Genetic Variability, Population Differentiation, and Correlations for Thermal Tolerance Indices in the Minute Wasp, Trichogramma cacoeciae.

Temperature is a main driver of the ecology and evolution of ectotherms. In particular, the ability to move at sub-lethal low temperatures can be described through three thermal tolerance indices-critical thermal minimum (CTmin), chill coma temperature (CCT), and activity recovery (AR). Although these indices have proven relevant for inter-specific comparisons, little is known about their intraspecific variability as well as possible genetic correlations between them. We thus investigated these two topics (intraspecific variability and genetic correlations between thermal tolerance indices) using the minute wasp, Trichogramma cacoeciae. Strains from T. cacoeciae were sampled across three geographic regions in France-two bioclimatic zones along a sharp altitudinal cline in a Mediterranean context (meso-Mediterranean at low elevations and supra-Mediterranean at higher elevations) and a more northwestern area characterized by continental or mountainous climates. Our results evidenced a significant effect of both the longitude and the severity of the cold during winter months on CCT. Results were however counter-intuitive since the strains from the two bioclimatic zones characterized by more severe winters (northwestern area and supra-Mediterranean) exhibited opposite patterns. In addition, a strong positive correlation was observed between CCT and CTmin. Neither strain differentiation nor the covariations between traits seem to be linked with the molecular diversity observed on the part of the mitochondrial marker COI.

Its all about connections: hubs and invasion in habitat networks.

During the early stages of invasion, the interaction between the features of the invaded landscape, notably its spatial structure, and the internal dynamics of an introduced population has a crucial impact on establishment and spread. By approximating introduction areas as networks of patches linked by dispersal, we characterised their spatial structure with specific metrics and tested their impact on two essential steps of the invasion process: establishment and spread. By combining simulations with experimental introductions of Trichogramma chilonis (Hymenoptera: Trichogrammatidae) in artificial laboratory microcosms, we demonstrated that spread was hindered by clusters and accelerated by hubs but was also affected by small-population mechanisms prevalent for invasions, such as Allee effects. Establishment was also affected by demographic mechanisms, in interaction with network metrics. These results highlight the importance of considering the demography of invaders as well as the structure of the invaded area to predict the outcome of invasions.

Evidence for an optimal level of connectivity for establishment and colonization.

Dispersal is usually associated with the spread of invasive species, but it also has two opposing effects, one decreasing and the other increasing the probability of establishment. Indeed, dispersal both slows population growth at the site of introduction and increases the likelihood of surrounding habitat being colonized. The connectivity of the introduction site is likely to affect dispersal, and, thus, establishment, according to the dispersal behaviour of individuals. Using individual-based models and microcosm experiments on minute wasps, we demonstrated the existence of a hump-shaped relationship between connectivity and establishment in situations in which individual dispersal resembled a diffusion process. These results suggest that there is an optimal level of connectivity for the establishment of introduced populations locally at the site of introduction, and regionally over the whole landscape.

Temporal autocorrelation in host density increases establishment success of parasitoids in an experimental system.

Environmental variation is classically expected to affect negatively population growth and to increase extinction risk, and it has been identified as a major determinant of establishment failures in the field. Yet, recent theoretical investigations have shown that the structure of environmental variation and more precisely the presence of positive temporal autocorrelation might alter this prediction. This is particularly likely to affect the establishment dynamics of biological control agents in the field, as host-parasitoid interactions are expected to induce temporal autocorrelation in host abundance. In the case where parasitoid populations display overcompensatory dynamics, the presence of such positive temporal autocorrelation should increase their establishment success in a variable environment. We tested this prediction in laboratory microcosms by introducing parasitoids to hosts whose abundances were manipulated to simulate uncorrelated or positively autocorrelated variations in carrying capacity. We found that environmental variability decreased population size and increased parasitoid population variance, which is classically expected to extinction risk. However, although exposed to significant environmental variation, we found that parasitoid populations experiencing positive temporal autocorrelation in host abundance were more likely to persist than populations exposed to uncorrelated variation. These results confirm that environmental variation is a key determinant of extinction dynamics that can have counterintuitive effects depending on its autocorrelation structure.

