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International audience; Metacommunity structure reflects the interplay of various processes, including niche filtering, extinction/colonization and interspecific interactions. Spatial patterns of species distributions are often analyzed to infer these processes. However, such inferences rely on often unrealistic equilibrium assumptions, and remain ambiguous, as different processes can produce similar patterns. Temporal data may improve these inferences. For example, stochastic species turnover may occur in local communities, while, on the long run, temporal changes are kept within limits set by locally available niches. Our objective is to explore how the joint analysis of spatial and temporal patterns can clarify the contribution of different processes to metacommunity structure. We recorded the occurrences of 21 freshwater mollusc species, and environmental data, in 250 sites over 17 successive years in a network of ponds in Guadeloupe (Lesser Antilles). We analyzed variation in α and β-diversities in space and time, and used a joint-species distribution mode to characterize species–environment and species–species relationships. Local communities showed pronounced temporal variation reflecting both imperfect species detection and true stochastic species turnover. On the long term however, local communities were largely controlled by niche filtering along two main environmental gradients, one driven by site connectivity, the other by hydrological stability and aquatic vegetation. Two gastropod clades, caenogastropods and pulmonates, showed contrasted spatio–temporal distributions resulting from different responses to these gradients, and these distributions seemed little altered by interspecific competition. Our study illustrates the benefit of using spatiotemporal metacommunity data to discern long-term impacts of niche filtering and species interactions behind short-term stochasticity.

Although metacommunity ecology has been a major field of research in the last decades, with both conceptual and empirical outputs, the analysis of the temporal dynamics of metacommunities has only emerged recently and still consists mostly of repeated static analyses. Here we propose a novel analysis framework to assess metacommunity processes using path analyses of spatial and temporal diversity turnovers. We detail the principles and practical aspects of this framework and apply it to simulated datasets to illustrate its ability to decipher the respective contributions of entangled drivers of metacommunity dynamics. We then apply it to four real datasets. Empirical results support the view that metacommunity dynamics may be generally shaped by multiple ecological processes acting in concert, with environmental filtering being variable across both space and time. These results reinforce our call to go beyond static analyses of metacommunities that are blind to the temporal part of environmental variability. This article has been peer-reviewed and recommended by Peer Community in Ecology

WOS:000779278700001; International audience; Characterizing the diversity of demographic strategies among species can inform research in topics such as trait syndromes, community stability, coexistence, and ecological succession. However, this diversity can depend on the spatial scale considered: at the landscape scale, species often form metapopulations, that is sets of local, sometimes short-lived, populations, inhabiting discrete habitat patches. Metapopulation dynamics are most frequently analyzed in individual species or pairs of interacting species because of the large amount of data required for multiple species, and because species vary in their perceptions of what constitutes a favorable or unfavorable habitat. Here we evaluate, using a case study, whether a metapopulation model can be used to generate accurate estimates of demographic parameters and to describe the diversity of dynamics, responses to environment, and prospects of long-term persistence in a guild of species inhabiting a common fragmented landscape. We applied this approach to a guild of 22 mollusk species that inhabit freshwater habitats on two islands of Guadeloupe, to compare metapopulation dynamics among species. We analyzed a 15-year time series of occupancy records for 278 sites using a multistate occupancy model that estimated colonization and extinction rates as a function of site-specific and year-specific environmental covariates, then used model results to simulate future island metapopulation dynamics. Despite the diverse array of metapopulation trajectories—a mix of species with either stable, increasing, declining, or fluctuating metapopulations—and the inherent challenges associated with such data (e.g., imperfect detection, spatial and temporal heterogeneity), our model accurately captured among-patch variation in suitability for many mollusk taxa. The dynamics of rare species or species with habitat preferences not fully captured by the retained set of covariates were less well described. For several species, we detected a negative correlation between extinction and colonization. This variation in habitat suitability created species-specific extinction-resistant pockets in the landscape. Our comparative analysis also revealed that species had distinct strategies for metapopulation dynamics, such as “fast-turnover” species with both a high proportion of occupied sites and a high rate of site extinction in the landscape.

Plant sexual dimorphism is thought to evolve in response to sex-specific selection associated with competition for access to mates or resources, both of which may be density dependent. In wind-pollinated plants in particular, vegetative traits such as plant size and architecture may influence resource acquisition and both pollen dispersal and receipt, with potential conflict between these two components of fitness. We evaluated the role of plant density in shaping plant traits by measuring evolutionary responses in experimental populations of the sexually dimorphic wind-pollinated plant Mercurialis annua. After three generations of evolution, we observed divergence between high- and low-density populations in several vegetative traits, whereas there was no divergence for reproductive traits. A reversal in the direction of sexually dimorphic traits expressed in young plants evolved in both low- and high-density populations compared to the original population (stored as seeds). Compared to the source population, males at high density evolved to be taller when young, whereas at low density young females tended to become smaller. These results demonstrate that a simple change in plant density can induce age-dependent and sex-specific evolution in the ontogeny of vegetative organs, and illustrates the power of experimental evolution for investigating plant trait evolution.

Coexistence with related species poses evolutionary challenges to which populations may react in diverse ways. When exposed to similar environments, sympatric populations of two species may adopt similar phenotypic trait values. However, selection may also favour trait divergence as a way to reduce competition for resources or mates. The characteristics of external body parts, such as coloration and external morphology, are involved to varying degrees in intraspecific signalling as well as in the adaptation to the environment and consequently may be diversely affected by interspecific interactions in sympatry. Here, we studied the effect of sympatry on various colour and morphological traits in males and females of two related newt species Lissotriton helveticus and L. vulgaris. Importantly, we did not only estimate how raw trait differences between species respond to sympatry, but also the marginal responses after controlling for environmental variation. We found that dorsal and caudal coloration converged in sympatry, likely reflecting their role in adaptation to local environments, especially concealment from predators. In contrast, aspects of male and female ventral coloration, which harbours sexual signals in both species, diverged in sympatry. This divergence may reduce opportunities for interspecific sexual interactions and the associated loss of energy, suggesting reproductive character displacement (RCD). Our study emphasizes the contrasting patterns of traits involved in different functions and calls for the need to consider this diversity in evolutionary studies.

Understanding how environmental change affects genetic variances and covariances of reproductive traits is key to formulate firm predictions on evolutionary responses. This is particularly true for sex-specific variance in reproductive success, which has been argued to affect how populations can adapt to environmental change. Our current knowledge on the impact of environmental stress on sex-specific genetic architecture of fitness components is still limited and restricted to separate-sexed organisms. However, hermaphroditism is widespread across animals and may entail interesting peculiarities with respect to genetic constraints imposed on the evolution of male and female reproduction. We explored how food restriction affects the genetic variance-covariance (G) matrix of body size and reproductive success of the simultaneously hermaphroditic freshwater snail Physa acuta. Our results provide strong evidence that the imposed environmental stress elevated the opportunity for selection in both sex functions. However, the G matrix remained largely stable across the tested food treatments. Importantly, our results provide no support for cross-sex genetic correlations suggesting no strong evolutionary coupling of male and female reproductive traits. We discuss potential implications for the adaptation to changing environments and highlight the need for more quantitative genetic studies on male and female fitness components in simultaneous hermaphrodites. Fil: Janicke, Tim. Technische Universität Dresden; Alemania. Centre National de la Recherche Scientifique; Francia Fil: Chapuis, Elodie. Université de Montpellier; Francia Fil: Meconcelli, Stefania. Centre National de la Recherche Scientifique; Francia. Università di Torino; Italia Fil: Bonel, Nicolás. Centre National de la Recherche Scientifique; Francia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Bahía Blanca. Centro de Recursos Naturales Renovables de la Zona Semiárida. Universidad Nacional del Sur. Centro de Recursos Naturales Renovables de la Zona Semiárida; Argentina Fil: Delahaie, Boris. University of Cambridge; Estados Unidos Fil: David, Patrice. Centre National de la Recherche Scientifique; Francia

