Population genetics of freshwater snails

Freshwater snails have attracted the attention of biologists for a long time, because they are intermediate hosts of schistosomes, agents of bilharziases. However, population-genetic studies of freshwater snails have been undertaken only during the past decade, covering topics such as the relative roles of genetic drift and gene flow in subdivided populations and the roles of extinction and recolonization events in determining population structure. Other studies in freshwater snails have investigated the maintenance of sex and the evolution of selling, widening a debate restricted mainly to plant populations. The possible role of parasites in freshwater-snail population genetics has also been investigated.     

Coevolution of species colonisation rates controls food-chain length in spatially structured food webs

How the complexity of food webs depends on environmental variables is a long-standing ecological question. It is unclear though how food-chain length should vary with adaptive evolution of the constitutive species. Here we model the evolution of species colonisation rates and its consequences on occupancies and food-chain length in metacommunities. When colonisation rates can evolve, longer food-chains can persist. Extinction, perturbation and habitat loss all affect evolutionarily stable colonisation rates, but the strength of the competition-colonisation trade-off has a major role: weaker trade-offs yield longer chains. Although such eco-evo dynamics partly alleviates the spatial constraint on food-chain length, it is no magic bullet: the highest, most vulnerable, trophic levels are also those that least benefit from evolution. We provide qualitative predictions regarding how trait evolution affects the response of communities to disturbance and habitat loss. This highlights the importance of eco-evolutionary dynamics at metacommunity level in determining food-chain length.     

Eco-evolution from deep time to contemporary dynamics: The role of timescales and rate modulators

Eco-evolutionary dynamics, or eco-evolution for short, are often thought to involve rapid demography (ecology) and equally rapid heritable phenotypic changes (evolution) leading to novel, emergent system behaviours. We argue that this focus on contemporary dynamics is too narrow: Eco-evolution should be extended, first, beyond pure demography to include all environmental dimensions and, second, to include slow eco-evolution which unfolds over thousands or millions of years. This extension allows us to conceptualise biological systems as occupying a two-dimensional time space along axes that capture the speed of ecology and evolution. Using Hutchinson's analogy: Time is the ‘theatre’ in which ecology and evolution are two interacting ‘players’. Eco-evolutionary systems are therefore dynamic: We identify modulators of ecological and evolutionary rates, like temperature or sensitivity to mutation, which can change the speed of ecology and evolution, and hence impact eco-evolution. Environmental change may synchronise the speed of ecology and evolution via these rate modulators, increasing the occurrence of eco-evolution and emergent system behaviours. This represents substantial challenges for prediction, especially in the context of global change. Our perspective attempts to integrate ecology and evolution across disciplines, from gene-regulatory networks to geomorphology and across timescales, from today to deep time.     

Population biology of the invasive freshwater snail Physa acuta approached through genetic markers, ecological characterization and demography

Abstract The respective role of factors acting on population functioning can be inferred from a variety of approaches, including population genetics and demography. We here investigated the role of four of these factors (mating systems, population size, bottlenecks and migration) in the hermaphroditic freshwater snail Physa acuta. Twenty-four populations were sampled either around Montpellier (local scale), or at the scale of France (global scale). At local scale, eight populations were sampled twice, before and after summer drying out. The genetic structure of these populations was studied using microsatellite loci. Populations were classified according to openness (ponds vs. rivers) and water regime (permanent vs. temporary) allowing predictions on genetic patterns (e.g. diversity within populations and differentiation). At local scale, progeny-arrays analysis of the selfing rate was conducted, and size distributions of individuals were followed over two years. Results with regard to the four factors mentioned above were: (i) Estimates of population selfing rates derived from inbreeding coefficients were only slightly higher than those from progeny-arrays. (ii) More variation was detected in rivers than in ponds, but no influence of water regime was detected. One reason might be that permanent populations are not going less often through low densities than those from temporary habitats at the time scale studied. (iii) There was limited evidence for genetic bottlenecks which is compatible with the fact that even marked reduction in water availability was not necessarily associated with demographic bottlenecks. More generally, bottlenecks reducing genetic variation probably occur at population foundation. (iv) Lower genetic differentiation was detected among rivers than among ponds which might be related to limitations on gene flow. Demographic and temporal genetic data further indicates that flooding in rivers is unlikely to induce marked gene flow explaining the strong genetic differentiation at short geographical scale in such habitats. Finally, the demographic data suggest that some populations are transitory and subject to recurrent recolonization, a pattern that was also detected through genetic data.     

