found no differences in males. Our findings support most predictions adapted from the anticancer hypothesis, but none of those from the resource allocation trade-off hypothesis. Results are in line with an increasing body of evidence suggesting that other evolutionary forces than resource allocation trade-offs shape the diversity of telomere length in adult somatic cells and the relationships between telomere length and life-histories.

Linking genetic, morphological, and behavioural divergence between inland island and mainland deer mice.

The island syndrome hypothesis (ISH) stipulates that, as a result of local selection pressures and restricted gene flow, individuals from island populations should differ from individuals within mainland populations. Specifically, island populations are predicted to contain individuals that are larger, less aggressive, more sociable, and that invest more in their offspring. To date, tests of the ISH have mainly compared oceanic islands to continental sites, and rarely smaller spatial scales such as inland watersheds. Here, using a novel set of genome-wide SNP markers in wild deer mice (Peromyscus maniculatus) we conducted a genomic assessment of predictions underlying the ISH in an inland riverine island system: analysing island-mainland population structure, and quantifying heritability of phenotypes thought to underlie the ISH. We found clear genomic differentiation between the island and mainland populations and moderate to high marker-based heritability estimates for overall variation in traits previously found to differ in line with the ISH between mainland and island locations. FST outlier analyses highlighted 12 loci associated with differentiation between mainland and island populations. Together these results suggest that the island populations examined are on independent evolutionary trajectories, the traits considered have a genetic basis (rather than phenotypic variation being solely due to phenotypic plasticity). Coupled with the previous results showing significant phenotypic differentiation between the island and mainland groups in this system, this study suggests that the ISH can hold even on a small spatial scale.

Interacting effects of cold snaps, rain, and agriculture on the fledging success of a declining aerial insectivore.

Climate change predicts the increased frequency, duration, and intensity of inclement weather periods such as unseasonably low temperatures (i.e., cold snaps) and prolonged precipitation. Many migratory species have advanced the phenology of important life history stages and, as a result, are likely to be exposed to these periods of inclement spring weather more often, therefore risking reduced fitness and population growth. For declining avian species, including aerial insectivores, anthropogenic landscape changes such as agricultural intensification are another driver of population declines. These landscape changes may affect the foraging ability of food provisioning parents and reduce the survival of nestlings exposed to inclement weather through, for example, pesticide exposure impairing thermoregulation and punctual anorexia. Breeding in agro-intensive landscapes may therefore exacerbate the negative effects of inclement weather under climate change. We observed that a significant reduction in the availability of insect prey occurred when daily maximum temperatures fell below 18.3°C, and thereby defined any day when the maximum temperature fell below this value as a day witnessing a cold snap. We then combined daily information on the occurrence of cold snaps and measures of precipitation to assess their impact on the fledging success of Tree Swallows (Tachycineta bicolor) occupying a nest box system placed across a gradient of agricultural intensification. Estimated fledging success of this declining aerial insectivore was 36.2% lower for broods experiencing 4 cold-snap days during the 12 days post-hatching period versus broods experiencing none, and this relationship was worsened when facing more precipitation. We further found that the overall negative effects of a brood experiencing periods of inclement weather was exacerbated in more agro-intensive landscapes. Our results indicate that two of the primary hypothesized drivers of many avian population declines may interact to further increase the rate of declines in certain landscape contexts.

Assessing pesticides exposure effects on the reproductive performance of a declining aerial insectivore.