Estimation of the dispersal of a major pest of maize by cline analysis of a temporary contact zone between two invasive outbreaks.

Dispersal is a key factor in invasion and in the persistence and evolution of species. Despite the importance of estimates of dispersal distance, dispersal measurement remains a real methodological challenge. In this study, we characterized dispersal by exploiting a specific case of biological invasion, in which multiple introductions in disconnected areas lead to secondary contact between two differentiated expanding outbreaks. By applying cline theory to this ecological setting, we estimated σ, the standard deviation of the parent-offspring distance distribution, of the western corn rootworm, Diabrotica virgifera virgifera, one of the most destructive pests of maize. This species is currently invading Europe, and the two largest invasive outbreaks, in northern Italy and Central Europe, have recently formed a secondary contact zone in northern Italy. We identified vanishing clines at 12 microsatellite loci throughout the contact zone. By analysing both the rate of change of cline slope and the spatial variation of linkage disequilibrium at these markers, we obtained two σ estimates of about 20 km/generation(1/2). Simulations indicated that these estimates were robust to changes in dispersal kernels and differences in population density between the two outbreaks, despite a systematic weak bias. These estimates are consistent with the results of direct methods for measuring dispersal applied to the same species. We conclude that secondary contact resulting from multiple introductions is very useful for the inference of dispersal parameters and should be more widely used in other species.

Time-lag in extinction dynamics in experimental populations: evidence for a genetic Allee effect?

1. Propagule pressure, i.e. the number of individuals introduced, is thought to be a major predictor of the establishment success of introduced populations in the field. Its influence in laboratory experimental systems has however been questioned. In fact, other factors involved in long-term population persistence, like habitat size, were usually found to explain most of the dynamics of experimental populations. 2. To better understand the respective influence of short- and long-term factors and their potential interaction on extinction dynamics in experimental systems, we investigated the influence of propagule pressure, habitat size and genetic background on the early dynamics of laboratory-based populations of a hymenopteran parasitoid. 3. The amount of demographic variance differed between establishment and persistence phase and was influenced by habitat size and genetic background (geographic strain), but independent of propagule pressure. In contrast, the probability of extinction within five generations depended on the genetic background and on the interaction between propagule pressure and habitat size. Vulnerability to extinction in small size habitats was increased when populations were founded with a small number of individuals, but this effect was delayed until the third to fifth generations. 4. These results indicate that demographic stochasticity is influential during population establishment, but is not affected by the genetic variability of propagules. On the other hand, extinction might be influenced by a genetic Allee effect triggered by the combination of low propagule pressure and genetic drift. Finally, we documented consistent differences between genetic backgrounds in both deterministic and stochastic population dynamics patterns, with major consequences on extinction risk and ultimately population establishment.

The biology of small, introduced populations, with special reference to biological control.

Populations are introduced into novel environments in different contexts, one being the biological control of pests. Despite intense efforts, less than half introduced biological control agents establish. Among the possible approaches to improve biological control, one is to better understand the processes that underpin introductions and contribute to ecological and evolutionary success. In this perspective, we first review the demographic and genetic processes at play in small populations, be they stochastic or deterministic. We discuss the theoretical outcomes of these different processes with respect to individual fitness, population growth rate, and establishment probability. Predicted outcomes differ subtly in some cases, but enough so that the evaluating results of introductions have the potential to reveal which processes play important roles in introduced populations. Second, we attempt to link the theory we have discussed with empirical data from biological control introductions. A main result is that there are few available data, but we nonetheless report on an increasing number of well-designed, theory-driven, experimental approaches. Combining demography and genetics from both theoretical and empirical perspectives highlights novel and exciting avenues for research on the biology of small, introduced populations, and great potential for improving both our understanding and practice of biological control.

Intraspecific variability in the parasitoid wasp Trichogramma chilonis: can we predict the outcome of hybridization?