A molecular phylogeography approach to biological invasions of the New World by parthenogenetic Thiarid snails

The parthenogenetic snail Melanoides tuberculata, present in tropical fresh waters of most of the Old World before 1950, has now invaded the Neotropical area. The phylogeography of this snail was studied to evaluate the pathways and number of such invasions. Because of parthenogenetic reproduction, individuals are structured into genetical clones. Within populations from both the original and invaded areas, several morphologically distinct clones (referred to as morphs) often coexist but the amount of genetic divergence among morphs is unknown. Individuals from 27 morphs and 40 populations world-wide were sequenced at two mitochondrial genes (12S and 16S). Our phylogenetic reconstruction suggests that (i) most of the morphological variation observed in the New World predates invasion, (ii) at least six independent introductions have occurred, and (iii) invasive clones are found throughout most of the phylogenetic tree and do not come from a particular region of the area of origin. Two ideas are discussed in the light of these results. The first lies with the specificities of parthenogenesis in an invasion context. While in sexual species, independently introduced populations eventually merge into a single invasive population, in a parthenogenetic species independently introduced clones have distinct invasion dynamics and possibly exclude each other. Second, although repeated invasions in Melanoides may have an impact on indigenous molluscan faunas, their most likely effect is the world-wide homogenization of the invasive taxon itself.     

Ontogenetic reaction norm for binary traits: the timing of phallus development in the snail Bulinus truncatus

The ontogenetic trajectory of plastic binary traits may provide valuable insights into their evolutionary rate of change. In this paper, the timing of the plastic response of a temperature-dependent sexual polymorphism, aphally, is investigated in the freshwater snail Bulinus truncatus. Aphally is defined as the loss of the male copulatory organ in otherwise hermaphroditic animals. Individuals from two inbred lines were switched at various times during their early development between 25 and 30 degrees C, and their phally status ascertained, in order to evaluate the parameters characterising the ontogenetic reaction norm of aphally to temperature. A series of nested models including parameters for the onset, offset, and the intensity of the response to temperature were fitted to the data, allowing for a wide range of reaction norms. One genotype did not show any variation in aphally ratio with switching temperature, while a switch-point model (onset and offset corresponding to the same developmental point in time) best fitted the second genotype. The results suggest that the plasticity of aphally is expressed before eggs hatch. Their consequences on the evolution of aphally are discussed. More generally, the methodology proposed here can be used to analyse variation in ontogenetic parameters of discrete traits.     

Evolutionary history and phylogeography of the schistosome-vector freshwater snail Biomphalaria glabrata based on nuclear and mitochondrial DNA sequences

The phylogeography of the freshwater snail Biomphalaria glabrata remains poorly known, although this species is the major vector of schistosomiasis in the New World. It was here investigated in South America and the Lesser Antilles, based on partial mitochondrial large ribosomal subunit (16S rDNA) and nuclear internal transcribed spacer-2 (ITS-2) gene sequences. Sampling included 17 populations from a large part of the current geographic range of the species (Brazil, Venezuela and Lesser Antilles). Substantial variability was detected, as well as a high amount of phylogenetically informative signal. The molecular phylogeny inferred splits B. glabrata into Northern and Southern clades separated by the Amazon river, and may even suggest a supra-specific status for B. glabrata. Brazilian populations were the most diverse and appeared basal to the other populations. Venezuelan haplotypes formed a single clade, albeit not strongly supported. Two Venezuelan haplotypes appear rather similar to Brazilian haplotypes. Similarly, Lesser Antilles haplotypes clustered in the same monophyletic clade, which suggests that the recent colonisation of the Antilles has a northern South American origin. However, the estimated divergence time between Antilles and Venezuelan sequences is extremely large (conservatively higher than 10(5) years). These results are discussed in the light of (i) phylogeographic patterns at South American scale, and (ii) recurrent introduction of molluscs, especially in the Antilles, as a consequence of human activities.     

The Effects of a Bottleneck on Inbreeding Depression and the Genetic Load

We study the effects of a population bottleneck on the inbreeding depression and genetic load caused by deleterious mutations in an outcrossing population. The calculations assume that loci have multiplicative fitness effects and that linkage disequilibrium is negligible. Inbreeding depression decreases immediately after a sudden reduction of population size, but the drop is at most only several percentage points, even for severe bottlenecks. Highly recessive mutations experience a purging process that causes inbreeding depression to decline for a number of additional generations. On the basis of available parameter estimates, the absolute fall in inbreeding depression may often be only a few percentage points for bottlenecks of 10 or more individuals. With a very high lethal mutation rate and a very slow population growth, however, the decline may be on the order of 25%. We examine when purging might favor a switch from outbreeding to selfing and find it occurs only under very limited conditions unless population growth is very slow. In contrast to inbreeding depression, a bottleneck causes an immediate increase in the genetic load. Purging causes the load to decline and then overshoot its equilibrium value. The changes are typically modest: the absolute increase in the total genetic load will be at most a few percentage points for bottlenecks of size 10 or more unless the lethal mutation rate is very high and the population growth rate very slow.