In the context of increasing global environmental changes, it has become progressively important to understand the effects of human activity on wildlife populations. Declines in several avian populations have been observed since the 1970s, especially with respect to many farmland and grassland birds, which also include some aerial insectivores. Changes in farming practices referred to as agricultural intensification coincide with these major avian declines. Among those practices, increased pesticide use is hypothesized to be a key driver of avian population declines as it can lead to both toxicological and trophic effects. While numerous laboratory studies report that birds experience acute and chronic effects upon consuming pesticide treated food, little is known about the effects of the exposure to multiple pesticides on wildlife in natural settings. We monitored the breeding activities of Tree Swallows (Tachycineta bicolor) on 40 farms distributed over a gradient of agricultural intensification in southern Québec, Canada, to evaluate the presence of pesticides in their diet and quantify the exposure effects of those compounds on their reproductive performance between 2013 and 2018. We first assessed the presence of 54 active agents (or derivatives) found in pesticides in 2,081 food boluses (insects) delivered to nestlings by parents and documented their spatial distribution within our study area. Second, we assessed the effect of pesticide exposure through food (number of active agents detected and number of contaminated boluses on a given farm for a given year, while controlling for sampling effort) on clutch size as well as hatching and fledging successes and nestling's mass upon fledging. Pesticides were ubiquitous in our study system and nearly half (46%) of food boluses were contaminated by at least one active agent. Yet we found no relationship between our proxies of food contamination by pesticides and Tree Swallow reproductive performance. More studies are needed to better understand the putative role of pesticides in the decline of farmland birds and aerial insectivores as potential sublethal effects of pesticides can carry over to later life stages and impact fitness.

Consumption of red maple in anticipation of beech mast-seeding drives reproduction in eastern chipmunks.

Understanding the determinants of reproduction is a central question in evolutionary ecology. In pulsed-resource environments, the reproduction and population dynamics of seed consumers are driven by pulsed production of seeds by trees or mast-seeding. In Southern Québec, eastern chipmunks Tamias striatus exclusively reproduce during the summer before and the spring after a mast-seeding event of American beech. They thus seem to anticipate beech mast by reproducing during early summer, so that juveniles can emerge at the time of maximum beechnut abundance during late summer. However, the cues allowing chipmunks to anticipate beech mast remain unknown, and the existence of the anticipation process itself has been questioned. To tackle those issues, we investigated the links between the nutritional ecology and reproduction of adult chipmunks and compared their spring diet in mast- versus post-mast years. We monitored female reproductive status (N = 446), analysed cheek pouch contents at capture (n = 3,761 captures) and recorded seed production by deciduous trees on three different sites in Mont-Sutton from 2006 to 2018. Results revealed a systematic shift in chipmunk diet towards red maple seeds in springs preceding a beech mast, with red maple seeds composing more than 77% of chipmunk diet. However, red maple consumption was unrelated to red maple production, but was related to beech seed production in the upcoming fall. We also found that red maple consumption best predicted the proportion of females in summer oestrus. Our results confirm that chipmunks anticipate beech mast-seeding and highlight a key role of red maple consumption in that anticipation. Results also suggest that red maple seeds may contain nutrients or secondary plant components essential to sustain or trigger the summer reproduction in chipmunks, which allow them to remain synchronized with pulsed productions of both red maple and beech and improve their fitness.

Development and characterization of 14 microsatellites for the eastern chipmunk, Tamias striatus.

The number of microsatellite markers currently available for the eastern chipmunk Tamias striatus provides limited capacity to achieve sufficient pedigree building for the study of their genetic structure and relatedness patterns. We developed microsatellite loci for the eastern chipmunk, a small rodent commonly found in eastern North America. More specifically, we report data for 14 loci and 50 individuals genotyped from a population in southern Québec, Canada. We found a number of alleles ranging from 5 to 21 and there was no linkage disequilibrium among locus. One locus deviated from Hardy-Weinberg equilibrium and had a high proportion of null alleles. Those loci will be used in addition to previously developed loci to improve the precision of parentage assignment and population genetics studies on this species.

Individual and environmental determinants of Cuterebra bot fly parasitism in the eastern chipmunk (Tamias striatus).