Cryptic species can present a significant challenge to the application of systematic and biogeographic principles, especially if they are invasive or transmit parasites or pathogens. Detecting cryptic species requires a pluralistic approach in which molecular markers facilitate the detection of coherent taxonomic units that can then be analyzed using various traits (e.g., internal morphology) and crosses. In asexual or self-fertilizing species, the latter criteria are of limited use. We studied a group of cryptic freshwater snails (genusGalba)from the family Lymnaeidae that have invaded almost all continents, reproducing mainly by self-fertilization and transmitting liver flukes to humans and livestock. We aim to clarify the systematics, distribution and phylogenetic relationships of these species with an integrative approach that includes morphology (shell and reproductive anatomy), molecular markers, wide-scale sampling across America, and data retrieved from GenBank (to include Old World samples). Our phylogenetic analysis suggests that the genusGalbaoriginated ca. 22 Myr ago and today comprises six clusters of species. Four of these clusters (G.truncatula,G.cubensis/viator,G.humilisandG.schirazensis) are morphologically cryptic and constitute species or species complexes with wide variation in their genetic diversity, geographic distribution and invasiveness. The other two clusters constitute a single species or a species complex (Galba cousini/meridensis) that demonstrate more geographically restricted distributions and exhibit an alternative morphology more phylogenetically derived than the cryptic one. Further genetic studies are required to clarify the status of bothG. cousini/meridensisandG.cubensis/viator. We emphasize that noGalbaspecies should be identified without molecular markers and that additional sampling is required, especially in North America, Eurasia and Africa to clarify remaining questions in systematics and biogeography. We also discuss several hypotheses that can explain crypsis inGalba, such as convergence and morphological stasis, and hypothesize a role for stabilizing selection in amphibious (rather than strictly freshwater) habitats.

We studied the population genetic structure of the freshwater snail Galba schirazensis (Küster, 1862), a potential vector of infectious diseases such as fascioliasis. Galba schirazensis has now a worldwide distribution but a poorly known origin because this species has been distinguished only recently from the morphologically similar and cosmopolitan Galba truncatula (O.F. Müller, 1774). We developed specific microsatellite markers and sequenced a mitochondrial gene (cytochrome oxidase subunit I (CO1)) to study individuals of G. schirazensis from the Old World and the New World. We found very low genetic diversity within populations, no heterozygotes, and marked population structure — a pattern observed in other highly selfing lymnaeid species with recently enlarged distributions as a result of biological invasions. The total lack of observed heterozygosity in the few populations of G. schirazensis that displayed some allelic diversity suggests high selfing rates. We also found that the center of diversity, and by extension the origin area of this species, should be found in the New World, whereas Old World populations should rather result from a recent introduction of a genetically uniform population. The microsatellite markers developed here will help to clarify the history of expansion of G. schirazensis and might help to understand its role as a potential vector of infectious diseases. Fil: Lounnas, Manon. Université de Montpellier. Maladies Infectieuses et Vecteurs: Ecologie, Génétique, Evolution et Contrôle; Francia. Centre National de la Recherche Scientifique. Institut de Recherche pour le Développement; Francia Fil: Correa, Ana Cristina. Centre National de la Recherche Scientifique. Institut de Recherche pour le Développement; Francia. Université de Montpellier. Maladies Infectieuses et Vecteurs: Ecologie, Génétique, Evolution et Contrôle; Francia Fil: Alda, Maria del Pilar. Centre National de la Recherche Scientifique. Institut de Recherche pour le Développement; Francia. Université de Montpellier. Maladies Infectieuses et Vecteurs: Ecologie, Génétique, Evolution et Contrôle; Francia. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - La Plata. Centro de Estudios Parasitológicos y de Vectores. Universidad Nacional de La Plata. Facultad de Ciencias Naturales y Museo. Centro de Estudios Parasitológicos y de Vectores; Argentina Fil: David, Patrice. Université de Montpellier. Centre d’Ecologie Fonctionnelle et d’Evolution; Francia Fil: Dubois, Marie Pierre. Université de Montpellier. Centre d’Ecologie Fonctionnelle et d’Evolution; Francia Fil: Calvopiña, Manuel. Universidad Central del Ecuador, Carrera de Medicina; Ecuador Fil: Caron, Yannick. Université de Liège; Bélgica Fil: Celi Erazo, Maritza. Universidad Central del Ecuador; Ecuador Fil: Bui, Dung Thi. Université de Liège; Bélgica Fil: Jarne, Philippe. Université de Montpellier. Centre d’Ecologie Fonctionnelle et d’Evolution; Francia Fil: Loker, Eric. University Of New Mexico. Department Of Biology; Estados Unidos Fil: Noya, Oscar. Universidad Central de Venezuela; Venezuela Fil: Rodríguez Hidalgo, Richar. Universidad Central del Ecuador; Ecuador Fil: Toty, Céline. Université de Montpellier. Maladies Infectieuses et Vecteurs: Ecologie, Génétique, Evolution et Contrôle; Francia Fil: Uribe, Nelson. Universidad Industrial Santander; Colombia Fil: Pointier, Jean Pierre. Université de Perpignan; Francia. Centre National de la Recherche Scientifique; Francia Fil: Hurtrez Boussès, Sylvie. Université de Montpellier. Maladies Infectieuses et Vecteurs: Ecologie, Génétique, Evolution et Contrôle; Francia

International audience; Biological invasions offer interesting situations for observing how novel interactions between closely related, formerly allopatric species may trigger phenotypic evolution in situ. Assuming that successful invaders are usually filtered to be competitively dominant, invasive and native species may follow different trajectories. Natives may evolve traits that minimize the negative impact of competition, while trait shifts in invasives should mostly reflect expansion dynamics, through selection for colonization ability and transiently enhanced mutation load at the colonization front. These ideas were tested through a large-scale common-garden experiment measuring life-history traits in two closely related snail species, one invasive and one native, co-occurring in a network of freshwater ponds in Guadeloupe. We looked for evidence of recent evolution by comparing uninvaded or recently invaded sites with long-invaded ones. The native species adopted a life history favoring rapid population growth (i.e., increased fecundity, earlier reproduction, and increased juvenile survival) that may increase its prospects of coexistence with the more competitive invader. We discuss why these effects are more likely to result from genetic change than from maternal effects. The invader exhibited slightly decreased overall performances in recently colonized sites, consistent with a moderate expansion load resulting from local founder effects. Our study highlights a rare example of rapid life-history evolution following invasion.

Batesian mimicry, with its highly colorful species and astonishing mimic-model resemblance, is a canonical example of evolution by natural selection. However, Batesian mimicry could also occur in inconspicuous species and rely on subtle resemblance. Although potentially widespread, such instances have been rarely investigated, such that the real frequency of Batesian mimicry has remained largely unknown. To fill this gap, we developed a new approach using deep learning to quantify the resemblance between putative mimics and models from photographs. We applied this method to quantify the frequency of Batesian mimicry in Western Palearctic snakes. Potential mimics were revealed by an excess of resemblance with sympatric venomous snakes compared to random expectations. We found that 8% of the non-venomous species were potential mimics, among which all were imperfect mimics. This study is the first to quantify the frequency of Batesian mimicry in a whole community of vertebrates, and shows that even concealed species can be reliably identified as potential models. Our approach should prove useful to detect mimicry in other communities, and more generally it highlights the benefits of deep learning for quantitative studies of phenotypic resemblance.

