Extended incubation recesses in sanderlings are impacted by temperature and body condition.

Complex incubation strategies have evolved to solve the trade-off between parent survival and care for their eggs with often brief departures (recesses) that maximize egg survival, and infrequent extended recesses maximizing adult condition. Here we examined incubation behaviour of sanderlings (Calidris alba), a species that exhibits both biparental and uniparental incubation behaviour. During 11 breeding seasons in Greenland, we have quantified incubation variability with thermologgers placed in nests. We estimated the impact of environmental conditions and individual characteristics on the occurrence and the duration of recesses. We found that extended recesses are a unique feature of uniparentals, and their frequency and duration increased in colder temperatures. The relationship was mediated by body condition, with individuals in poor condition performing longer extended recesses in colder temperatures. This suggests that extended recesses may represent a shift towards self-maintenance at the expense of the egg care, allowing birds to continue incubating under unfavourable conditions. Our study illustrates how extended recesses may be a key breeding strategy to overcome high energetic costs associated with incubation. Quantifying such behavioural flexibility paves the way for tracking future behavioural responses of individuals in the face of changing environments.

Coccidial oocyst release: once a day or all day long? Tropical bird hosts shed new light on the adaptive significance of diurnal periodicity in parasite output.

Many parasites spend part of their life cycle as infectious forms released from an infected host in the external environment, where they may encounter and infect new hosts. The emergence of infectious life stages often occurs once a day to minimize mortality in adverse environments. In bird hosts, intestinal parasites such as coccidia are generally released with feces in the late afternoon. This dynamic is adaptive since it allows avoiding desiccation and ultraviolet (UV) radiation, thus reducing mortality of oocysts in the environment until transmission to the next host. If this circadian rhythm is the result of natural selection to increase oocyst survival, we may hypothesize that oocysts will appear in feces at different times depending on the environment where hosts live. Particularly, in an environment where UV radiation and desiccation are very low, we may expect oocyst circadian release to disappear since the main selective pressure would be relaxed. We sampled different species of birds in tropical and temperate forests in spring and investigated coccidian oocyst output. A strong circadian variation in the prevalence of hosts shedding coccidian oocyst was detected for species caught in the temperate forest with an increase in prevalence in the late afternoon, whereas prevalence of birds shedding oocysts was constant over the course of the day for most species sampled in the tropical rain forest. These results are consistent with the hypothesis that oocysts' circadian output is maintained by natural selection to increase oocyst survival. We discuss the adaptive significance of diurnal periodicity in parasite output.

Pesticide impacts on avian species with special reference to farmland birds: a review.

For decades, we have observed a major biodiversity crisis impacting all taxa. Avian species have been particularly well monitored over the long term, documenting their declines. In particular, farmland birds are decreasing worldwide, but the contribution of pesticides to their decline remains controversial. Most studies addressing the effects of agrochemicals are limited to their assessment under controlled laboratory conditions, the determination of lethal dose 50 (LD50) values and testing in a few species, most belonging to Galliformes. They often ignore the high interspecies variability in sensitivity, delayed sublethal effects on the physiology, behaviour and life-history traits of individuals and their consequences at the population and community levels. Most importantly, they have entirely neglected to test for the multiple exposure pathways to which individuals are subjected in the field (cocktail effects). The present review aims to provide a comprehensive overview for ecologists, evolutionary ecologists and conservationists. We aimed to compile the literature on the effects of pesticides on bird physiology, behaviour and life-history traits, collecting evidence from model and wild species and from field and lab experiments to highlight the gaps that remain to be filled. We show how subtle nonlethal exposure might be pernicious, with major consequences for bird populations and communities. We finally propose several prospective guidelines for future studies that may be considered to meet urgent needs.

Storage of Carotenoids in Crustaceans as an Adaptation to Modulate Immunopathology and Optimize Immunological and Life-History Strategies.

Why do some invertebrates store so much carotenoids in their tissues? Storage of carotenoids may not simply be passive and dependent on their environmental availability, as storage variation exists at various taxonomic scales, including among individuals within species. While the strong antioxidant and sometimes immune-stimulating properties of carotenoids may be beneficial enough to cause the evolution of features improving their assimilation and storage, they may also have fitness downsides explaining why massive carotenoid storage is not universal. Here, the functional and ecological implications of carotenoid storage for the evolution of invertebrate innate immune defenses are examined, especially in crustaceans, which massively store carotenoids for unclear reasons. Three testable hypotheses about the role of carotenoid storage in immunological (resistance and tolerance) and life-history strategies (with a focus on aging) are proposed, which may ultimately explain the storage of large amounts of these pigments in a context of host-pathogen interactions.