Understanding the interactions between parasites, hosts, and their shared environment is central to ecology. Variation in infestation prevalence may be the result of varying environmental and population characteristics; however, variations in parasitism may also depend on individual characteristics that influence both the exposure and susceptibility to parasites. Using 12 years of data from a population of wild eastern chipmunks relying on pulsed food resources, we investigated the determinants of bot fly parasitism at both the population and individual level. We assessed the relationship between infestation prevalence and weather conditions, population size and food abundance. Then, we assessed the relationship between infestation intensity and chipmunk behavior, sex, age, body mass and food abundance. Precipitation, temperature and population size were positively related to infestation prevalence, while beech masts were negatively related to infestation prevalence, highlighting the importance of local environmental conditions on hosts and parasites. We also found that the influence of activity and exploration on infestation intensity varied according to sex in adults. More active and faster exploring males had more parasites compared to females, suggesting that reproductive behaviors may influence parasite exposure. For juveniles, infestation intensity was greater when juveniles emerged in the spring as opposed to fall, possibly because spring emergence is synchronized with the peak of bot fly eggs in the environment, low food availability and longer activity period. Our results suggest that the environmental, population and host characteristics that are advantageous for reproduction and resource acquisition may come at the cost of increasing parasitism.

A range-wide domino effect and resetting of the annual cycle in a migratory songbird.

Latitudinal differences in timing of breeding are well documented but how such differences carry over to influence timing of events in the annual cycle of migratory birds is not well understood. We examined geographical variation in timing of events throughout the year using light-level geolocator tracking data from 133 migratory tree swallows ( Tachycineta bicolor) originating from 12 North American breeding populations. A swallow's breeding latitude influenced timing of breeding, which then carried over to affect breeding ground departure. This resulted in subsequent effects on the arrival and departure schedules at autumn stopover locations and timing of arrival at non-breeding locations. This 'domino effect' between timing events was no longer apparent by the time individuals departed for spring migration. Our range-wide analysis demonstrates the lasting impact breeding latitude can have on migration schedules but also highlights how such timing relationships can reset when individuals reside at non-breeding sites for extended periods of time.

Current spring warming as a driver of selection on reproductive timing in a wild passerine.

Evolutionary adaptation as a response to climate change is expected for fitness-related traits affected by climate and exhibiting genetic variance. Although the relationship between warmer spring temperature and earlier timing of reproduction is well documented, quantifications and predictions of the impact of global warming on natural selection acting on phenology in wild populations remain rare. If global warming affects fitness in a similar way across individuals within a population, or if fitness consequences are independent of phenotypic variation in key-adaptive traits, then no evolutionary response is expected for these traits. Here, we quantified the selection pressures acting on laying date during a 24-year monitoring of blue tits in southern Mediterranean France, a hot spot of climate warming. We explored the temporal fluctuation in annual selection gradients and we determined its temperature-related drivers. We first investigated the month-specific warming since 1970 in our study site and tested its influence on selection pressures, using a model averaging approach. Then, we quantified the selection strength associated with temperature anomalies experienced by the blue tit population. We found that natural selection acting on laying date significantly fluctuated both in magnitude and in sign across years. After identifying a significant warming in spring and summer, we showed that warmer daily maximum temperatures in April were significantly associated with stronger selection pressures for reproductive timing. Our results indicated an increase in the strength of selection by 46% for every +1°C anomaly. Our results confirm the general assumption that recent climate change translates into strong selection favouring earlier breeders in passerine birds. Our findings also suggest that differences in fitness among individuals varying in their breeding phenology increase with climate warming. Such climate-driven influence on the strength of directional selection acting on laying date could favour an adaptive response in this trait, since it is heritable.

Environmental determinants of haemosporidian parasite prevalence in a declining population of Tree swallows.

