Habitat-quality effects on metapopulation dynamics in greater white-toothed shrews, Crocidura russula.

The effects of patch size and isolation on metapopulation dynamics have received wide empirical support and theoretical formalization. By contrast, the effects of patch quality seem largely underinvestigated, partly due to technical difficulties in properly assessing quality. Here we combine habitat-quality modeling with four years of demographic monitoring in a metapopulation of greater white-toothed shrews (Crocidura russula) to investigate the role of patch quality on metapopulation processes. Together, local patch quality and connectivity significantly enhanced local population sizes and occupancy rates (R2 = 14% and 19%, respectively). Accounting for the quality of patches connected to the focal one and acting as potential sources improved slightly the model explanatory power for local population sizes, pointing to significant source-sink dynamics. Local habitat quality, in interaction with connectivity, also increased colonization rate (R2 = 28%), suggesting the ability of immigrants to target high-quality patches. Overall, patterns were best explained when assuming a mean dispersal distance of 800 m, a realistic value for the species under study. Our results thus provide evidence that patch quality, in interaction with connectivity, may affect major demographic processes.

Conservation of a threatened European tree frog (Hyla arborea) metapopulation

Summary: Amphibians are among the most vulnerable animals of the world. One third of all species are currently threatened with extinction. Habitat loss is the major menace to pond- and stream-breeding species in the old world. In highly urbanized landscape like the Swiss Plateau, most species suffer from habitat reduction and fragmentation. Among all indigenous species, the European tree frog (Hyla arborea L., 1758) is one of the most endangered. It experienced an alarming decline during the last century and its regional long-term persistence is not guaranteed. We developed a monitoring framework based on calling male counts which included multiple visits to each wetland during the reproduction period in order to precisely determine its distribution on the Lemanic coast. Our results indicate that visiting populations 3 limes under suitable climatic conditions (temperature >20°C) provides reliable presence/absence data. Based on our monitoring data, we analyzed the species requirements regarding its breeding habitat. It appeared that anthropogenic activities had paradoxical effects on the species. On one hand, urbanization, traffic and intensive agriculture had a strong detrimental effect on tree frog distribution. On the other hand, large tree frog populations were frequently associated with gravel pits and military training grounds. Our results allowed us to create a habitat suitability map taking into account detrimental landscape elements around ponds (>1100m away from urban areas and >500m away from first class roads). In parallel, we developed a metapopulation model of the European tree frog in order to identify the critical threats to the long term persistence of the species. Our results indicated that suitable pond density is at the low end of the species requirements. Pond creation must therefore be considered an essential complementary approach to pond conservation and restoration. Our model also provided a mapping solution permitting the location of the must suitable area for pond creation from a metapopulation perspective. As many other amphibians, the European tree frog is not only exposed to an aquatic habitat (breeding and larval period), but also to a terrestrial stage (summer and overwintering habitats). Unfortunately, animals in their terrestrial phase are less conspicuous and, as a consequence, their terrestrial needs are relatively unknown. Using a recent tracking method (the Harmonic Direction Finder), we followed post-breeding frogs and identified favored terrestrial habitats, thus providing another practical conservation tool. We conclude that only the combination of multiple spatially explicit approaches (landscape-scale habitat suitability, metapopulation dynamics and terrestrial needs) is likely to provide wildlife managers with effective tools for the conservation of highly endangered amphibians. Résumé: Les amphibiens font partie des animaux les plus vulnérables du monde. Un tiers des espèces est actuellement menacé d'extinction. Dans l'ancien monde, la disparition des habitats constitue la principale menace pour les grenouilles, crapauds, tritons et salamandres. Dans les paysages fortement urbanisés comme le Plateau Suisse, la plupart des espèces souffrent d'une réduction et d'une fragmentation de leurs habitats. Parmi toutes les espèces indigènes, la rainette verte (Hyla arborea L., 1758) est l'une des plus menacée. Sa distribution a régressé de manière alarmante durant le siècle passé et sa survie régionale à long terme n'est pas assurée. Nous avons développé une méthode de suivi des populations se basant sur le comptage des mâles chanteurs durant la période de reproduction. Cette méthode requiert plusieurs visites à chaque plan d'eau de manière à déterminer précisément la distribution de l'espèce. Nos résultats démontrent que 3 visites par population dans des conditions climatiques favorable (température >20°C) permettent d'obtenir des données de présence/ absence valables. Sur la base de nos comptages sur la Côte lémanique, nous avons analysé les exigences de l'espèce concernant ses sites de reproduction. Il est apparu que les activités humaines avaient un effet paradoxal sur l'espèce. D'une part, l'urbanisation, le trafic routier et l'intensification de l'agriculture ont un effet fortement préjudiciable, tandis que d'autre part les plus grandes populations sont souvent associées à des gravières et autres places d'armes. Nos résultats ont permis de créer une carte de qualité d'habitat prenant en compte les éléments paysagers préjudiciables à la rainette (situé à plus de 1100m de zones urbaines et à plus de 500m de routes de première classe). En parallèle, nous avons développé un modèle métapopulationnel (incluant l'ensemble des populations) de manière à identifier les menaces prépondérantes sur la survie à long terme de l'espèce. Nos résultats ont permis de déterminer que la densité actuelle de plans d'eau adéquats est à la limite inférieure des exigences de l'espèce. La création d'étangs doit donc être considérée comme une approche indispensable et complémentaire à la protection et à la restauration des sites existants. Notre modèle a également fourni des résultats cartographiables permettant l'identification des sites les plus appropriés dans une perspective métapopulationnelle. Comme de nombreux autres amphibiens, la rainette verte est exposée à un habitat aquatique (reproduction et développement larvaire) ainsi qu'à un habitat terrestre (été et hiver). Les animaux étant particulièrement cryptiques dans cette seconde phase, leurs besoins terrestres sont relativement mal connus. Nous avons donc développé une nouvelle méthode de télémétrie basée sur le goniomètre harmonique. Cette méthode nous a permis de suivre des rainettes dans leurs migrations jusqu'à leurs habitats d'été et d'établir ainsi des recommandations pratiques pour la conservation de la rainette. Nous concluons que la combinaison de multiples approches spatialement explicites (qualité d'habitat, dynamique de métapopulation et habitats terrestres) est seule à même de produire des outils efficaces pour la conservation des espèces menacées d'amphibiens.