Reproductive senescence and parental effects in an indeterminate grower.

Reproductive senescence is the decrease of reproductive performance with increasing age and can potentially include trans-generational effects as the offspring produced by old parents might have a lower fitness than those produced by young parents. This negative effect may be caused either by the age of the father, mother or the interaction between the ages of both parents. Using the common woodlouse Armadillidium vulgare, an indeterminate grower, as a biological model, we tested for the existence of a deleterious effect of parental age on fitness components. Contrary to previous findings reported from vertebrate studies, old parents produced both a higher number and larger offspring than young parents. However, their offspring had lower fitness components (by surviving less, producing a smaller number of clutches or not reproducing at all) than offspring born to young parents. Our findings strongly support the existence of trans-generational senescence in woodlice and contradict the belief that old individuals in indeterminate growers contribute the most to recruitment and correspond thereby to the key life stage for population dynamics. Our work also provides rare evidence that the trans-generational effect of senescence can be stronger than direct reproductive senescence in indeterminate growers.

Where you come from matters: temperature influences host-parasitoid interaction through parental effects.

Temperature alters host suitability for parasitoid development through direct and indirect pathways. Direct effects depend on ambient temperatures experienced by a single host individual during its lifetime. Indirect effects (or parental effects) occur when thermal conditions met by a host parental generation affect the way its offspring will interact with parasitoids. Using the complex involving eggs of the moth Lobesia botrana as hosts for the parasitoid Trichogramma cacoeciae, we developed an experimental design to disentangle the effects of (1) host parental temperature (temperature at which the host parental generation developed and laid host eggs) and (2) host offspring temperature (temperature at which host eggs were incubated following parasitism, i.e. direct thermal effects) on this interaction. The host parental generation was impacted by temperature experienced during its development: L. botrana females exposed to warmer conditions displayed a lower pupal mass but laid more host eggs over a 12-h period. Host parental temperature also affected the outcomes of the interaction. Trichogramma cacoeciae exhibited lower emergence rates but higher hind tibia length on emergence from eggs laid under warm conditions, even if they were themselves exposed to cooler temperatures. Such indirect thermal effects might arise from a low nutritional quality and/or a high immunity of host eggs laid in warm conditions. By contrast with host parental temperature, offspring temperature (direct thermal effects) did not significantly affect the outcomes of the interaction. This work emphasises the importance of accounting for parental thermal effects to predict the future of trophic dynamics under global warming scenarios.

Trans-generational Immune Priming Protects the Eggs Only against Gram-Positive Bacteria in the Mealworm Beetle.

In many vertebrates and invertebrates, offspring whose mothers have been exposed to pathogens can exhibit increased levels of immune activity and/or increased survival to infection. Such phenomena, called "Trans-generational immune priming" (TGIP) are expected to provide immune protection to the offspring. As the offspring and their mother may share the same environment, and consequently similar microbial threats, we expect the immune molecules present in the progeny to be specific to the microbes that immune challenged the mother. We provide evidence in the mealworm beetle Tenebrio molitor that the antimicrobial activity found in the eggs is only active against Gram-positive bacteria, even when females were exposed to Gram-negative bacteria or fungi. Fungi were weak inducers of TGIP while we obtained similar levels of anti-Gram-positive activity using different bacteria for the maternal challenge. Furthermore, we have identified an antibacterial peptide from the defensin family, the tenecin 1, which spectrum of activity is exclusively directed toward Gram-positive bacteria as potential contributor to this antimicrobial activity. We conclude that maternal transfer of antimicrobial activity in the eggs of T. molitor might have evolved from persistent Gram-positive bacterial pathogens between insect generations.

Personality, immune response and reproductive success: an appraisal of the pace-of-life syndrome hypothesis.