The prevalence of vector-borne parasites such as haemosporidian species is influenced by several environmental factors. While the negative effects of parasitism on hosts are well documented, these can also be amplified by interactions with environmental stressors, many of which are anthropogenic. Yet, we know little about the possible effects of anthropogenic perturbations on parasite prevalence. The goals of this study were to assess the prevalence and environmental determinants of haemosporidian parasites in a declining population of Tree swallows (Tachycineta bicolor) living in an agricultural landscape in southern Québec, Canada. Overall, a low prevalence and a moderate lineage diversity were identified in both adults and nestlings, confirming that transmission can occur during the breeding period. Anthropic areas, extensive cultures (hayfields and pastures) and forest cover within 500 km of nest boxes, as well as daily temperature fluctuations, were all related to infection by haemosporidian parasites. These findings suggest that anthropogenic alterations of landscape composition can modulate the prevalence of haemosporidian parasites in Tree swallows. Our results represent a baseline for future comparative studies assessing haemosporidian parasite prevalence in human-modified landscapes.

Introgressive hybridization between wild and domestic individuals and its relationship with parasitism in brook charr Salvelinus fontinalis.

The effects of introgression on parasitism in brook charr Salvelinus fontinalis were investigated in 28 lakes with various levels of stocking in Québec, Canada. No effect of genetic background on parasitism was found at the individual level. Body length seemed to explain most of the variation observed at this level, with largest fish being more infected. However, lakes with the greater average domestic genetic background were found to display significantly lower parasite prevalence and diversity. Since our results indicate no effect of domestic genes at the individual level, the negative association with introgression found at the population level may be mainly attributed to differences in intrinsic environmental quality of lakes (e.g. fishing pressure, availability of food resources, abiotic characteristics).

Patterns of Diversity and Spatial Variability of ß-Defensin Innate Immune Genes in a Declining Wild Population of Tree Swallows.

Assessing the genetic variation and distribution of immune genes across heterogeneous environmental conditions in wild species is essential to further our understanding of the role of pathogen pressure and potential resistance or prevalence in hosts. Researchers have recently investigated ß-defensin genes in the wild, because their variability suggests that they may play an important role in innate host defense. This study investigated the variation occurring at 6 innate immune genes of the ß-defensin family in a declining population of tree swallows (Tachycineta bicolor) in southern Québec, Canada (N = 160). We found that all 6 genes showed synonymous and nonsynonymous single nucleotide polymorphisms (SNPs) within the exon coding for the mature peptide. These results indicated that this group of genes was diverse in tree swallows. Our results suggested a potential interaction of this group of genes with fluctuating pathogen diversity, however, we found no sign of positive or negative selection. We assessed whether or not the distribution of genetic diversity of ß-defensin genes in our study population differed between 2 regions that strongly differ in their level of agricultural intensification. Adults are highly philopatric to their breeding sites and their immunological responses differ between these 2 regions. However, we found little evidence that the level and distribution of genetic variability differed between these heterogeneous environmental conditions. Further studies should aim to assess the link between genetic diversity of ß-defensin genes and fitness-related traits in wild populations.

Severe recent decrease of adult body mass in a declining insectivorous bird population.

Migratory bird species that feed on air-borne insects are experiencing widespread regional declines, but these remain poorly understood. Agricultural intensification in the breeding range is often regarded as one of the main drivers of these declines. Here, we tested the hypothesis that body mass in breeding individuals should reflect habitat quality in an aerial insectivore, the tree swallow (Tachycineta bicolor), along a gradient of agricultural intensity. Our dataset was collected over 7 years (2005-2011) and included 2918 swallow captures and 1483 broods. Analyses revealed a substantial decline of the population over the course of the study (-19% occupancy rate), mirrored by decreasing body mass. This trend was especially severe in females, representing a total loss of 8% of their mass. Reproductive success was negatively influenced by intensive agriculture, but did not decrease over time. Interestingly, variation in body mass was independent of breeding habitat quality, leading us to suggest that this decline in body mass may result from carry-over effects from non-breeding areas and affect population dynamics through reduced survival. This work contributes to the growing body of evidence suggesting that declines in migratory aerial insectivores are driven by multiple, complex factors requiring better knowledge of year-round habitat use.

Drawing ecological inferences from coincident patterns of population- and community-level biodiversity.

