Ecological strategies in the ancient asexual animal group Darwinulidae (Crustacea, Ostracoda)

• 1 We investigated the relationship between geographical distribution and ecological tolerance within the ancient asexual family Darwinulidae. Distribution maps were compiled based on data from the literature, the Non‐marine Ostracod Distribution in Europe database and personal collections. Ecological tolerance was assessed experimentally by exposing individual ostracods to a combination of eight different salinities (range from 0 to 30 g L^?1) and three different temperatures (10, 20 and 30 °C).
• 2 The type species of the family, Darwinula stevensoni, is ubiquitous and cosmopolitan; the two species Penthesilenula brasiliensis and Microdarwinula zimmeri also have an intercontinental distribution. Two other darwinulid species tested here (Vestalenula molopoensis and P. aotearoa) are known only from their type localities. The latter is also true for most extant darwinulids.
• 3 Darwinula stevensoni and P. brasiliensis had a broad salinity tolerance, tolerating distilled water and also salinity up to 25–30 g L^?1, whereas the maximum salinity tolerance of V. molopoensis was 12 g L^?1 and of P. aotearoa, 20 g L^?1.
• 4 The results indicate that both ecological specialists and generalists, as well as intermediate forms, exist in the Darwinulidae and that taxa with the broadest ecological tolerance also have the widest distribution.

     

Diverse,endemic and polyphyletic clones in mixed populations of a freshwater snail (Potamopyrgus antipodarum)

Understanding the source and diversity of clones is necessary to resolve the complicated issues surrounding the apparent evolutionary stability of sexual reproduction. The source of clones is important because present theory is based on an “all else equal” assumption, which is predicated on the idea that clonal mutants are derived from and compete with local sexual populations. Clonal diversity is important because it reduces the advantage of sexual reproduction under either soft selection (the Tangled Bank Hypothesis) or under strict frequency-dependent selection (the Red Queen Hypothesis). In the present study, protein electrophoresis was used to determine the source and diversity of clones in a freshwater snail (Potamopyrgus antipodarum) in four glacial lakes in which sexual and clonal females were thought to coexist. The results showed (1) that the populations were mixtures of diploid sexual and triploid asexual individuals, (2) that genotypic diversity of clonal populations is very high in all four lakes (but lower than in the sympatric sexual populations), and (3) that the clones are polyphyletically derived from their sympatric sexual populations. Consequently, repeated mutation to parthenogenetic reproduction since the Pleistocene has introduced a different and diverse set of clones in all four lakes. Such diversity may provide a challenge for the ecological theories of sex that rely on frequency-dependent selection.     

Clonal diversity and polyphyletic origins of hybrid and spontaneous parthenogenetic Campeloma (Gastropoda: Viviparidae) from the south-eastern United States

Some theories for the maintenance of sexual reproduction indicate that parthenogens may persist if there is high clonal diversity and high dispersal rates. Using allozymic variation, we report on the origin, clonal diversity and population structure of hybrid and spontaneous parthenogens from south-eastern United States populations of the freshwater snail Campeloma. Independent origins of triploid hybrid parthenogens in the Florida panhandle occurred by hybridization between an Atlantic coastal species (C. limum or C. floridense) and the Florida sexual species (C. geniculum). Allozyme genotypic diversity is similar between these hybrid parthenogens and sexuals. Diploid spontaneous parthenogens originated multiple times from nonlocal C.limum sexual populations in Atlantic coastal rivers, and levels of genotypic diversity are significantly higher in sexual C. limum. How parthenogens originate, the degree of clonal diversity, and their subsequent dispersal influence whether basic assumptions of evolution-of-sex models are met.     