The pace-of-life syndrome (POLS) hypothesis is an extended concept of the life-history theory that includes behavioural traits. The studies challenging the POLS hypothesis often focus on the relationships between a single personality trait and a physiological and/or life-history trait. While pathogens represent a major selective pressure, few studies have been interested in testing relationships between behavioural syndrome, and several fitness components including immunity. The aim of this study was to address this question in the mealworm beetle, Tenebrio molitor, a model species in immunity studies. The personality score was estimated from a multidimensional syndrome based of four repeatable behavioural traits. In a first experiment, we investigated its relationship with two measures of fitness (reproduction and survival) and three components of the innate immunity (haemocyte concentration, and levels of activity of the phenoloxidase including the total proenzyme and the naturally activated one) to challenge the POLS hypothesis in T. molitor. Overall, we found a relationship between behavioural syndrome and reproductive success in this species, thus supporting the POLS hypothesis. We also showed a sex-specific relationship between behavioural syndrome and basal immune parameters. In a second experiment, we tested whether this observed relationship with innate immunity could be confirmed in term of differential survival after challenging by entomopathogenic bacteria, Bacillus thuringiensis. In this case, no significant relationship was evidenced. We recommend that future researchers on the POLS should control for differences in evolutionary trajectory between sexes and to pay attention to the choice of the proxy used, especially when looking at immune traits.

Immune benefits from alternative host plants could maintain polyphagy in a phytophagous insect.

The tritrophic interactions hypothesis, integrating bottom-up (plant-herbivore) and top-down (herbivore-natural enemies) effects, predicts that specialist herbivores should outcompete generalists. However, some phytophagous insects have generalist diets, suggesting that maintenance of a diverse diet may confer certain fitness advantages that outweigh diet specialization. In field conditions, the European grapevine moth, Lobesia botrana, feeds on diverse locally rare alternative host plants (AHPs) although grapevines are a highly abundant and predictable food source. The laboratory studies presented here show that survival, growth, and constitutive levels of immune defences (concentration of haemocytes and phenoloxidase activity) of L. botrana larvae were significantly enhanced when they were fed AHPs rather than grape. These results indicated a strong positive effect of AHPs on life history traits and immune defences of L. botrana. Such positive effects of AHPs should be advantageous to the moth under heavy selective pressure by natural enemies and, as a consequence, favour the maintenance of a broad diet preference in this species. We therefore believe that our results account for the role of immunity in the maintenance of polyphagy in phytophagous insects.

Pesticide contamination patterns in Montagu's harrier (Circus pygargus) chicks.

Biomonitoring of persistent pesticides in birds of prey has been carried out for decades, but few studies have investigated their relevance for the monitoring of non-persistent pesticides. Herein, we determined the contamination patterns of multiple pesticides in Montagu's harrier (Circus pygargus) chicks in an intensive farming area of southwestern France. Blood samples from 55 chicks belonging to 22 nests in 2021 were assessed for 104 compounds (herbicides, fungicides, insecticides, safeners and synergists). All chicks had at least one herbicide in their blood, and half had at least two compounds. The 28 compounds detected comprised 10 herbicides, 12 fungicides, 5 insecticides and 1 synergist. Mixtures in blood were predominantly composed of herbicides, and six chicks presented a mixture of the three pesticide classes. The most prevalent compounds were sulcotrione (96% of chicks), tebutam (44%) and chloridazon (31%), of which the latter two had been banned in France for 19 and 3 years, respectively, at the time of sampling. Most compounds are considered non-acutely toxic, but sulcotrione is potentially carcinogenic, mutagenic and reprotoxic, raising questions about the effects on the health of nestlings. Biomonitoring of multiple pesticides through Montagu's harrier chicks in agroecosystems is clearly relevant because it reflects the general pattern of agricultural pesticide use in the study area. It also raises questions about exposure pathways in chicks, and further investigations are needed to disentangle the roles of dietary routes and maternal transfer for the established pesticide contamination patterns.

Interaction between females and males grapevine moth Lobesia botrana modifies further mating preference.