International audience; Sexual selection may contribute to the evolution of plant sexual dimorphism by favoring architectural traits in males that improve pollen dispersal to mates. In both sexes, larger individuals may be favored by allowing the allocation of more resources to gamete production (a ‘budget’ effect of size). In wind‐pollinated plants, large size may also benefit males by allowing the liberation pollen from a greater height, fostering its dispersal (a ‘direct’ effect of size). To assess these effects and their implications for trait selection, we measured selection on plant morphology in both males and females of the wind‐pollinated dioecious herb Mercurialis annua in two separate experimental common gardens at contrasting density. In both gardens, selection strongly favored males that disperse their pollen further. Selection for pollen production was observed in the high‐density garden only and was weak. In addition, male morphologies associated with increased mean pollen dispersal differed between the two gardens, as elongated branches were favored in the high‐density garden while shorter plants with longer inflorescence stalks were favored in the low‐density garden. Larger females were selected in both gardens. Our results point to the importance of both a direct effect of selection on male traits that affect pollen dispersal, and, to a lesser extent, a budget effect of selection on pollen production.

1. Heterozygosity fitness correlations (HFCs) have been used extensively to explore the impact of inbreeding on individual fitness. Initially, most studies used small panels of microsatellites, but more recently with the advent of next generation sequencing, large SNP datasets are becoming increasingly available and these provide greater power and precision to quantify the impact of inbreeding on fitness. 2. Despite the popularity of HFC studies, effect sizes tend to be rather small. One reason for this may be a low variation in inbreeding level across individuals. Using genetic markers, it is possible to measure variance in inbreeding through the strength of correlation in heterozygosity across marker loci, termed identity disequilibrium (ID). 3. ID can be quantified using the measure g2 which is also a central parameter in HFC theory that can be used within a wider framework to estimate the direct impact of inbreeding on both marker heterozygosity and fitness. However, no software exists to calculate g2 for large SNP datasets nor to implement this framework. 4. inbreedR is an R package that provides functions to calculate g2 based on microsatellite and SNP markers with associated p-values and confidence intervals. Within the framework of HFC theory, inbreedR also estimates the impact of inbreeding on marker heterozygosity and fitness. Moreover, we implemented easy-to-use simulations to explore the precision and magnitude of estimates based on different numbers of genetic markers. We hope this package will facilitate good practice in the analysis of HFCs and help to deepen our understanding of inbreeding effects in natural populations.

Basic models of mating-system evolution predict that hermaphroditic organisms should mostly either cross-fertilize, or self-fertilize, due to self-reinforcing coevolution of inbreeding depression and outcrossing rates. However transitions between mating systems occur. A plausible scenario for such transitions assumes that a decrease in pollinator or mate availability temporarily constrains outcrossing populations to self-fertilize as a reproductive assurance strategy. This should trigger a purge of inbreeding depression, which in turn encourages individuals to self-fertilize more often and finally to reduce male allocation. We tested the predictions of this scenario using the freshwater snail Physa acuta, a self-compatible hermaphrodite that preferentially outcrosses and exhibits high inbreeding depression in natural populations. From an outbred population, we built two types of experimental evolution lines, controls (outcrossing every generation) and constrained lines (in which mates were often unavailable, forcing individuals to self-fertilize). After ca. 20 generations, individuals from constrained lines initiated self-fertilization earlier in life and had purged most of their inbreeding depression compared to controls. However, their male allocation remained unchanged. Our study suggests that the mating system can rapidly evolve as a response to reduced mating opportunities, supporting the reproductive assurance scenario of transitions from outcrossing to selfing.

Sexual selection operates through consecutive episodes of selection that ultimately contribute to the observed variance in reproductive success between individuals. Understanding the relative importance of these episodes is challenging, particularly because the relevant postcopulatory fitness components are often difficult to assess. Here, we investigate different episodes of sexual selection on the male sex function, by assessing how (precopulatory) mating success, and (postcopulatory) sperm-transfer efficiency and sperm-fertilizing efficiency contribute to male reproductive success. Specifically, we used a transgenic line of the transparent flatworm, Macrostomum lignano, which expresses green fluorescent protein (GFP) in all cell types, including sperm cells, enabling in vivo sperm tracking and paternity analysis. We found that a large proportion of variance in male reproductive success arose from the postcopulatory episodes. Moreover, we also quantified selection differentials on 10 morphological traits. Testis size and seminal vesicle size showed significant positive selection differentials, which were mainly due to selection on sperm-transfer efficiency. Overall, our results demonstrate that male reproductive success in M. lignano is not primarily limited by the number of matings achieved, but rather by the ability to convert matings into successful fertilizations, which is facilitated by producing many sperm.

Sex allocation theory predicts that simultaneous hermaphrodites evolve to an evolutionary stable resource allocation, whereby any increase in investment to male reproduction leads to a disproportionate cost on female reproduction and vice versa. However, empirical evidence for sexual trade-offs in hermaphroditic animals is still limited. Here, we tested how male and female reproductive traits evolved under conditions of reduced selection on either male or female reproduction for 40 generations in a hermaphroditic snail. This selection favors a reinvestment of resources from the sex function under relaxed selection toward the other function. We found no such evolutionary response. Instead, juvenile survival and male reproductive success significantly decreased in lines where selection on the male function (i.e., sexual selection) was relaxed, while relaxing selection on the female function had no effect. Our results suggest that most polymorphisms under selection in these lines were not sex-antagonistic. Rather, they were deleterious mutations affecting juvenile survival (thus reducing both male and female fitness) with strong pleiotropic effects on male success in a sexual selection context. These mutations accumulated when sexual selection was relaxed, which supports the idea that sexual selection in hermaphrodites contributes to purge the mutation load from the genome as in separate-sex organisms.

Modeling the dynamics of competition and coexistence between species is cru- cial to predict long-term impacts of invasive species on their native congeners. However, natu- ral environments are often fragmented and variable in time and space. In such contexts, regional coexistence depends on complex interactions between competition, niche differentia- tion and stochastic colonization–extinction dynamics. Quantifying all these processes at land- scape scale has always been a challenge for ecologists. We propose a new statistical framework to evaluate metapopulation parameters (colonization and extinction) in a two-species system and how they respond to environmental variables and interspecific competition. It requires spatial surveys repeated in time, but does not assume demographic equilibrium. We apply this model to a long-term survey of two snails inhabiting a network of freshwater habitats in the West Indies. We find evidence of reciprocal competition affecting colonization or extinction rates, modulated by species-specific sensitivity to environmental variables. Simulations using model estimates allow us to predict species dynamics and explore the role of various coexis- tence mechanisms described by metacommunity theory in our system. The two species are pre- dicted to stably coexist, because niche partitioning, source–sink dynamics and interspecific differences in extinction–colonization parameters all contribute to reduce the negative impacts of competition. However, none of these mechanisms is individually essential. Regional coexis- tence is primarily facilitated by transient co-occurrence of the two species within habitat patches, a possibility generally not considered in theoretical metacommunity models. Our framework is general and could be extended to guilds of several competing species.

