Isolation by distance and sharp discontinuities in gene frequencies: implications for the phylogeography of an alpine insect species, Carabus solieri

Analysis of genetic isolation by distance (IBD) is of prime importance for the study of processes responsible for spatial population genetic structure and is thus frequently used in case studies. However, the identification of a significant IBD pattern does not necessarily imply the absence of sharp discontinuities in gene frequencies. Therefore, identifying barriers to gene flow and/or secondary contact between differentiated entities remains a major challenge in population biology. Geographical genetic structure of 41 populations (1080 individuals) of an alpine insect species, Carabus solieri, was studied using 10 microsatellite loci. All populations were significantly differentiated and spatially structured according to IBD over the entire range. However, clustering analyses clearly identified three main clusters of populations, which correspond to geographical entities. Whereas IBD also occurs within each cluster, population structure was different according to which group of populations was considered. The southernmost cluster corresponds to the most fragmented part of the range. Consistently, it was characterized by relatively high levels of differentiation associated with low genetic diversity, and the slope of the regression of genetic differentiation against geographical distances was threefold those of the two other clusters. Comparisons of within-cluster and between-cluster IBD patterns revealed barriers to gene flow. A comparison of the two approaches, IBD and clustering analyses, provided us with valuable information with which to infer the phylogeography of the species, and in particular to propose postglacial colonization routes from two potential refugia located in Italy and in southeastern France. Our study highlights strongly the possible confounding contribution of barriers to gene flow to IBD pattern and emphasizes the utility of the model-based clustering analysis to identify such barriers.     

Genetic consequences of Pleistocene glaciations for the tundra vole (Microtus oeconomus) in Beringia

Repeated glacial events during the Pleistocene fragmented and displaced populations throughout the northern continents. Different models of the effects of these climate-driven events predict distinct phylogeographic and population genetic outcomes for high-latitude faunas. The role of glaciations in (i) promoting intraspecific genetic differentiation and (ii) influencing genetic diversity was tested within a phylogeographic framework using the rodent Microtus oeconomus. The spatial focus for the study was Beringia, which spans eastern Siberia and northwestern North America, and was a continental crossroads and potential high arctic refugium during glaciations. Variation in mitochondrial DNA (cytochrome b and control region; 214 individuals) and nuclear DNA (ALDH1 intron; 63 individuals) was investigated across the Beringian region. Close genetic relationships among populations on either side of the Bering Strait are consistent with a history of periodic land connections between North America and Asia. A genetic discontinuity observed in western Beringia between members of a Central Asian clade and a Beringian clade is geographically congruent with glacial advances and with phylogeographic discontinuities identified in other organisms. Divergent island populations in southern Alaska were probably initially isolated by glacial vicariance, but subsequent differentiation has resulted from insularity. Tests of the genetic effects of postglacial colonization were largely consistent with expansion accompanied by founder effect bottlenecking, which yields reduced diversity in populations from recently deglaciated areas. Evidence that populations in the Beringian clade share a history of expansion from a low-diversity ancestral population suggests that Beringia was colonized by a small founder population from central Asia, which subsequently expanded in isolation.     

Past and current gene flow in the selfing, wind-dispersed species Mycelis muralis in western Europe

The distribution of genetic diversity in Mycelis muralis, or wall lettuce, was investigated at a European scale using 12 microsatellite markers to infer historical and contemporary forces from genetic patterns. Mycelis muralis has the potential for long-distance seed dispersal by wind, is mainly self-pollinated, and has patchily distributed populations, some of which may show metapopulation dynamics. A total of 359 individuals were sampled from 17 populations located in three regions, designated southern Europe (Spain and France), the Netherlands, and Sweden. At this within-region scale, contemporary evolutionary forces (selfing and metapopulation dynamics) are responsible for high differentiation between populations (0.34 < F(ST) < 0.60) but, contrary to expectation, levels of within-population diversity, estimated by Nei's unbiased expected heterozygosity (H(E)) (0.24 < H(E) < 0.68) or analyses of molecular variance (50% of the variation found within-populations), were not low. We suggest that the latter results, which are unusual in selfing species, arise from efficient seed dispersal that counteracts population turnover and thus maintains genetic diversity within populations. At the European scale, northern regions showed lower allelic richness (A = 2.38) than populations from southern Europe (A = 3.34). In light of postglacial colonization hypotheses, these results suggest that rare alleles may have been lost during recolonization northwards. Our results further suggest that mutation has contributed to genetic differentiation between southern and northern Europe, and that Sweden may have been colonized by dispersers originating from at least two different refugia.     