During reproduction, females may boost their fitness by being selective based on direct material benefits provided by the males, such as nuptial gifts. In Lepidoptera, male provides a spermatophore containing nutrients. However, virgin males produce a bigger spermatophore, containing spermatozoa and nutrients, allowing higher female fertility. Lepidoptera females that could detect the sexual status of males may thus prefer a male without previous mating experience (i.e. a virgin male). This mate selection could be achieved by the use of chemical indices, such as sexual pheromones and cuticular compounds, known to be possibly exchanged during reproduction, and which can be indicators of a previous mating experience and known to be possibly sources of information exchanged. In this study, we experimentally presented Lobesia botrana virgin males with females in order for them to be exposed to females' natural sexual pheromones or cuticular compounds. 12 or 48 h after the exposure of males to either females' sexual pheromones or cuticular compounds, these males were confronted to naïve females, which have a choice between them or a virgin non-exposed males. We highlighted that, despite producing a spermatophore of similar volume, all exposed virgin males were less likely to mate with females 12 h after exposure, while after 48 h of exposure this is only the case for virgin males exposed to sexual pheromones. L. botrana females may thus discriminate male sexual experience based on chemical cues (either from cues transferred directly from females to males, or from changes in the cuticular or pheromone males' profile) indicating past mating experiences. Mating duration was longer for males exposed to sexual pheromones after 12 h only, and for males exposed to cuticular compounds after 48 h only. Pheromones signal might be more persistent over time and seems to more easily gather information for males. The physiological reasoning behind this result still needs to be investigated.

Organic farming reduces pesticide load in a bird of prey.

Human activities have led to the contamination of all environmental compartments worldwide, including bird species. In birds, both the environment and maternal transfer lead to high inter-brood variability in contamination levels of pollutants, whereas intra-brood variability is generally low. However, most existing studies focused on heavy metals or persistent compounds and none, to our knowledge, addressed the variability in contamination levels of multiple pesticides and the factors influencing it. In this study, the number of pesticides detected (of 104 compounds searched) and the sum of their concentrations in the blood of 55 Montagu's harrier (Circus pygargus) nestlings from 22 nests sampled in 2021 were used as metrics of contamination levels. We investigated the effect of organic farming at the size of male's home range (i.e., 14 km2) and chicks' sex and hatching order on contamination levels. We did not find a difference between inter-brood and intra-brood variability in pesticide contamination levels, suggesting a different exposure of siblings through food items. While chicks' sex or rank did not affect their contamination level, we found that the percentage of organic farming around the nests significantly decreased the number of pesticides detected, although it did not decrease the total concentrations. This finding highlights the potential role of organic farming in reducing the exposure of birds to a pesticide cocktail.

Comparison of pesticide contamination between captive-reared and wild grey partridges: insights into environmental exposure disparities.

Pesticide contamination is often cited as a key factor in the global decline of farmland birds. However, the majority of studies on pesticide exposure in non-target fauna are not representative of what happens in nature because they are limited to artificial conditions. The aim of this study was to define and compare, for the first time, pesticide contamination in grey partridges (Perdix perdix) from two different contexts, i.e., captivity vs. the wild. Blood samples taken from 35 captive and 54 wild partridges in 2021-2022 were analysed for 94 pesticides most commonly used in French agriculture. Captive partridges had 29 molecules detected in their blood (12 herbicides, 14 fungicides, and three insecticides) compared to wild partridges, which had 50 molecules (13 herbicides, 23 fungicides, and 14 insecticides). Of these pesticide compounds found in individuals, 26 were banned. Captive partridges had significantly fewer pesticide molecules than wild partridges, with one to 14 pesticides per captive individual and 8 to 20 pesticides per wild individual. Nineteen molecules were common to both groups, with concentrations up to three times higher in wild partridges than in captive partridges. Our results thus show multiple exposures for most of our individuals, especially in wild partridges, which can lead to cocktail effects, which are never considered. Furthermore, the difference in contamination between the wild and captive partridges reflects the multiple routes of contamination in nature, in particular, due to the use of a wide range of habitats by wild partridges.

Pest management facing warming and chemical stresses: Multi-stress effects on the biological agent Trichogramma oleae.