Theory and empirical data showed that two processes can boost selection against deleterious mutations, thus facilitating the purging of the mutation load: inbreeding, by exposing recessive deleterious alleles to selection in homozygous form, and sexual selection, by enhancing the relative reproductive success of males with small mutation loads. These processes tend to be mutually exclusive because sexual selection is reduced under mating systems that promote inbreeding, such as self‐fertilization in hermaphrodites. We estimated the relative efficiency of inbreeding and sexual selection at purging the genetic load, using 50 generations of experimental evolution, in a hermaphroditic snail (Physa acuta). To this end, we generated lines that were exposed to various intensities of inbreeding, sexual selection (on the male function), and nonsexual selection (on the female function). We measured how these regimes affected the mutation load, quantified through the survival of outcrossed and selfed juveniles. We found that juvenile survival strongly decreased in outbred lines with reduced male selection, but not when female selection was relaxed, showing that male‐specific sexual selection does purge deleterious mutations. However, in lines exposed to inbreeding, where sexual selection was also relaxed, survival did not decrease, and even increased for self‐fertilized juveniles, showing that purging through inbreeding can compensate for the absence of sexual selection. Our results point to the further question of whether a mixed strategy combining the advantages of both mechanisms of genetic purging could be evolutionary stable. Fil: Noël, Elsa. Institut National de la Recherche Agronomique; Francia. Centre National de la Recherche Scientifique; Francia Fil: Fruitet, Elise. University of Amsterdam; Países Bajos. Max Planck Institute for Chemical Ecology; Alemania. Centre National de la Recherche Scientifique; Francia Fil: Lelaurin, Dennyss. Centre National de la Recherche Scientifique; Francia Fil: Bonel, Nicolás. Universidad Nacional del Sur; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Centre National de la Recherche Scientifique; Francia Fil: Ségard, Adeline. Centre National de la Recherche Scientifique; Francia Fil: Sarda, Violette. Centre National de la Recherche Scientifique; Francia Fil: Jarne, Philippe. Centre National de la Recherche Scientifique; Francia Fil: David, Patrice. Centre National de la Recherche Scientifique; Francia

[Departement_IRSTEA]Territoires [TR1_IRSTEA]SEDYVIN; International audience; Community genetics aims at understanding how within-species variation, species diversity and environmental factors interact to shape community assembly. An approach that emerged a few years ago has been to quantify the correlation between the neutral genetic diversity of a focal species and species diversity of the surrounding community (species–genetic diversity correlations, or SGDCs). We here review this approach and discuss its interpretative framework in a community ecology context. First, we show that the case for mostly positive SGDCs is probably overstated due to publication bias – only 11% are significantly positive, a fraction comparable to the significantly negative ones. This suggests that variation in area and connectivity among habitat patches, theoretically leading to positive SGDCs, is not the only factor affecting SGDCs. Second, building upon previous contributions, we propose a general framework to identify the multiple factors underpinning SGDCs, and argue that it will help deepen our understanding of community assembly, especially with regard to the ecological factors playing at metacommunity scale. Our framework distinguishes between site and community factors which can affect SGDCs either positively or negatively, depending on whether the focal species and the rest of the community are similar or dissimilar, in terms of realized niches and dispersal abilities. Empirical studies should thus go beyond simply computing SGDCs, and we provide statistical methods (e.g. structural equation modelling) to decompose SGDCs into the multiple contributions of site and community factors. As an example, we use a published dataset (freshwater snail metacommunity), and show how the role of focal population size on SGDCs had hitherto not been detected. We further discuss how considering several focal species and various delimitations of the community may help one to identify clusters of ecologically similar species. We eventually highlight the benefit that SGDC studies would get from integrating beta-diversities.

The differential plasticity hypothesis suggests that sexual dimorphism in dioecious plants could evolve in response to sex-specific resource requirements for reproduction (i.e., high carbon requirements for ovules and high nitrogen demands for pollen). When resources become limiting during growth, males and females should, therefore, adjust their allocation to resource-harvesting organs differently. To investigate the potential for plants to respond to resource limitation late in life and to test the differential plasticity hypothesis, we grew male and female individuals of the annual wind-pollinated plant Mercurialis annua in a common garden. Late in the growth season, we simulated a change in competition by decreasing plant density in half of the replicates. We measured both allocation to vegetative and reproductive traits and analyzed the relative allocation to reproduction vs. growth. Males and females differentially adjusted their resource allocation in response to varying plant densities, despite the fact that they were reproductively mature. Males maintained the same relative allocation of resource to reproductive vs. vegetative tissues at both densities. In contrast, females reduced vegetative growth proportionally less than seed production at the higher density. Our results highlight the dynamic nature of allocation decisions taken by plants, which respond quickly and in a sexually dimorphic way to changes in their competitive circumstances. The existence of resource ‘currencies’ limiting male and female functions differently have potentially led to the evolution of sex-specific strategies of resource acquisition and deployment, with females conserving resources for vegetative organs to ensure their future carbon-rich reproduction. Electronic supplementary material The online version of this article (doi:10.1007/s00442-017-3966-5) contains supplementary material, which is available to authorized users.

Explaining the strong variation in lifespan among organisms remains a major challenge in evolutionary biology. Whereas previous work has concentrated mainly on differences in selection regimes and selection pressures, we hypothesize that differences in genetic drift may explain some of this variation. We develop a model to formalize this idea and show that the strong positive relationship between lifespan and genetic diversity predicted by this model indeed exists among populations of Daphnia magna, and that ageing is accelerated in small populations. Additional results suggest that this is due to increased drift in small populations rather than adaptation to environments favoring faster life histories. First, the correlation between genetic diversity and lifespan remains significant after statistical correction for potential environmental covariates. Second, no trade-offs are observed; rather, all investigated traits show clear signs of increased genetic load in the small populations. Third, hybrid vigor with respect to lifespan is observed in crosses between small but not between large populations. Together, these results suggest that the evolution of lifespan and ageing can be strongly affected by genetic drift, especially in small populations, and that variation in lifespan and ageing may often be nonadaptive, due to a strong contribution from mutation accumulation.

Sexual selection operates on a sequence of events, from mating to offspring production. Which stages in this sequence undergo stronger selection, especially the relative importance of pre- versus postcopulatory processes, are intensely debated issues. Unequal siring success among mates of polyandrous females is classically taken as evidence for a large contribution of postcopulatory processes to the variance in male reproductive success (var(RSm )). However, paternity skews also depend on the timing and number of copulations, a source of variation that should be considered precopulatory rather than postcopulatory. We develop a method for decomposing var(RSm ) accounting for copulatory activity and apply it to experimental mating groups of the snail Physa acuta. In our experiment, 40% of var(RSm ) emerges at the precopulatory stage, only half of which depends on variation in mating success (number of partners). Ignoring copulation characteristics can therefore lead to severe underestimation of precopulatory sexual selection. Moreover, although only 36% of var(RSm ) arises at the postcopulatory stage, this is when sexual selection on body weight mostly occurs. Finally, trade-offs were detected between different components of precopulatory success, whereas pre- and postcopulatory success appear independent. Our study opens the way to a detailed quantitative understanding of sexual selection in polyandrous species.

Quantifying metapopulation dynamics is a challenging task. Difficulties particularly arise in species that possess unobservable resistance forms that bias the estimation of colonization and persistence rates. Here, we develop a general multistate occupancy model that allows estimation of species persistence for both normal and resistant forms, even when the latter are not detectable. We apply this model to an 11-year data set on the tropical freshwater snail Drepanotrema depressissimum in a network of 229 ponds. These ponds frequently dry out, and the snails can persist by aestivating in the ground, where they are not detected. Our model adequately captures this feature because it infers a high persistence rate for the resistant form, while models ignoring aestivation tend to overestimate extinction and colonization. In addition, we find that, surprisingly, colonization and persistence are even higher in sites prone to desiccation and during years with low rainfall than in more humid sites and years, suggesting that D. depressissimum favors unstable sites where competitors are rare and where it can rely on aestivation to persist. Our model has the potential to provide valuable insights into the metapopulation dynamics of many species that otherwise could hardly be studied because of the existence of undetectable life forms.