Genetic structure of kestrel populations and colonization of the Cape Verde archipelago

Genetic diversity and population structure were studied in eight populations of the kestrel Falco tinnunculus to identify the genetic consequences of spatial distribution and to infer the colonization patterns of the Cape Verde archipelago. We studied genetic differentiation and gene flow among seven island populations and one mainland population using nine microsatellite loci. Within the archipelago, differentiation was strong and genetic diversity and heterozygosity were low but variable among populations. Two subspecies F. tinnunculus neglectus on the northwestern islands and F. tinnunculus alexandri on all the other islands were identified as genetically distinct units. F. t. alexandri could be further separated into two groups on eastern and southern islands. Populations are probably founded by birds originating from the mainland. Immigration is more likely to the eastern and southern populations, whereas the northwestern islands with the lowest genetic diversity and highest differentiation are likely to exhibit fewer founding events by immigrants. The number of founding events on each island may depend not only on geographical distance to neighbouring populations, but also on directional immigration due to the northeastern trade winds. This may explain differences in genetic differentiation and diversity between populations and subspecies and may enable allopatric speciation.     

Long-distance colonization,isolation by distance,and historical demography in a relictual Mexican pinyon pine (Pinus nelsonii Shaw) as revealed by paternally inherited genetic markers (cpSSRs)

Pinus nelsonii is a relictual pinyon pine distributed across a wide altitudinal range in semiarid zones in Mexico near the border between the States of Nuevo León and Tamaulipas. It also occurs in small patches in the State of San Luis Potosí. Pinus nelsonii is classified in the monotypic subsection Nelsoniae, separated from other pinyon pines (subsection Cembroides), because it possesses several distinctive characters including persistent fascicle sheaths, connate needles, and a distinctive wood anatomy. In the present study, chloroplast simple sequence repeats (cpSSRs) were used to estimate genetic variation in most known populations (nine) of P. nelsonii. The genetic variation (HT = 0.73; 27 haplotypes in 256 individuals) is moderate when compared to other pine species. Population differentiation ranged between low and moderate (FST = 0.13 and RST = 0.05), as did the Nei and Goldstein genetic distances between populations. However, this pattern varied depending on whether the infinite alleles or stepwise mutation model was used. In the former case a significant isolation by distance was found, but not in the latter. A significant association between geographical and genetic structure in one clade, through a nested clade analysis, was found, which suggested long-distance colonization between 125000 and 309000 years ago. We found weak evidence for a population expansion. A mismatch distribution suggests that P. nelsonii populations underwent an expansion 4.25 times their size between 59000 and 146000 years ago. On the other hand, the populations' star-like phylogeny and a slight parabolic relationship between coalescence times and lineage number also suggest weak population expansion. Overall, this species appears to have been in demographic stasis for a large proportion of the time detected by the markers used.     

Evidence for male-biased effective sex ratio and recent step-by-step colonization in the bivalve Pinctada mazatlanica

This paper presents a comparison of the geographical distribution of genetic variability at mitochondrial and nuclear loci among pearl oyster populations from the tropical American Pacific coast (Pinctada mazatlanica). Surprisingly, both mitochondrial and nuclear gene variability decreased regularly from north to south of the studied area, which, altogether with a significant correlation between genetic and geographical distances for mtDNA, suggests a recent colonization or re-colonization of the southern areas. However, the loss of diversity between north and south was much more important for mitochondrial than for nuclear DNA, and this did not translate into measurable fixation index at nuclear loci (theta = 0.03, n.s.), contrary to the mitochondrial data (theta = 0.18*). Smaller effective size of mtDNA accentuated by a strong male-biased effective sex ratio and step-by-step colonization from northern areas can explain this discrepancy among natural populations of this protandric species.     

Radiation, diversity, and host-plant interactions among island and continental legume-feeding psyllids

Island archipelagos and insect-plant associations have both independently provided many useful systems for evolutionary study. The arytainine psyllid (Sternorrhyncha: Hemiptera) radiation on broom (Fabaceae: Genisteae) in the Canary Island archipelago provides a discrete system for examining the speciation of highly host-specific phytophagous insects in an island context. Phylogenetic reconstructions based on three datasets (adult and nymph morphological characters, and two mitochondrial DNA regions: part of the small subunit rRNA, and part of cytochrome oxidase I, cytochrome oxidase II and the intervening tRNA leucine) are generally consistent. The combined molecular tree provides a well-supported estimate of psyllid relationships and shows that there have been several colonizations of the Macaronesian islands but that only one has resulted in a significant radiation. Psyllid diversification has apparently been constrained by the presence of suitable host groups within the genistoid legumes, and the diversity, distribution, and abundance of those groups. The phylogeny, by indicating pairs of sister species, allows putative mechanisms of speciation to be assessed. The most common conditions associated with psyllid speciation are geographical allopatry with a host switch to closely related hosts (six examples), or geographical allopatry on the same host (four examples). Where allopatric speciation involves a host switch, these have all been to related hosts. There is some evidence that switches between unrelated host plants may be more likely in sympatry. Only one sister pair (Aryrtainilla cytisi and A. telonicola) and the putative host races of Arytinnis modica are sympatric but on unrelated hosts, which may be a necessary condition for sympatric speciation in these insects. Where several psyllids share the same host, resources appear to be partitioned by ecological specialization and differing psyllid phenology.     