Global change is affecting plant-insect interactions in agroecosystems and can have dramatic consequences on yields when causing non-targeted pest outbreaks and threatening the use of pest natural enemies for biocontrol. The vineyard agroecosystem is an interesting system to study multi-stress conditions: on the one hand, agricultural intensification comes with high inputs of copper-based fungicides and, on the other hand, temperatures are rising due to climate change. We investigated interactive and bottom-up effects of both temperature increase and copper-based fungicides exposure on the important Lepidopteran vineyard pest Lobesia botrana and its natural enemy, the oophagous parasitoid Trichogramma oleae. We exposed L. botrana larvae to three increasing copper sulfate concentrations under two fluctuating thermal regimes, one current and one future. Eggs produced by L. botrana were then exposed to T. oleae. Our results showed that the survival of L. botrana, was only reduced by the highest copper sulfate concentration and improved under the warmer regime. The development time of L. botrana was strongly reduced by the warmer regime but increased with increasing copper sulfate concentrations, whereas pupal mass was reduced by both thermal regime and copper sulfate. T. oleae F1 emergence rate was reduced and their development time increased by combined effects of the warmer regime and increasing copper sulfate concentrations. Size, longevity and fecundity of T. oleae F1 decreased with high copper sulfate concentrations. These effects on the moth pest and its natural enemy are probably the result of trade-offs between the survival and the development of L. botrana facing multi-stress conditions and implicate potential consequences for future biological pest control. Our study supplies valuable data on how the interaction between pests and biological control agents is affected by multi-stress conditions.

Adverse effects of the Bordeaux mixture copper-based fungicide on the non-target vineyard pest Lobesia botrana.

BACKGROUND: Bordeaux mixture is a copper-based fungicide commonly used in vineyards to prevent fungal and bacterial infections in grapevines. However, this fungicide may adversely affect the entomological component, including insect pests. Understanding the impacts of Bordeaux mixture on the vineyard pest Lobesia botrana is an increasing concern in the viticultural production. RESULTS: Bordeaux mixture had detrimental effects on the development and reproductive performance of L. botrana. Several physiological traits were adversely affected by copper-based fungicide exposure, including a decrease in larval survival and a delayed larval development to moth emergence, as well as a reduced reproductive performance through a decrease in female fecundity and fertility and male sperm quality. However, we did not detect any effect of Bordeaux mixture on the measured reproductive behaviors (mating success, pre-mating latency and mating duration). CONCLUSION: Ingestion by larvae of food contaminated with Bordeaux mixture had a negative effect on the reproductive performance of the pest L. botrana, which could affect its population dynamics in vineyards. Although this study highlighted collateral damage of Bordeaux mixture on L. botrana, the potential impact of copper-based fungicides on vineyard diversity, including natural predators is discussed and needs to be taken in consideration in integrated pest management. © 2024 The Author(s). Pest Management Science published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.

Geographical variation in parasitism shapes larval immune function in a phytophagous insect.

Two of the central goals of immunoecology are to understand natural variation in the immune system among populations and to identify those selection pressures that shape immune traits. Maintenance of the immune system can be costly, and both food quality and parasitism selection pressure are factors potentially driving immunocompetence. In tritrophic interactions involving phytophagous insects, host plants, and natural enemies, the immunocompetence of phytophagous insects is constrained by selective forces from both the host plants and the natural enemies. Here, we assessed the roles of host plants and natural enemies as selective pressures on immune variation among natural populations of Lobesia botrana. Our results showed marked geographical variation in immune defenses and parasitism among different natural populations. Larval immune functions were dependent of the host plant quality and were positively correlated to parasitism, suggesting that parasitoids select for greater investment into immunity in moth. Furthermore, investment in immune defense was negatively correlated with body size, suggesting that it is metabolically expensive. The findings emphasize the roles of host plants and parasitoids as selective forces shaping host immune functions in natural conditions. We argue that kinds of study are central to understanding natural variations in immune functions, and the selective forces beyond.

Neonicotinoids: Still present in farmland birds despite their ban.

Neonicotinoids (neonics) are the most widely used insecticides worldwide and are considered to be of low risk to non-target organisms such as vertebrates. Further, they are reported to be rapidly excreted and metabolized, reducing their potential toxicity. Nevertheless, growing evidence of adverse effects of neonics on farmland bird species raise questions about the purported harmless nature of these pesticides. We attempted to search for pesticide residues in species of different trophic levels and at different life stages, by using multiple bird monitoring programs on a Long-Term Socio-Ecological Research (LTSER) platform. Three passerine birds-the blackbird (Turdus merula), cirl bunting (Emberiza cirlus), and common nightingale (Luscinia megarhynchos)-that feed on seeds and invertebrates were monitored during their reproductive period, and the grey partridge (Perdix perdix) that feeds on seeds was monitored during its wintering period. We also monitored chicks of an apex predator-the Montagu's harrier (Circus pygargus)-that preys mostly upon common voles but also upon insects. We found that the birds' blood samples showed presence of residues of five neonics: three banned since 2018 in France-clothianidin, thiacloprid, and thiamethoxam-and two-dinotefuran and nitenpyram-used for veterinary purposes only. While none of these neonics was detected in blackbirds, all were present in grey partridges. Clothianidin was detected in all species, except blackbirds. Concentrations of the three banned neonics were similar or higher than concentrations found in birds monitored elsewhere before the ban. These findings raise questions about the persistence of neonics within the environment and the mode of exposure to wild fauna. Future investigations on the sublethal effects of these neonics on life-history traits of these farmland birds may help in providing a better understanding of the effects of exposure of bird populations to these insecticides, and also to the consequent effect on human health.