A metapopulation model for the introgression from genetically modified plants into their wild relatives

Most models on introgression from genetically modified (GM) plants have focused on small spatial scales, modelling gene flow from a field containing GM plants into a single adjacent population of a wild relative. Here, we present a model to study the effect of introgression from multiple plantations into the whole metapopulation of the wild relative. The most important result of the model is that even very low levels of introgression and selection can lead to a high probability that the transgene goes to fixation in the metapopulation. Furthermore, the overall frequency of the transgene in the metapopulation, after a certain number of generations of introgression, depends on the population dynamics. If there is a high rate of migration or a high rate of population turnover, the overall transgene frequency is much higher than with lower rates. However, under an island model of population structure, this increased frequency has only a very small effect on the probability of fixation of the transgene. Considering these results, studies on the potential ecological risks of introgression from GM plants should look not only at the rate of introgression and selection acting on the transgene, but also at the metapopulation dynamics of the wild relative.

The fixation of locally beneficial alleles in a metapopulation.

Extinction, recolonization, and local adaptation are common in natural spatially structured populations. Understanding their effect upon genetic variation is important for systems such as genetically modified organism management or avoidance of drug resistance. Theoretical studies on the effect of extinction and recolonization upon genetic variance started appearing in the 1970s, but the role of local adaptation still has no good theoretical basis. Here we develop a model of a haploid species in a metapopulation in which a locally adapted beneficial allele is introduced. We study the effect of different spatial patterns of local adaptation, and different metapopulation dynamics, upon the fixation probability of the beneficial allele. Controlling for the average selection pressure, we find that a small area of positive selection can significantly increase the global probability of fixation. However, local adaptation becomes less important as extinction rate increases. Deme extinction and recolonization have a spatial smoothing effect that effectively reduces spatial variation in fitness.

Effects of cognitive abilities on metapopulation connectivity

Connectivity among demes in a metapopulation depends on both the landscape's and the focal organism's properties (including its mobility and cognitive abilities). Using individual-based simulations, we contrast the consequences of three different cognitive strategies on several measures of metapopulation connectivity. Model animals search suitable habitat patches while dispersing through a model landscape made of cells varying in size, shape, attractiveness and friction. In the blind strategy, the next cell is chosen randomly among the adjacent ones. In the near-sighted strategy, the choice depends on the relative attractiveness of these adjacent cells. In the far-sighted strategy, animals may additionally target suitable patches that appear within their perceptual range. Simulations show that the blind strategy provides the best overall connectivity, and results in balanced dispersal. The near-sighted strategy traps animals into corridors that reduce the number of potential targets, thereby fragmenting metapopulations in several local clusters of demes, and inducing sink-source dynamics. This sort of local trapping is somewhat prevented in the far-sighted strategy. The colonization success of strategies depends highly on initial energy reserves: blind does best when energy is high, near-sighted wins at intermediate levels, and far-sighted outcompetes its rivals at low energy reserves. We also expect strong effects in terms of metapopulation genetics: the blind strategy generates a migrant-pool mode of dispersal that should erase local structures. By contrast, near- and far-sighted strategies generate a propagule-pool mode of dispersal and source-sink behavior that should boost structures (high genetic variance among- and low variance within local clusters of demes), particularly if metapopulation dynamics is also affected by extinction-colonization processes. Our results thus point to important effects of the cognitive ability of dispersers on the connectivity, dynamics and genetics of metapopulations.