Summary 1.How species co-exist is a central question in ecology. Niche differentiation can prevent two species from excluding each other by competition. However, this interaction can vary in space because of internal factors such as intraspecific genetic variation, or external ones such as the presence of a third species. While these effects have been repeatedly observed, their joint effect on the outcome of competition has rarely been considered. 2.We experimentally investigated how a dominant ecosystem engineer (the shrub Thymus vulgaris) affects interactions between two co-occurring and closely related annuals (Medicago minima and M. rigidula). We ask first if the outcome of their interaction depends on the genetic identity of the competitors, and in particular if intraspecific interaction differs between genetically related versus unrelated conspecifics. Second, we ask if co-existence of the two species is modified by the presence of thyme plants altering local soil conditions. 3.We grew genotypes collected in nature of both annual species in inter- and intraspecific competition experiments using soil collected directly underneath thyme plant (thyme soil), and soil collected away from thyme plants (no-thyme soil). 4.We found that thyme soil affected interactions in a way that shifted the species’ rank. Crucial for this result was the division of intraspecific competition between genetically related and unrelated individuals. Intraspecific competition was significantly less detrimental to kin than to unrelated conspecifics. In highly structured plant populations, such effects could favor local dominance of a patch by one species. However this kin effect occurred for each species only in its preferred soil (thyme versus no-thyme), promoting a stabilizing co-existence mechanism at the landscape level. 5.Synthesis: This study shows that the importance of intraspecific competition relative to interspecific competition can be highly dependent on the genotype identity of intraspecific competitors and of the local environment in which the interaction occurs. The local environment can itself be modified by the presence of a third species in the community. These results emphasize how, in order to understand the overall problem of species co-existence, it can be insightful to divide this into smaller local scale problems of co-existence. This article is protected by copyright. All rights reserved.

Environmental changes, such as climate change, lead to the opening of new niches. In such situations, species that adapt to new niches can survive and/or expand their ranges. However, gene flow can hamper genetic adaptation to new environments. Alternatively, recent models have highlighted the importance of phenotypic plasticity in tracking environmental change. Here, we investigate whether phenotypic plasticity or genetic evolution (or both) allows an amphibian species to exploit two divergent climatic niches. In the Mediterranean region, the parsley frog Pelodytes punctatus breeds both in spring, as do most other species, and in autumn, a temporal niche not exploited by most other species, but which may become increasingly important with global warming. Conditions of development are dramatically different between the two seasons and deeply impact tadpole life-history traits. To determine whether these temporal niches are exploited by two genetically differentiated subpopulations, or whether the bimodal phenology arises in a panmictic population displaying plastic life-history traits, we use two complementary approaches. We measure both molecular genetic differentiation and quantitative-trait differentiation between spring and autumn cohorts, using microsatellites and common garden experiments, respectively. Seasonal cohorts were not genetically differentiated and differences in tadpole life history between cohorts were not maintained in laboratory conditions. We conclude that phenotypic plasticity, rather than genetic adaptation, allows Parsley frog to exploit two contrasting temporal niches.

Assortative mating by size has been argued to be widespread in the animal kingdom. However, the strength of size-assortative mating is known to vary considerably between species and the underlying mechanisms promoting this inter-specific variation remain largely unexplored. Size-assortative mating has been proposed to be particularly strong in simultaneous hermaphrodites, i.e. organisms that produce male and female gametes at the same time. Here, we build on this hypothesis by arguing that size-assortative mating mediated by sexual selection is generally stronger in reciprocally mating hermaphrodites compared with unilaterally mating species and separate-sexed organisms. We report a series of empirical tests suggesting that size-assortative mating in the unilaterally copulating freshwater snail Physa acuta is caused by spatial clustering of similar-sized individuals and not by mate choice. In addition, we present a meta-analysis testing, for the first time, the hypothesis that sexual selection-mediated size-assortative mating is stronger in reciprocally copulating simultaneous hermaphrodites. Overall, we found significant size-assortative mating across 18 tested species and substantial inter-specific variation. Importantly, part of this variation can be explained by mating type, providing support for the hypothesis that size-assortative mating is stronger in reciprocally mating hermaphrodites compared with unilaterally mating species. We highlight potential pitfalls when testing for sexual selection-mediated size-assortative mating and discuss the need for more experimental and comparative approaches in order to resolve the observed variation in the strength of size-assortative mating among species., Centro de Estudios Parasitológicos y de Vectores

One of the most general patterns in sexual selection is stronger selection on mating activity in males than in females. This asymmetry is thought to result from the higher energetic cost of producing one female compared to one male gamete (anisogamy). However, most studies focused on gonochoric species with strong sexual dimorphism, in which males and females are necessarily under different selection regimes. The question remains whether anisogamy alone would suffice to produce such differences. In simultaneous hermaphrodites one can compare sexual selection on the male and female functions in the absence of sexual dimorphism. Here we quantify sexual selection in the hermaphroditic freshwater snail Physa acuta under laboratory conditions. We combine exhaustive behavioral records of mating activity in mating groups and molecular paternity assignment to measure the mating success and reproductive success of 120 individuals. Our results validate the prediction of stronger selection to gain mating partners in the male than in the female function. Moreover, we did not detect cross-sex effects on fitness, or correlations between male and female production of offspring over the course of our experiment. We conclude that with respect to sexual selection P. acuta is comparable to gonochorists, confirming that anisogamy is a sufficient explanation for the differences in sexual selection regimes between sexes.

Genome-wide heterozygosity inferred from neutral markers such as microsatellites is often expected to (i) reflect individual inbreeding and (ii) covary positively with fitness, generating positive heterozygosity-fitness correlations (HFCs). The often forgotten other end of the inbreeding-outbreeding continuum is outbreeding depression: past a certain degree of heterozygosity, heterozygotes tend to have lower fitness than homozygotes. Outbreeding depression arises from the breakup of co-adapted gene complexes and/or the introgression of nonlocally adapted genes. Provided that a correlation in heterozygosity exists across loci, outbreeding depression will be reflected in negative HFCs. In this issue, Olano-Marin et al. (2011a) describe negative heterozygosity-fitness correlations (HFCs) in blue tits Cyanistes caeruleus (Fig. 1), whereby heterozygosity has a significant, negative effect on female hatching success and recruitment. This study, together with a similar study by the same authors published in Evolution (Olano-Marin et al. 2011b), forms an original contribution in two respects. First, in the same population, positive and negative HFCs were recorded, revealing both inbreeding and outbreeding depression depending on the trait studied (whereby both processes were reliant on unknown, and possibly different, sets of coding loci). Second, a large number of microsatellite markers were split into two functional groups: microsatellite markers were either designed using zebra finch expressed sequence tags (ESTs) or derived using traditional cloning methods and presumed to be neutral. Contrasting large classes of loci and their varying levels of polymorphism, rather than looking for one locus that would stand out among tens of randomly selected markers, pave the way for a more elegant and powerful approach to explore how HFCs vary across traits and among regions of the genome. [Figure: see text].