Breaking taboos in the tropics: incest promotes colonization by wood-boring beetles

• 1 Inbreeding and parthenogenesis are especially frequent in colonizing species of plants and animals, and inbreeding in wood‐boring species in the weevil families Scolytinae and Platypodidae is especially common on small islands. In order to study the relationship between colonization success, island attributes and mating system in these beetles, we analysed the relative proportions of inbreeders and outbreeders for 45 Pacific and Old World tropical islands plus two adjacent mainland sites, and scored islands for size, distance from nearest source population, and maximum altitude.
• 2 The numbers of wood‐borer species decreased with decreasing island size, as expected; the degree of isolation and maximum island altitude had negligible effects on total species numbers.
• 3 Numbers of outbreeding species decreased more rapidly with island size than did those of inbreeders. Comparing species with similar ecology (e.g. ambrosia beetles) showed that this difference was best explained by differential success in colonization, rather than by differences in resource utilization or sampling biases. This conclusion was further supported by analyses of data from small islands, which suggested that outbreeding species have a higher degree of endemism and that inbreeding species are generally more widespread.
• 4 Recently established small populations necessarily go through a period of severe inbreeding, which should affect inbreeding species much less than outbreeding ones. In addition, non‐genetic ecological and behavioural (‘Allee’) effects are also expected to reduce the success of outbreeding colonists much more than that of inbreeders: compared with inbreeders, outbreeders are expected to have slower growth rates, have greater difficulties with mate‐location and be vulnerable to random extinction over a longer period.

     

The impact of stocking on the genetic integrity of Arctic charr (Salvelinus) populations from the Alpine region

There is a long tradition of artificially stocking lakes and rivers with fish in the hope to increase the quality and amount of fish that can be harvested. The animals used for stocking often originate in distant hatcheries or hatcheries that breed fish from remote regions. This stocking practice could have effects on the genetic integrity of resident populations. We have analysed here a case of the influence of stocking on Alpine populations of the Arctic charr (usually included into Salvelinus alpinus, but revised to Salvelinus umbla Kottelat 1997 ) within a unique geographical and historical setting. The Königssee in the Bavarian Alps (Berchtesgaden) was heavily stocked several times during the last century. However, a sample of the ancestral Königssee population still exists in the Grünsee, which lies close to Königssee, but 1000 m higher. To trace the influence of stocking in Königssee we have analysed more than 300 individuals from 10 lake populations, including the source populations of the fish used for stocking. From these we have sequenced a part of the mitochodrial control region and have typed them at six microsatellite loci. The differential distribution of haplotypes, as well as assignment tests, show that the influence of stocking on the genetic integrity of the Königssee population has been negligible. However, our data reveal that in another lake included in our study (Starnberger See), the ancestral population was apparently replaced completely by the populations used for stocking. The major difference between the lakes is the relative preservation of ecological integrity. Königssee was ecologically stable in the past, whereas Starnberger See was heavily polluted at one point, with charr approaching extinction. Interestingly, in a lake neighbouring Starnberger See, the Ammersee, which was also subject to strong pollution but not stocked, the ancestral population has recovered. Our data suggest that the practice of artificial stocking should be reconsidered, or at least monitored for effectiveness.     

Individual-based modelling of Pinus sylvestris invasion after grazing abandonment in the French Massif Central

In the Chaîne des Puys, a mid-elevation volcanic mountain of the French Massif Central, Scots pine proves to be an invasive species colonizing abandoned lawns or heathlands, and forms in a few years monospecific natural forests. Most of the abandonment occurred 30 to 40 years ago and this process has now stopped. Thus, we lack data on the very first phase of tree colonization. We anticipate that a simulation tool could bring an appreciable help in (i) rebuilding the entire colonization process – including the initial phase – of pine settlement and (ii) answering questions about the origin of the narrow and unimodal distributions of age of pine stands we observed. In addition, such a simulator could help managers to forecast extension of Scots pine and to predict growth and evolution of present secondary forests. A spatially explicit individual-based model is presented. The model takes into account both space and time and includes growth of trees, seed production and seed dispersal, death and competition between individuals. The influence of the initial parameters are analyzed and elements of validation given. The model was then used to predict tree settlement and stand establishment using the initial conditions from a natural stand studied in the field whose characteristics before abandonment were known (number and age-distribution of trees, location of mother trees, time of abandonment). Three simulations were achieved by using the same initial conditions but following different scenarios for the recruitment process. The scenario of a fluctuating resistance of the resident vegetation (that controls the susceptibility of the environment to tree establishment) seemed as one of the most probable to explain the actual stand characteristics. We thus concluded that dynamic models could be improved by taking into account the resistance of the vegetation to colonization as a fluctuating parameter instead of a static and permanent attribute.