Evolution in heterogeneous populations: from migration models to fixation probabilities.

Although dispersal is recognized as a key issue in several fields of population biology (such as behavioral ecology, population genetics, metapopulation dynamics or evolutionary modeling), these disciplines focus on different aspects of the concept and often make different implicit assumptions regarding migration models. Using simulations, we investigate how such assumptions translate into effective gene flow and fixation probability of selected alleles. Assumptions regarding migration type (e.g. source-sink, resident pre-emption, or balanced dispersal) and patterns (e.g. stepping-stone versus island dispersal) have large impacts when demes differ in sizes or selective pressures. The effects of fragmentation, as well as the spatial localization of newly arising mutations, also strongly depend on migration type and patterns. Migration rate also matters: depending on the migration type, fixation probabilities at an intermediate migration rate may lie outside the range defined by the low- and high-migration limits when demes differ in sizes. Given the extreme sensitivity of fixation probability to characteristics of dispersal, we underline the importance of making explicit (and documenting empirically) the crucial ecological/ behavioral assumptions underlying migration models.

The scope of Baker's law.

656 I. 657 II. 658 III. 660 IV. 661 V. 663 VI. 663 VII. 664 VIII. 664 665 References 665 SUMMARY: Baker's law refers to the tendency for species that establish on islands by long-distance dispersal to show an increased capacity for self-fertilization because of the advantage of self-compatibility when colonizing new habitat. Despite its intuitive appeal and broad empirical support, it has received substantial criticism over the years since it was proclaimed in the 1950s, not least because it seemed to be contradicted by the high frequency of dioecy on islands. Recent theoretical work has again questioned the generality and scope of Baker's law. Here, we attempt to discern where the idea is useful to apply and where it is not. We conclude that several of the perceived problems with Baker's law fall away when a narrower perspective is adopted on how it should be circumscribed. We emphasize that Baker's law should be read in terms of an enrichment of a capacity for uniparental reproduction in colonizing situations, rather than of high selfing rates. We suggest that Baker's law might be tested in four different contexts, which set the breadth of its scope: the colonization of oceanic islands, metapopulation dynamics with recurrent colonization, range expansions with recurrent colonization, and colonization through species invasions.

Mother-offspring competition promotes colonization success.

Colonization is the crucial process underlying range expansions, biological invasions, and metapopulation dynamics. Which individuals leave their natal population to colonize empty habitats is a crucial question and is presently unresolved. Dispersal is the first step in colonization. However, not all dispersing individuals are necessarily good colonizers. Indeed, in some species, the phenotype of dispersers differs depending on the selective pressures that induce dispersal. In particular, kin-based interactions, a factor driving social evolution, should induce different social response profiles in nondispersing and dispersing individuals. Kin competition (defined here as between the mother and offspring) has been proven to produce dispersers with a particular phenotype that may enhance their colonizing ability. By using the common lizard (Lacerta vivipara), we conducted a multipopulation experiment to study the effect of kin competition on dispersal and colonization success. We manipulated mother-offspring interactions, which are the most important component of kin competition in the studied species, at the family and population levels and measured the consequences on colonization success. We demonstrate that mother-offspring competition at the population level significantly influences colonization success. Increased competition at the population level enhanced the colonization rate of the largest juveniles as well as the growth and survival of the colonizers. Based on these results, we calculated that kin-induced colonization halves the extinction probability of a newly initiated population. Because interactions between relatives are likely to affect the ability of a species to track habitat modifications, kin-based dispersal should be considered in the study of invasion dynamics and metapopulation functioning.

Keeping the balance? Management of oxidative stress, body mass and reproduction under energetic constraints by dispersing and philopatric collared flycatchers.