Genomic scans of multiple populations often reveal marker loci with greatly increased differentiation between populations. Often this differentiation coincides in space with contrasts in ecological factors, forming a genetic–environment association (GEA). GEAs imply a role for local adaptation, and so it is tempting to conclude that the strongly differentiated markers are themselves under ecologically based divergent selection, or are closely linked to loci under such selection. Here, we highlight an alternative and neglected explanation: intrinsic (i.e. environment-independent) pre- or post-zygotic genetic incompatibilities rather than local adaptation can be responsible for increased differentiation. Intrinsic genetic incompatibilities create endogenous barriers to gene flow, also known as tension zones, whose location can shift over time. However, tension zones have a tendency to become trapped by, and therefore to coincide with, exogenous barriers due to ecological selection. This coupling of endogenous and exogenous barriers can occur easily in spatially subdivided populations, even if the loci involved are unlinked. The result is that local adaptation explains where genetic breaks are positioned, but not necessarily their existence, which can be best explained by endogenous incompatibilities. More precisely, we show that (i) the coupling of endogenous and exogenous barriers can easily occur even when ecological selection is weak; (ii) when environmental heterogeneity is fine-grained, GEAs can emerge at incompatibility loci, but only locally, in places where habitats and gene pools are sufficiently intermingled to maintain linkage disequilibria between genetic incompatibilities, local-adaptation genes and neutral loci. Furthermore, the association between the locally adapted and intrinsically incompatible alleles (i.e. the sign of linkage disequilibrium between endogenous and exogenous loci) is arbitrary and can form in either direction. Reviewing results from the literature, we find that many predictions of our model are supported, including endogenous genetic barriers that coincide with environmental boundaries, local GEA in mosaic hybrid zones, and inverted or modified GEAs at distant locations. We argue that endogenous genetic barriers are often more likely than local adaptation to explain the majority of Fst-outlying loci observed in genome scan approaches – even when these are correlated to environmental variables. (Resume d'auteur)

Quantifying the impacts of inbreeding and genetic drift on fitness traits in fragmented populations is becoming a major goal in conservation biology. Such impacts occur at different levels and involve different sets of loci. Genetic drift randomly fixes slightly deleterious alleles leading to different fixation load among populations. By contrast, inbreeding depression arises from highly deleterious alleles in segregation within a population and creates variation among individuals. A popular approach is to measure correlations between molecular variation and phenotypic performances. This approach has been mainly used at the individual level to detect inbreeding depression within populations and sometimes at the population level but without consideration about the genetic processes measured. For the first time, we used in this study a molecular approach considering both the interpopulation and intrapopulation level to discriminate the relative importance of inbreeding depression vs. fixation load in isolated and non-fragmented populations of European tree frog (Hyla arborea), complemented with interpopulational crosses. We demonstrated that the positive correlations observed between genetic heterozygosity and larval performances on merged data were mainly caused by co-variations in genetic diversity and fixation load among populations rather than by inbreeding depression and segregating deleterious alleles within populations. Such a method is highly relevant in a conservation perspective because, depending on how populations lose fitness (inbreeding vs. fixation load), specific management actions may be designed to improve the persistence of populations.

In hermaphrodites, traits that influence the selfing rate can coevolve with inbreeding depression, leading to the emergence of evolutionary syndromes. Theory predicts a negative correlation between inbreeding depression and selfing rate across species. This prediction has only been examined and validated in vascular plants. Furthermore, selfing rates are often influenced by environmental conditions (e.g., lack of mates or pollinators), and species are predicted to evolve mechanisms to buffer this variation. We extend previous studies of mating-system syndromes in two ways. First, we assembled a new dataset on Basommatophoran snails (17 species, including new data on 12 species). Second, we measured how species responded to variation in mate availability. Specifically, we quantified the waiting time before selfing (i.e., how long the onset of reproduction is delayed in the absence of mates). Selfing rates were negatively correlated with both inbreeding depression and the waiting time. Species with stronger inbreeding depression exhibited longer waiting times. These patterns obtained on Basommatophorans still hold when including eight other hermaphroditic animals. Our results support the hypothesis that selection drives the evolution of mating-system syndromes in animals. The reaction norm of selfing rates to mate availability is a key target of natural selection in this context. © 2011 The Author(s). Evolution© 2011 The Society for the Study of Evolution.

The detrimental effects of genetic erosion on small isolated populations are widely recognized contrary to their interactions with environmental changes. The ability of genotypes to plastically respond to variability is probably essential for the persistence of these populations. Genetic erosion impact may be exacerbated if inbreeding affects plastic responses or if their maintenance were at higher phenotypic costs. To understand the interplay 'genetic erosion-fitness-phenotypic plasticity', we experimentally compared, in different environments, the larval performances and plastic responses to predation of European tree frogs (Hyla arborea) from isolated and connected populations. Tadpoles from isolated populations were less performant, but the traits affected were environmental dependant. Heterosis observed in crosses between isolated populations allowed attributing their low fitness to inbreeding. Phenotypic plasticity can be maintained in the face of genetic erosion as inducible defences in response to predator were identical in all populations. However, the higher survival and developmental costs for isolated populations in harsh conditions may lead to an additional fitness loss for isolated populations.

Numerous models have been designed to understand how dispersal ability evolves when organisms live in a fragmented landscape. Most of them predict a single dispersal rate at evolutionary equilibrium, and when diversification of dispersal rates has been predicted, it occurs as a response to perturbation or environmental fluctuation regimes. Yet abundant variation in dispersal ability is observed in natural populations and communities, even in relatively stable environments. We show that this diversification can operate in a simple island model without temporal variability: disruptive selection on dispersal occurs when the environment consists of many small and few large patches, a common feature in natural spatial systems. This heterogeneity in patch size results in a high variability in the number of related patch mates by individual, which, in turn, triggers disruptive selection through a high per capita variance of inclusive fitness. Our study provides a likely, parsimonious and testable explanation for the diversity of dispersal rates encountered in nature. It also suggests that biological conservation policies aiming at preserving ecological communities should strive to keep the distribution of patch size sufficiently asymmetric and variable.

We characterized 15 new variable microsatellites in the freshwater snail Pseudosuccinea (Lymnaea) columella, as well as conditions for multiplexing and simultaneously genotyping sets of loci. Two to six alleles were detected per locus over the six populations studied. Gene diversity ranged from 0.000 to 0.498, but essentially no heterozygous individuals were observed. This resulted in extremely high F(IS) estimates, and therefore high selfing rates. The F(ST) estimates ranged from 0.18 to 1 among populations, but was generally high. These markers will constitute efficient tools for investigating the population structure of this invasive species. Cross-species amplification was on the whole unsuccessful.

Inbreeding depression, one of the main factors driving mating system evolution, can itself evolve as a function of the mating system (the genetic purging hypothesis). Classical models of coevolution between mating system and inbreeding depression predict negative associations between inbreeding depression and selfing rate, but more recent approaches suggest that negative correlations should usually be too weak or transient to be detected within populations. Empirical results remain unclear and restricted to plants. Here, we evaluate, for the first time, the within-population genetic correlation between inbreeding depression and a trait that controls the amount of self-fertilization (the waiting time) in a self-fertile hermaphroditic animal, the freshwater snail Physa acuta. Using a large quantitative-genetic design (36 grand-families and 348 families), we observe abundant within-population family-level genetic variation for both inbreeding depression (estimated for survival, fecundity, and size) and the degree of behavioral selfing avoidance. However, we detected no correlation between waiting time and inbreeding depression across families. In agreement with recent models, this result shows that mutational variance rather than differential purging accounts for most of the genetic variance in inbreeding depression within a population.

Because domestication rarely leads to speciation, domesticated populations often hybridize with wild relatives when they occur in close proximity. Little work has focused on this question in clonally propagated crops. If selection on the capacity for sexual reproduction has been relaxed, these crops would not be expected to hybridize with their wild relatives as frequently as seed-propagated crops. Cassava is one of the most important clonally propagated plants in tropical agriculture. Gene flow between cassava and wild relatives has often been postulated, but never demonstrated in nature. We studied a population of a wild Manihot sp. in French Guiana, which was recently in contact with domesticated cassava, and characterized phenotypes (10 morphological traits) and genotypes (six microsatellite loci) of individuals in a transect parallel to the direction of hypothesized gene flow. Wild and domesticated populations were strongly differentiated at microsatellite loci. We identified many hybrids forming a continuum between these two populations, and phenotypic variation was strongly correlated with the degree of hybridization as determined by molecular markers. Analysis of linkage disequilibrium and of the diversity of hybrid pedigrees showed that hybridization has gone on for at least three generations and that no strong barrier prevents admixture of the populations. Hybrids were more heterozygous than either wild or domesticated individuals, and phenotypic comparisons suggested heterosis in vegetative traits. Our results also suggest that this situation is not uncommon, at least in French Guiana, and demonstrate the need for integrated management of wild and domesticated populations even in clonally propagated crops.