Improbable but true: the invasive inbreeding ambrosia beetle Xylosandrus morigerus has generalist genotypes

The wide distribution and dominance of invasive inbreeding species in many forest ecosystems seems paradoxical in face of their limited genetic variation. Successful establishment of invasive species in new areas is nevertheless facilitated by clonal reproduction: parthenogenesis, regular self-fertilization, and regular inbreeding. The success of clonal lineages in variable environments has been explained by two models, the frozen niche variation (FNV) model and the general-purpose genotype (GPG) model. We tested these models on a widely distributed forest pest that has been recently established in Costa Rica-the sibling-mating ambrosia beetle Xylosandrus morigerus. Two deeply diverged mitochondrial haplotypes coexist at multiple sites in Costa Rica. We find that these two haplotypes do not differ in their associations with ecological factors. Overall the two haplotypes showed complete overlap in their resource utilization; both genotypes have broad niches, supporting the GPG model. Thus, probable or not, our findings suggest that X. morigerus is a true ecological generalist. Clonal aspects of reproduction coupled with broad niches are doubtless important factors in the successful colonization of new habitats in distant regions.     

Adaptation in the Asexual False Spider Mite <Emphasis Type="Italic">Brevipalpus phoenicis</Emphasis>: Evidence for Frozen Niche Variation

Because asexual species lack recombination, they have little opportunity to produce genetically variable offspring and cannot adapt to changes in their environment. However, a number of asexual species are very successful and appear to contradict this general view. One such species is the phytophagous mite Brevipalpus phoenicis (Geijskes), a species that is found in a wide range of environments. There are two general explanations for this pattern, the General Purpose Genotype (GPG) and Frozen Niche Variation (FNV). According to the GPG model, an asexual species consists of clones that can all survive and reproduce in all the different niches. Alternatively, the FNV model postulates that different clones are specialized to different niches. We have performed a test to distinguish between these models in B. phoenicis. Mites from three populations from three different host plant species (citrus, hibiscus and acerola) were transplanted to their own and the two alternative host plants and mite survival and egg production were measured. Additionally, the mite populations were genotyped using microsatellites. Fitness was seriously reduced when mites were transplanted to the alternative host plant species, except when the alternative host was acerola. We concluded that B. phoenicis clones are specialized to different niches and thus the FNV best describes the broad ecological niche of this species but that there is also some evidence for host plant generalization. This conclusion was strengthened by the observations that on each host plant species the native mite population performed better than the introduced ones, and that three microsatellite markers showed that the mite populations are genetically distinct.     

Environmental gradients structure Daphnia pulex × pulicaria clonal distribution

The rarity of eukaryotic asexual reproduction is frequently attributed to the disadvantage of reduced genetic variation relative to sexual reproduction. However, parthenogenetic lineages that evolved repeatedly from sexual ancestors can generate regional pools of phenotypically diverse clones. Various theories to explain the maintenance of this genetic diversity as a result of environmental and spatial heterogeneity [frozen niche variation (FNV), general-purpose genotype] are conceptually similar to community ecological explanations for the maintenance of regional species diversity. We employed multivariate statistics common in community ecological research to study population genetic structure in the freshwater crustacean, Daphnia pulex × pulicaria. This parthenogenetic hybrid arose repeatedly from sexual ancestors. Daphnia pulex × pulicaria populations harboured substantial genetic variation among populations and the clonal composition at each pond corresponded to nutrient levels and invertebrate predator densities. The interclonal selection process described by the FNV hypothesis likely structured our D. pulex × pulicaria populations.     

Are hybridogenetic complexes structured by habitat in water frogs?

The success and the evolutionary fate of hybridogenetic lineages are explained by both a generalistic heterosis hypothesis and an alternative hypothesis, the habitat segregation hypothesis. Because such hypotheses have rarely been tested at the level of whole habitats, our aim was to compare performances of two taxa within a hybridogenetic complex across diverse natural habitats. We took advantage of the waterfrog hybridogenetic complex (Rana esculenta and R. lessonae) by rearing tadpoles in natural contrasted habitats by means of enclosure experiments. We also monitored the frequency of each taxon in the waterfrog assemblages that naturally breed in the studied ponds. The hybridogenenetic taxon showed no evidence of broader tolerance as growth, development and physiology strongly varied in response to environmental heterogeneity. Our study reveals a differential success of the hybridogenetic taxon and its sexual host among environments. Moreover, hybridogenetic taxa rarely dominated the sexual species in natural assemblages. Consequently, our results show that the generalistic model does not explain the success of hybridogenetic lineages, but rather support the habitat segregation, among other alternative concepts.     