In the framework of biological control, the selection of effective natural enemies determines the final pest control. Thus, the genetic improvement of biocontrol agents could enhance the efficiency of biocontrol programs. Although promising, this approach has rarely been applied in this field. At the intraspecific level, hybridization between divergent populations of biocontrol agents is expected to promote hybrid vigor (heterosis), but it is not clear to what extent. An even more difficult task is the ability to predict the fitness of hybrids from the biological characteristics of their parents. We investigated these general questions by crossing seven populations of the parasitoid wasp Trichogramma chilonis (Hymenoptera: Trichogrammatidae). Our results show different levels of mating compatibilities among populations, including asymmetric or almost complete reproductive isolation. Hybrids' performance (fitness of the F(1) generation) ranges from inbreeding depression to heterosis. It was possible, to some extent, to predict hybrid fitness from pairwise genetic and phenotypic distances among parents, in accordance with the 'dominance' hypothesis. This may provide general guidelines for the genetic improvement of biological control agents.

Don'T fall off the adaptation cliff: when asymmetrical fitness selects for suboptimal traits.

The cliff-edge hypothesis introduces the counterintuitive idea that the trait value associated with the maximum of an asymmetrical fitness function is not necessarily the value that is selected for if the trait shows variability in its phenotypic expression. We develop a model of population dynamics to show that, in such a system, the evolutionary stable strategy depends on both the shape of the fitness function around its maximum and the amount of phenotypic variance. The model provides quantitative predictions of the expected trait value distribution and provides an alternative quantity that should be maximized ("genotype fitness") instead of the classical fitness function ("phenotype fitness"). We test the model's predictions on three examples: (1) litter size in guinea pigs, (2) sexual selection in damselflies, and (3) the geometry of the human lung. In all three cases, the model's predictions give a closer match to empirical data than traditional optimization theory models. Our model can be extended to most ecological situations, and the evolutionary conditions for its application are expected to be common in nature.

Maintenance of fungal pathogen species that are specialized to different hosts: allopatric divergence and introgression through secondary contact.

Sympatry of species that lack complete prezygotic isolation is ideal for the study of how species can be maintained in the face of potential gene flow. This is particularly important in the context of emerging diseases on new hosts because pathogen adaptation is facilitated by reduced gene flow from ancestral populations. Here, we investigated divergence and gene flow between two closely related fungal species, Microbotryum lychnidis-dioicae and M. silenes-dioicae, causing anther-smut disease on the wide-spread plant species Silene latifolia and S. dioica, respectively. Using model-based clustering algorithms on microsatellite data from samples across Europe, we identified rare disease transmission between the host species and rare pathogen hybrids. Using a coalescent-based approach and an isolation-with-migration model, the age of divergence between the two fungal species was estimated at approximately 4.2 × 10(5) years. Levels of gene flow were low and concentrated in very recent times. In addition, gene flow appeared unidirectional from M. silenes-dioicae to M. lychnidis-dioicae. Altogether, our findings are consistent with a scenario of recurrent introgressive hybridization but at a very low level and through secondary contact following initial divergence in allopatry. Asymmetry in the direction of gene flow mirrors previous findings on introgression between the two host plants. Our study highlights the consequences of bringing closely related pathogens into contact, which is increasing through modern global changes and favors cross-species disease transmission, hybridization, and introgression by pathogens.

Glacial refugia in pathogens: European genetic structure of anther smut pathogens on Silene latifolia and Silene dioica.

Climate warming is predicted to increase the frequency of invasions by pathogens and to cause the large-scale redistribution of native host species, with dramatic consequences on the health of domesticated and wild populations of plants and animals. The study of historic range shifts in response to climate change, such as during interglacial cycles, can help in the prediction of the routes and dynamics of infectious diseases during the impending ecosystem changes. Here we studied the population structure in Europe of two Microbotryum species causing anther smut disease on the plants Silene latifolia and Silene dioica. Clustering analyses revealed the existence of genetically distinct groups for the pathogen on S. latifolia, providing a clear-cut example of European phylogeography reflecting recolonization from southern refugia after glaciation. The pathogen genetic structure was congruent with the genetic structure of its host species S. latifolia, suggesting dependence of the migration pathway of the anther smut fungus on its host. The fungus, however, appeared to have persisted in more numerous and smaller refugia than its host and to have experienced fewer events of large-scale dispersal. The anther smut pathogen on S. dioica also showed a strong phylogeographic structure that might be related to more northern glacial refugia. Differences in host ecology probably played a role in these differences in the pathogen population structure. Very high selfing rates were inferred in both fungal species, explaining the low levels of admixture between the genetic clusters. The systems studied here indicate that migration patterns caused by climate change can be expected to include pathogen invasions that follow the redistribution of their host species at continental scales, but also that the recolonization by pathogens is not simply a mirror of their hosts, even for obligate biotrophs, and that the ecology of hosts and pathogen mating systems likely affects recolonization patterns.