Dispersal, i.e. individual movement between breeding sites, is a key process for metapopulation dynamics and gene flow. Its success can be modulated by phenotypic differences between dispersing and philopatric individuals, or dispersal syndromes. However, the environmental (external) and physiological (internal) constraints underlying such syndromes remain poorly known. This project aimed at clarifying the impact of environmental variation and oxidative constraints, linked to the reactive oxygen species produced during respiration, on phenotypes associated to dispersal in a passerine bird, the collared flycatcher Ficedula albicollis. Energetic demand was experimentally (i) increased through a wing load manipulation or (ii) relieved through food supplementation. The oxidative balance of breeding flycatchers was influenced by complex interactions of dispersal status and extrinsic factors (breeding density, year, experimental treatments). Interestingly, antioxidant capacity was influenced both by permanent individual differences and by food availability, whereas measures of pro-oxidants were highly variables within individuals. Environmental variation and energetic constraints also modulated the differences in reproduction associated with dispersal: dispersing and philopatric birds differ in their management of the oxidative balance when it is competing with reproductive investment. This thesis highlights that reaction norms, rather than fixed differences, often shape traits associated to dispersal. ----- Le déplacement d'un individu entre sites de reproduction, ou dispersion, est un processus clé pour la dynamique des métapopulations et les flux de gènes. Son succès peut être modulé par des différences de phénotype, ou syndromes de dispersion. Cependant, les contraintes environnementales et physiologiques qui sous-tendent ces syndromes restent mal connues. Ce projet vise à clarifier l'impact des variations environnementales et des contraintes oxydatives (liées aux espèces réactives de l'oxygène produites durant la respiration) sur les phénotypes associés à la dispersion chez un passereau, le gobemouche à collier Ficedula albicollis. La demande énergétique a été expérimentalement (i) augmentée en manipulant la surface alaire ou (ii) diminuée par une supplémentation en nourriture. L'équilibre oxydo-réducteur des gobemouches en reproduction est influencé par des interactions complexes entre statut de dispersion et facteurs extrinsèques (densité de couples reproducteurs, année, traitement expérimental). La capacité antioxydante dépend principalement de différences permanentes entre individus, alors que les pro-oxydants présentent de grandes variations intra-individu. Environnement et contraintes énergétiques modulent aussi les différences de reproduction liées à la dispersion : les oiseaux dispersants et philopatriques diffèrent dans leur gestion de l'équilibre oxydo-réducteur lorsqu'il est en compétition avec l'investissement reproducteur. Ce travail souligne que les traits associés à la dispersion sont souvent déterminés par des normes de réaction à l'environnement et non des différences fixées entre individus.

On metapopulation resistance to drift and extinction.

The spatial configuration of metapopulations (numbers, sizes, and localization of patches) affects their ability to resist demographic extinction and genetic drift, but sometimes with opposite effects. Small and isolated patches, for instance, contribute marginally to demography but may play a large role in genetics by maintaining a sizeable amount of genetic variance among demes. In source-sink systems, similarly, connectivity may be beneficial in terms of effective size, but detrimental in terms of survival, by lowering the reproductive value of source populations. How to reconcile these opposite effects? Here we propose an analytical framework that integrates fixation time (ability to resist genetic drift) and extinction time (ability to resist demographic extinction) into a single index of resistance, measuring the ability of a metapopulation to maintain its demo-genetic integrity. We then illustrate with numerical examples how conflicting demands may be resolved.

Viability and Management of an Endangered Capercaillie (Tetrao urogallus) Metapopulation in the Jura Mountains, Western Switzerland

The populations of Capercaillie (Tetrao urogallus), the largest European grouse, have seriously declined during the last century over most of their distribution in western and central Europe. In the Jura mountains, the relict population is now isolated and critically endangered (about 500 breeding adults). We developed a simulation software (TetrasPool) that accounts for age and spatial structure as well as stochastic processes, to perform a viability analysis and explore management scenarios for this population, capitalizing on a 24 years-long series of field data. Simulations predict a marked decline and a significant extinction risk over the next century, largely due to environmental and demographic stochasticity (average values of life-history parameters would otherwise allow stability). Variances among scenarios mainly stem from uncertainties about the shape and intensity of density dependence. Uncertainty analyses suggest to focus conservation efforts on enhancing, not only adult survival (as often advocated for long-lived species), but also recruitment. The juvenile stage matters when local populations undergo extinctions, because it ensures connectivity and recolonization. Besides limiting human perturbations, a silvicultural strategy aimed at opening forest structure should improve the quality and surface of available patches, independent of their size and localization. Such measures are to be taken urgently, if the population is to be saved.

How patch configuration affects the impact of disturbances on metapopulation persistence.