Temporal dynamics of genotypic diversity reveal strong clonal selection in the aphid Myzus persicae

Parthenogenetic organisms often harbour substantial genotypic diversity. This diversity may be the result of recurrent formations of new clones, or it may be maintained by environmental heterogeneity acting on ecological differences among clones. In aphids, both processes may be important because obligate and cyclical parthenogens can form mixed populations. Using microsatellites, I analysed the temporal dynamics of clonal diversity in such a population of the aphid Myzus persicae over a 1-year period. The frequency distribution of clonal genotypes was very skewed, with many rare and few common clones. The relative frequencies of common clones underwent strong and rapid changes indicative of intense clonal selection. Differences in their host associations suggest that these shifts may partly be caused by changes in the abundance of annual host plants. Other selective factors of potential importance are also discussed. New, sexually produced genotypes made a minor contribution to clonal diversity, consistent with the observed heterozygote excess characteristic of predominantly asexual populations in M. persicae.     

Vertical and horizontal transmission drive bacterial invasion

A huge variety of Arthropod species is infected with endosymbiotic Wolbachia bacteria that manipulate their host’s reproduction to invade populations. In addition to vertical transmission from mother to offspring through the egg cytoplasm, it has been demonstrated through phylogenetic analyses and natural transfer experiments that horizontal transmission of Wolbachia (i.e. contagion) can occur between Arthropod hosts. More recently, factors influencing horizontal transfer have also been explored. While it is clear that horizontal transmission between species plays a major role in the evolutionary history of Wolbachia infections among insects, its role in the spread of a new infection through a host population, notably through within‐species transfers, remained unknown. In this issue of Molecular Ecology, Kraaijeveld et al. (2011) present the first evidence that horizontal transmission played a key role in the early spread of parthenogenesis‐inducing Wolbachia through the parasitoid wasp Leptopilina clavipes. To support their finding, the authors studied genetic variation in three types of markers, including host nuclear DNA, mitochondrial DNA and Wolbachia DNA. Specifically, they examined potential associations between their diversity patterns. No diversity was detected in Wolbachia genes, indicating that a single Wolbachia strain must have infected and spread through L. clavipes. In addition, a correlation between substantial variation in mitochondrial and nuclear genotypes suggested that horizontal transmission played an important role in the current clonal genetic variation in this wasp. Such horizontal transmission could be facilitated by a specific host ecology (e.g. parasitoid wasps sharing the same host resource) and potentially impact co‐evolution between host and symbiont.     

Asexual and sexual reproductive strategies in clonal plants

Most plants can reproduce both sexually and asexually (or vegetatively),and the balance between the two reproductive modes may vary widely between and within species.Extensive clonal growth may affect the evolution of life history traits in many ways.First,in some clonal species,sexual reproduction and sex ratio vary largely among populations.Variation in sexual reproduction may strongly affect plant's adaptation to local environments and the evolution of the geographic range.Second,clonal growth can increase floral display,and thus pollinator attraction,while it may impose serious constraints and evolutionary challenges on plants through geitonogamy that may strongly influence pollen dispersal.Geitonogamous pollination can bring a cost to plant fitness through both female and male functions.Some co-evolutionary interactions,therefore,may exist between the spatial structure and the mating behavior of clonal plants.Finally,a trade-off may exist between sexual reproduction and clonal growth.Resource allocation to the two reproductive modes may depend on environmental conditions,competitive dominance,life span,and genetic factors.If different reproductive modes represent adaptive strategies for plants in different environments,we expect that most of the resources should be allocated to sexual reproduction in habitats with fluctuating environmental conditions and strong competition,while clonal growth should be dominant in stable habitats.Yet we know little about the consequence of natural selection on the two reproductive modes and factors which control the balance of the two reproductive modes.Future studies should investigate the reproductive strategies of clonal plants simultaneously from both sexual and asexual perspectives.