Sexual audience affects male's reproduction investment without consequences on reproductive outputs.

Males evolved plastic strategies to respond to male-male competition and exhibit adaptive traits and behaviors maximizing their access to the females and limiting sperm competition. Mating behaviors allow males to express quick responses to current sexual audience, that is, the number of nearby conspecifics prone to mate. In contrast, physiological responses are frequently delayed because they are constrained by the time and resources having to be mobilized to produce and export sperm and associated products. This is especially critical in species for which males produce spermatophores. Here we investigated in what extend moth males (the tortricid moth Lobesia botrana) producing spermatophores exhibit plastic behavioral and physiological responses to different sexual audiences before and during mating and the consequences for their reproductive output. We found that males adjusted their mating behaviors and spermatophore size to a potentially elevated risk of sperm competition perceived before mating. In addition, males responded to the closed presence of females during mating by reducing their mating duration. Surprisingly, the various behavioral and physiological responses we highlighted here were not fully reflected in their reproductive performance as we did not reveal any effect on fecundity and fertility of their mate. The selective pressure exerted on males experiencing male-male competition could thus be sufficient to trigger adjustment in male mating behaviors but constrains physiological responses according to the perception of competition.

Feeding on grains containing pesticide residues is detrimental to offspring development through parental effects in grey partridge.

Numerous toxicological studies have shown that ingestion of pesticides can induce physiological stress in breeding birds, with adverse consequences on egg laying parameters and offspring quality through parental effects. However, previous studies do not mimic current levels of pesticide residues in typical landscapes, and they do not consider potential cocktail effects of pesticides as they occur in the wild. Herein, we explored whether realistic pesticide exposure affected reproduction parameters and offspring condition through parental effects in Grey partridge. We fed 24 breeding pairs with either seeds from conventional agriculture crops treated with various pesticides during cropping, or organic grains without pesticide residues as controls. The conventional and organic grain diets mimicked food options potentially encountered by wild birds in the field. The results showed that ingesting low pesticide doses over a long period had consequences on reproduction and offspring quality without altering mortality in parents or chicks. Compared with organic pairs, conventional pairs yielded smaller chicks at hatching that had a lower body mass index at 24 days old. Additionally, these chicks displayed lower haematocrit when body mass index was higher. Therefore, ingestion of conventional grains by parents resulted in chronic exposure to pesticide residues, even at low doses, and this had detrimental consequences on offspring. These results demonstrate a sublethal effect of pesticide residues through parental effects. The consequences of parental exposure on chicks might partly explain the decline in wild Grey partridge populations, which raises questions for avian conservation and demography if current agrosystem approaches are continued.

Consistent variations in personality traits and their potential for genetic improvement in biocontrol agents: Trichogramma evanescens as a case study.

Improvements in the biological control of agricultural pests require improvements in the phenotyping methods used by practitioners to select efficient biological control agent (BCA) populations in industrial rearing or field conditions. Consistent inter-individual variations in behaviour (i.e. animal personality) probably affect BCA efficiency, but have never been taken into account in the development of phenotyping methods, despite having characteristics useful for phenotyping: repeatable (by definition), often heritable, etc. We developed a video-tracking method targeting animal personality traits and evaluated the feasibility of its use for genetic improvement in the BCA Trichogramma evanescens, by phenotyping 1,049 individuals from 24 isogenic lines. We found consistent individual variations in boldness, activity and exploration. Personality differences between the 24 isogenic lines suggested a genetic origin of the variations in activity and exploration (broad-sense heritability estimates of 0.06 to 0.11) and revealed a trade-off between exploration and fecundity.