Disturbances affect metapopulations directly through reductions in population size and indirectly through habitat modification. We consider how metapopulation persistence is affected by different disturbance regimes and the way in which disturbances spread, when metapopulations are compact or elongated, using a stochastic spatially explicit model which includes metapopulation and habitat dynamics. We discover that the risk of population extinction is larger for spatially aggregated disturbances than for spatially random disturbances. By changing the spatial configuration of the patches in the system--leading to different proportions of edge and interior patches--we demonstrate that the probability of metapopulation extinction is smaller when the metapopulation is more compact. Both of these results become more pronounced when colonization connectivity decreases. Our results have important management implication as edge patches, which are invariably considered to be less important, may play an important role as disturbance refugia.

Effects of colonization asymmetries on metapopulation persistence.

Ocean currents, prevailing winds, and the hierarchical structures of river networks are known to create asymmetries in re-colonization between habitat patches. The impacts of such asymmetries on metapopulation persistence are seldom considered, especially rarely in theoretical studies. Considering three classical models (the island, the stepping stone and the distance-dependent model), we explore how metapopulation persistence is affected by (i) asymmetry in dispersal strength, in which the colonization rate between two patches differs in direction, and (ii) asymmetry in connectivity, in which the overall colonization pattern displays asymmetry (circulating or dendritic networks). Viability can be drastically reduced when directional bias in dispersal strength is higher than 25%. Re-colonization patterns that allow for strong local connectivity provide the highest persistence compared to systems that allow circulation. Finally, asymmetry has relatively weak effects when metapopulations maintain strong general connectivity.

Mdx myotubes have normal excitability but show reduced contraction-relaxation dynamics.

The pathogenesis of Duchenne muscular dystrophy (DMD), characterised by lack of the cytoskeletal protein dystrophin, is not completely understood. An early event in the degenerative process of DMD muscle could be a rise in cytosolic calcium concentration. In order to investigate whether this leads to alterations of contractile behaviour, we studied the excitability and contractile properties of cultured myotubes from control (C57BL/10) and mdx mice, an animal model for DMD. The myotubes were stimulated electrically and their motion was recorded photometrically. No significant differences were found between control and mdx myotubes with respect to the following parameters: chronaxy and rheobase (0.33 +/- 0.03 ms and 23 +/- 4 V vs. 0.39 +/- 0.07 ms and 22 +/- 2 V for C57 and mdx myotubes, respectively), tetanisation frequency (a similar distribution pattern was found between 5 and 30 Hz), fatigue during tetanus (found in 35% of both types of myotubes) and post-tetanic contracture. In contrast, contraction and relaxation times were longer (P < 0.005) in mdx (36 +/- 2 and 142 +/- 13 ms, respectively) than in control myotubes (26 +/- 1 and 85 +/- 9 ms, respectively). Together with our earlier findings, these results suggest a decreased capacity for calcium removal in mdx cells leading, in particular, to alterations of muscle relaxation.

The dynamics of interfaces: Seven authors in search of encounters across levels of description of an event involving a mother, father, and baby

The interfaces between the intrapsychic, interactional, and intergenerational domains are a new frontier. As a pilot, we exposed ourselves to a complex but controllable situation as viewed by people whose main interest is in one of the three interfaces; we also fully integrated the subjects in the team, to learn about their subjective perspectives and to provide them with an enriching experience. We started with a brief "triadification" sequence (i.e., moving from a "two plus one" to a "three together" family organization). Considering this sequence as representing at a micro level many larger family transitions, we proceeded with a microanalytic interview, a psychodynamic investigation, and a family interview. As expected, larger patterns of correspondences are emerging. Central questions under debate are: What are the most appropriate units at each level of description and what are their articulations between these levels? What is the status of "triadification"? Les interfaces entre les domaines intrapsychiques, interactionnels et intergénérationnels représentent une nouvelle frontiére. A titre exploratoire, nous nous sommes exposés à une situation complexe mais contrǒlable ainsi que le voient ceux dont I'intérět principal se porte sur l'une de ces trois interfaces. Nous avons aussi entièrement intégré les sujets dans l'équipe, de facon à comprendre leur perspective subjective et à leur offrir une expérience enrichissante. Nous avons commencé avec une brève séquence de "triadification," c'est-à-dire passer d'une organisation familiale "deux plus un" à Ltne organisation familiale "trois (add sentenc)ensemble." Considérant cette séquence comme representative à un niveau microscopique de transitions familiales bien plus larges, nous avons procedé à l'entretien microanalytique, à une enquěte psychodynamique et à un entretien familial. Comme prévu, de grands patterns de correspondances émergent. Les questions essentielles sur lesquelles portent le débat sont: quelles les unités les plus appropiées à chaque niveau de description et quelles sont les articulations entre ces niveaux? Quel est le statut de la "triadification"?