Invasive species are often said to be r-selected. However, invaders must sometimes compete with related resident species. In this case invaders should present combinations of life-history traits that give them higher competitive ability than residents, even at the expense of lower colonization ability. We test this prediction by comparing life-history traits among four fruit fly species, one endemic and three successive invaders, in La Reunion Island. Recent invaders tend to produce fewer, but larger, juveniles, delay the onset but increase the duration of reproduction, survive longer, and senesce more slowly than earlier ones. These traits are associated with higher ranks in a competitive hierarchy established in a previous study. However, the endemic species, now nearly extinct in the island, is inferior to the other three with respect to both competition and colonization traits, violating the trade-off assumption. Our results overall suggest that the key traits for invasion in this system were those that favoured competition rather than colonization. (Resume d'auteur)

Population genetic studies are efficient for inferring the invasion history based on a comparison of native and invasive populations, especially when conducted at species scale. An expected outcome in invasive populations is variability loss, and this is especially true in self-fertilizing species. We here focus on the self-fertilizing Pseudosuccinea columella, an invasive hermaphroditic freshwater snail that has greatly expanded its geographic distribution and that acts as intermediate host of Fasciola hepatica, the causative agent of human and veterinary fasciolosis. We evaluated the distribution of genetic diversity at the largest geographic scale analysed to date in this species by surveying 80 populations collected during 16 years from 14 countries, using eight nuclear microsatellites and two mitochondrial genes. As expected, populations from North America, the putative origin area, were strongly structured by selfing and history and harboured much more genetic variability than invasive populations. We found high selfing rates (when it was possible to infer it), none-to-low genetic variability and strong population structure in most invasive populations. Strikingly, we found a unique genotype/haplotype in populations from eight invaded regions sampled allover the world. Moreover, snail populations resistant to infection by the parasite are genetically distinct from susceptible populations. Our results are compatible with repeated introductions in South America and flash worldwide invasion by this unique genotype/haplotype. Our study illustrates the population genetic consequences of bio- logical invasion in a highly selfing species at very large geographic scale. We discuss how such a large-scale flash invasion may affect the spread of fasciolosis

National audience; Because mating systems affect population genetics and ecology, they are expected to impact the molecular evolution of species. Self-fertilizing species experience reduced effective population size, recombination rates, and heterozygosity, which in turn should decrease the efficacy of natural selection, both adaptive and purifying, and the strength of meiotic drive processes such as GC-biased gene conversion. The empirical evidence is only partly congruent with these predictions, depending on the analyzed species, some, but not all, of the expected effects have been observed. One possible reason is that self-fertilization is an evolutionary dead-end, so that most current selfers recently evolved self-fertilization, and their genome has not yet been strongly impacted by selfing. Here, we investigate the molecular evolution of two groups of freshwater snails in which mating systems have likely been stable for several millions of years. Analyzing coding sequence polymorphism, divergence, and expression levels, we report a strongly reduced genetic diversity, decreased efficacy of purifying selection, slower rate of adaptive evolution, and weakened codon usage bias/GC-biased gene conversion in the selfer Galba compared with the outcrosser Physa, in full agreement with theoretical expectations. Our results demonstrate that self-fertilization, when effective in the long run, is a major driver of population genomic and molecular evolutionary processes. Despite the genomic effects of selfing, Galba truncatula seems to escape the demographic consequences of the genetic load. We suggest that the particular ecology of the species may buffer the negative consequences of selfing, shedding new light on the dead-end hypothesis.

Spatial patterns of biological diversity have been extensively studied in ecology and population genetics, because they reflect the forces acting on biodiversity. A growing number of studies have found that genetic (within-species) and species diversity can be correlated in space (the so-called species-gene diversity correlation [SGDC]), which suggests that they are controlled by nonindependent processes. Positive SGDCs are generally assumed to arise from parallel responses of genetic and species diversity to variation in site size and connectivity. However, this argument implicitly assumes a neutral model that has yet to be developed. Here, we build such a model to predict SGDC in a metacommunity. We describe how SGDC emerges from competition within sites and variation in connectivity and carrying capacity among sites. We then introduce the formerly ignored mutation process, which affects genetic but not species diversity. When mutation rate is low, our model confirms that variation in the number of migrants among sites creates positive SGDCs. However, when considering high mutation rates, interactions between mutation, migration, and competition can produce negative SGDCs. Neutral processes thus do not always contribute positively to SGDCs. Our approach provides empirical guidelines for interpreting these novel patterns in natura with respect to evolutionary and ecological forces shaping metacommunities.

International audience; Sexual selection is a potent evolutionary force that has been shown to vary in strength and direction depending on demographic factors such as density and sex ratio. However, the effect of other environmental factors on the mode of sexual selection remains largely unexplored. Here, we tested experimentally how food restriction affects the potential for sexual selection in the male and the female sex function of the simultaneously hermaphroditic freshwater snail Physa acuta. We manipulated food availability and compared Bateman's metrics of sexual selection between groups of five well-fed and five food-restricted snails. Food-restricted individuals had a reduced female reproductive output, suggesting that we successfully manipulated the reproductive resources. Importantly, food restriction reduced the male opportunity for sexual selection (in terms of a lowered variance in male mating success) and led to diminishing returns of mating in both sexes (in terms of nonsignificant Bateman gradients). Furthermore, we observed significant changes in the relative contribution of different fitness components, suggesting stronger postcopulatory selection in the male sex role and stronger fecundity selection in the female sex role under restricted food conditions. This study highlights the need to incorporate ecological factors to better understand how sexual selection operates in the wild.

1. Biological invasions have profound effects on community structure. The community composition following invasions can be influenced by the habitat diversity and the species' responses to abiotic factors. 2. We evaluated the tolerance to climatic factors and analysed the field distribution of four polyphagous fruit flies (Diptera: Tephritidae) of La Reunion Island (three exotic species that successively invaded the island and the endemic species Ceratitis catoirii) in order to evaluate the opportunities of coexistence by niche differentiation. 3. Atmospheric humidity and immersion in water in the laboratory greatly influence the survival of fruit fly pupae. While C. catoirii and C. rosa are very sensitive to desiccation, C. capitata and especially Bactrocera zonata are relatively tolerant. B. zonata also tolerated immersion in water much longer than did C. rosa and C. catoirii, that in turn were more resistant than C. capitata. Overall, field distributions agree with the predictions based on this study of humidity combined with previous data on the effects of temperature. 4. Climatic niche partitioning promotes coexistence between some but not all pairs of invasive species. Thus, C. rosa can coexist with both C. capitata and B. zonata at the regional scale, while climatic niches are not different enough to promote coexistence of the latter two species. The endemic species has no private climatic niche either and this now very rare species could be in the process of extinction. 5. By promoting coexistence or not, climatic diversity in invaded areas can directly affect the community composition following invasions. (Resume d'auteur)

Correspondance: Benoit.Facon@unil.ch; International audience; Studies of bioinvasions have revealed various strategies of invasion, depending on the ecosystem invaded and the alien species concerned. Here, we consider how migration (as a demographic factor), as well as ecological and evolutionary changes, affect invasion success. We propose three main theoretical scenarios that depend on how these factors generate the match between an invader and its new environment. Our framework highlights the features that are common to, or differ among, observed invasion cases, and clarifies some general trends that have been previously highlighted in bioinvasions. We also suggest some new directions of research, such as the assessment of the time sequence of demographic, genetic and environmental changes, using detailed temporal surveys