Frequency-dependent reproductive success in female common lizards: a real-life hawk-dove-bully game?

Alternative strategies are characterised by context-dependent fitness payoffs, which means that their fitness depends on the frequency and the nature of their interactions with one or more strategies. The analysis of the variation of the fitness of each strategy in different social environments can elucidate the evolutionary dynamics played by the strategies. In the common lizard, three female colour types (yellow, orange and mixed) are associated with alternative reaction norms in reproduction and social behaviour that signal alternative strategies. To clarify the nature of colour-specific interactions and their influence on female fitness, we analysed the response of female reproductive success to an experimental manipulation of colour frequencies in natural populations. We found that juvenile body condition at birth for all colour types was negatively affected by the local frequency of yellow females. In addition, we found that mixed females had higher clutch hatching success in the populations where orange females were frequent. These results prove that female reproduction is sensitive to the social environment, and are consistent with a scenario of a hawk-dove-bully game, in which yellow females are aggressive hawks, orange females non-aggressive doves, and mixed females have a context-dependent bully strategy. In this system, the plastic bully strategy would confer a reproductive advantage to putative heterozygotes in some social environments, which could allow the maintenance of the system through context-dependent overdominance effects.

Are dispersal-dependent behavioral traits produced by phenotypic plasticity?

Dispersal is a common response to deteriorating conditions such as intense competition, food limitation, predation or parasitism. Although it provides obvious advantages, dispersal is often assumed to be costly. Selection is therefore likely to have acted to decrease these costs, and indeed several studies demonstrated that dispersers and philopatric individuals differ in their morphology, physiology and/or behavior. Using the common lizard (Lacerta vivipara) as our model system, we examined the contribution of phenotypic plasticity to the establishment of dispersal-dependent behavioral traits. We used a reciprocal transplant experiment in which conditions at the maternal site of origin, during offspring development in utero, and at the release site were manipulated. We then compared activity, social interactions and foraging behavior between individuals that stayed philopatric and those that dispersed. Most behavioral traits were also measured at birth and after the dispersal phase.This study demonstrates that (a) 10 months after the dispersal phase, there were still marked behavioral differences between dispersing and philopatric individuals, (b) the reaction when confronted to another individual was also dispersal-status dependent, a result which strongly suggests that individuals are able to recognize the dispersal status of same-age conspecifics and (c) none of the behavioral characteristics were found to be dependent on the environmental conditions (maternal and natal environment) indicating a lack of phenotypic plasticity in the building of the dispersal-dependent behavioral traits examined.

Mother's timing and duration of corticosterone exposure modulate offspring size and natal dispersal in the common lizard (Lacerta vivipara).

Although multiple condition dependence in dispersal is common, the proximate mechanisms that integrate information from multiple sources remain largely unknown. In the common lizard (Lacerta vivipara), earlier studies have shown that maternal plasma corticosterone level interacts with maternal phenotype to affect offspring phenotype and dispersal strategy, and may reflect the mother's external and/or internal environment. However, the mechanism by which these two types of environmental information are integrated has not been identified. We explored the possibility that the timing and duration of the corticosterone signal are the key factors by which the message is modulated. We found that the timing of corticosterone exposure affects the juvenile phenotype: an exposure to corticosterone early in development has negative effects on juvenile size, weight, and body condition, that can nevertheless be restored in the case of a prolonged exposure. The duration of corticosterone exposure affects the dispersal strategy, although the precise effect depends on the sex ratio of the clutch. That is, in female-biased clutches, a prolonged exposure during gestation promotes philopatry of juveniles, while a short exposure enhances their dispersal, a result that is consistent with kin competition theory. Therefore, our results demonstrate that while corticosterone titer may signal a mother's external and/or internal environment to her developing young, differences in the timing and duration of this endocrine signal produce offspring with specialized phenotypes that exhibit different dispersal strategies.