Biodiversity is comprised of genetic and phenotypic variation among individual organisms, which might belong to the same species or to different species. Spatial patterns of biodiversity are of central interest in ecology and evolution for several reasons: to identify general patterns in nature (e.g. species-area relationships, latitudinal gradients), to inform conservation priorities (e.g. identifying hotspots, prioritizing management efforts) and to draw inferences about processes, historical or otherwise (e.g. adaptation, the centre of origin of particular clades). There are long traditions in ecology and evolutionary biology of examining spatial patterns of biodiversity among species (i.e. in multispecies communities) and within species, respectively, and there has been a recent surge of interest in studying these two types of pattern simultaneously. The idea is that examining both levels of diversity can materially advance the above-stated goals and perhaps lead to entirely novel lines of inquiry. Here, we review two broad categories of approach to merging studies of inter- and intraspecific variation: (i) the study of phenotypic trait variation along environmental gradients and (ii) the study of relationships between patterns of molecular genetic variation within species and patterns of distribution and diversity across species. For the latter, we report a new meta-analysis in which we find that correlations between species diversity and genetic diversity are generally positive and significantly stronger in studies with discrete sampling units (e.g. islands, lakes, forest fragments) than in studies with nondiscrete sampling units (e.g. equal-area study plots). For each topic, we summarize the current state of knowledge and key future directions.

Neutral and selective processes shape MHC gene diversity and expression in stocked brook charr populations (Salvelinus fontinalis).

The capacity of an individual to battle infection is an important fitness determinant in wild vertebrate populations. The major histocompatibility complex (MHC) genes are crucial for a host's adaptive immune system to detect pathogens. However, anthropogenic activities may disrupt natural cycles of co-evolution between hosts and pathogens. In this study, we investigated the dynamic sequence and expression variation of host parasite interactions in brook charr (Salvelinus fontinalis) in a context of past human disturbance via population supplementation from domestic individuals. To do so, we developed a new method to examine selection shaping MHC diversity within and between populations and found a complex interplay between neutral and selective processes that varied between lakes that were investigated. We provided evidence for a lower introgression rate of domestic alleles and found that parasite infection increased with domestic genomic background of individuals. We also documented an association between individual MHC alleles and parasite taxa. Finally, longer cis-regulatory minisatellites were positively correlated with MHC II down-regulation and domestic admixture, suggesting that inadvertent selection during domestication resulted in a lower immune response capacity, through a trade-off between growth and immunity, which explained the negative selection of domestic alleles at least under certain circumstances.

Pulsed resources and the coupling between life-history strategies and exploration patterns in eastern chipmunks (Tamias striatus).

Understanding the causes of animal personality (i.e. consistent behavioural differences) is a major aim of evolutionary studies. Recent theoretical work suggests that major personality traits may contribute to evolutionary trade-offs. However, such associations have only been investigated in a few study systems, and even less so in free ranging animal populations. Eastern chipmunks exhibit consistent individual differences in exploration, ranging from slow to fast. Birth cohorts also experience dramatic differences in age at first breeding opportunity due to annual differences in beech mast. Individuals may breed for the first time at 24, 33 or 50% of their average life span, depending on year of birth. Here, we used data from a long-term survey on a wild population to investigate the relationship between reproductive life history and consistent individual differences in exploration. We determined whether predictable differences in age at first breeding opportunity among birth cohorts were associated with exploration differences and favoured individuals with different exploration. Birth cohorts with a predictably earlier age at first breeding opportunity were faster explorers on average. Slower explorers displayed their highest fecundity (females) or highest fertilization success (males) later in their life compared with faster explorers. Overall, slow explorers attained a higher lifetime reproductive success than fast explorers when given an opportunity to reproduce later in their life. Our results suggest that the timing of mating seasons, associated with fluctuating food abundance, may favour individual variation in exploration and maintain population variation through its effects on reproductive life history. Together, our result shed light on how fluctuation in ecological conditions may maintain personality differences and on the nature of the relationships between animal personality and life history.