We study the influence of mate availability on the mating behavior of the self-fertile, preferentially outcrossing freshwater snail Physa acuta. Previous optimization theory indicated that mating system interacts with life-history traits to influence the age at first reproduction, providing three testable predictions. First, isolated individuals should reproduce later than individuals with available mates in the expectancy of finding a partner and avoiding the cost of inbreeding. Second, resource reallocation to future fecundity is needed for such reproductive delays to evolve. Third, the reproductive delay can be optimized with respect to life-history traits (e.g., survival, growth) and the mating system (inbreeding depression). Our results largely validate these predictions. First, reproduction is significantly delayed in isolated individuals ("selfers") as compared with individuals frequently exposed to mates ("outcrossers"). Second, delayed reproduction is associated with reallocation to future growth, survival, and fecundity, although fecundity is also affected by the mating system (selfing vs. outcrossing). Third, the reproductive delay found (approximately 2 wk) is consistent with quantitative predictions from optimization models. The delay is largely heritable, which might be partly explained by among-family differences in the amount of inbreeding depression (mating system) but not growth or survival.

Resource allocation is thought to play a key role in the coexistence of different sexual morphs within hermaphroditic species. Indeed, most models assume that sexual functions are subject to a balance between reproductive advantage and energetic cost. Various types of cost (e.g. organ construction, maintenance and utilization) and levels of trade-off (physiological and genetic) may be considered. We here examine physiological and genetic costs of phallus construction and maintenance in Bulinus truncatus, a snail species in which aphallic individuals (without phallus) coexist with regular hermaphrodites. We use a quantitative genetic design involving 37 inbred lines (four populations) known to produce different proportions of aphallics, to test for the existence of genetic and nongenetic correlations between aphally and a range of life-history traits over the totality of the life cycle. Our results show that aphallic and euphallic individuals of the same line do not show consistent differences in either growth, fecundity (including offspring survival), or longevity. Furthermore, none of these traits is genetically correlated across lines with the frequency of the aphallic morph. We conclude that the cost of the construction and maintenance of the phallus must be very low in this species. Future studies should investigate the cost associated with using the phallus (i.e. male outcrossing behaviour) to explain the maintenance of high frequencies of aphallic individuals in natural populations.

Assortative mating (prezygotic isolation) and reduced hybrid fitness (postzygotic isolation) are typically invoked to explain the stability of hybrid zones. In the tension zone model, these factors work in opposition to migration, which promotes genetic homogeneity. Many marine animals migrate over long distances through a planktonic larval stage. Therefore, strong reproductive isolation is needed to maintain stable marine hybrid zones. However, surprisingly little is known about mating preferences and hybrid fitness in marine organisms. Smooth-shelled mussels (Mytilus spp.) form a well-known species complex, with hybridization over extensive areas such as the contact zone of M. edulis and M. galloprovincialis around European Atlantic coasts. This paper reports direct experimental evidence of assortative fertilization, hybrid larval inviability, and early heterosis for growth rate in M. edulis and M. galloprovincialis. Four crosses between pure M. edulis and M. galloprovincialis were analyzed with a new polymerase-chain-reaction-based diagnostic marker. Gamete competition between taxa was allowed in two out of the four crosses. Genotype frequencies observed at an early stage (36 h after fertilization) unambiguously revealed assortative fertilization when gamete competition was allowed. A significant reduction in hybrid viability was subsequently observed during the larval stage. At the same stage an antagonistic effect, heterosis, was observed on growth rate. However, even if heterosis is observed in the F1, it is expected to vanish in subsequent hybrid generations. Although specialization for different habitats and asynchronous spawning have been mentioned as factors contributing to the maintenance of the blue mussel hybrid zone in Europe, we argue that assortative fertilization and reduced hybrid fitness are important factors that also contribute to the stabilization of this zone. These results emphasize that multiple factors may act concomitantly in a barrier to gene flow, especially in complex life cycles. Furthermore, they show that assortative mating through gamete preference, as already demonstrated for sea urchins, may play a role in speciation processes taking place in the sea.

International audience; Whether bioinvasions are associated with a loss of genetic diversity and a change in mating system is instrumental for understanding the evolutionary fate of invasive species. Little loss is expected under strong propagule pressure which might be a general situation in widespread, efficient invader. In hermaphroditic species, we have few examples of a transition between outcrossing and selfing as a consequence of invasion, though this is classically predicted (as a corollary to Baker's law). We estimated microsatellite variation in 44 populations of the widespread freshwater snail Physa acuta sampled at worldwide scale (including several populations from its native North America). Neither loss of variation (or bottleneck), nor increase in selfing rate was detected in invaded areas. Moreover there was no isolation by distance at large geographic scale, and limited geographic coherence in genetic patterns was detected using STRUCTURE software-the West Mediterranean area being an exception. Such patterns might be explained by (1) multiple introductions in the invaded areas, presumably fostered by aquarium trade, followed by regional spread in some cases-in which case mating partners might be numerous enough to prevent transition towards higher selfing rates, and (2) invasions from the whole native area. This suggests that P. acuta is an exceptionally efficient invader (which is not true of related species), but the reasons of its success remain elusive.

International audience; Inbreeding is expected to impair male and female reproductive performance, but little is known on how inbreeding depression varies between sexes and different levels of competition. We studied inbreeding depression in mating behavior and its reproductive consequences in a hermaphroditic freshwater snail and demonstrate that inbreeding depresses mating success in both sex functions. However, the magnitude of inbreeding depression does not differ between sex functions and is not affected by the opportunity for mate choice and male-male competition.Theoretical work predicts that the magnitude of inbreeding depression is particularly high in traits that are closely related to fitness. Despite the extensive work on inbreeding depression of male and female reproductive performance, relatively little is known on how inbreeding impairs male and female mating behavior. We studied inbreeding depression of male and female mating behavior in the simultaneously hermaphroditic freshwater snail Physa acuta to test 1) whether there is inbreeding depression of mating behavior, 2) whether the potential of mate competition and mate choice has an effect on the strength of inbreeding depression, 3) whether the magnitude of inbreeding depression differs between both sex functions, and 4) how inbreeding depression of mating behavior translates into inbreeding depression of reproductive success. For this, we compared the mating behavior between selfed (inbred) and outcrossed (outbred) focal snails in a series of mating trials, in which we manipulated experimentally the potential of mate competition and mate choice. Our results provide evidence for moderate inbreeding depression of the number of copulatory encounters, the number of copulations, and the total time spent mating in both sex functions. The magnitude of inbreeding depression did not differ between the levels of competition and between both sex functions. Finally, our results suggest that inbreeding depression of mating behavior only explains a small fraction of the observed inbreeding depression of reproductive success. We discuss the implications of these findings with respect to precopulatory sexual selection and sex-specific inbreeding depression.

The relationship between genotype and phenotype is not one to one. This statement is central to our understanding of how natural selection shapes phenotypic evolution. Here, we clarify the links between canalization, plasticity and developmental stability, the three major processes involved in the control of phenotypic variability. We present a short historical review, including the original definitions of these concepts, and then summarize their current meaning and use, highlighting possible sources of confusion. Some of the perspectives allowed by a more synthetic conceptual framework are presented, in the light of the recent advances in molecular and developmental genetics.

We characterized eight microsatellite loci to study spatial and temporal population structure of Pelodytes punctatus, a European anuran that has a peculiar breeding pattern among amphibians. The eight loci proved to be highly polymorphic with the number of alleles per locus ranging from two to 15 within two populations. Cross-amplification indicates that those markers may be also useful for closely related species from the same family.