Genetic differentiation and reduced genetic diversity at the northern range edge of two species with different dispersal modes

Theory predicts that genetic variation should be reduced at range margins, but empirical support is equivocal. Here, we used genotyping‐by‐sequencing technology to investigate genetic variation in central and marginal populations of two species in the marine gastropod genus Crepidula. These two species have different development and dispersal types and might therefore show different spatial patterns of genetic variation. Both allelic richness and the proportion of private alleles were highest in the most central populations of both species, and lower at the margin. The species with low dispersal, Crepidula convexa, showed high degrees of structure throughout the range that conform to the pattern found in previous studies using other molecular markers. The northernmost populations of the high‐dispersing species, Crepidula fornicata, are distinct from more central populations, although this species has been previously observed to have little genetic structure over much of its range. Although genetic diversity was significantly lower at the range margin, the absolute reduction in diversity observed with these genomewide markers was slight, and it is not yet known whether there are functional consequences for the marginal populations.     

Homogenous Population Genetic Structure of the Non-Native Raccoon Dog (Nyctereutes procyonoides) in Europe as a Result of Rapid Population Expansion

The extent of gene flow during the range expansion of non-native species influences the amount of genetic diversity retained in expanding populations. Here, we analyse the population genetic structure of the raccoon dog (Nyctereutes procyonoides) in north-eastern and central Europe. This invasive species is of management concern because it is highly susceptible to fox rabies and an important secondary host of the virus. We hypothesized that the large number of introduced animals and the species’ dispersal capabilities led to high population connectivity and maintenance of genetic diversity throughout the invaded range. We genotyped 332 tissue samples from seven European countries using 16 microsatellite loci. Different algorithms identified three genetic clusters corresponding to Finland, Denmark and a large ‘central’ population that reached from introduction areas in western Russia to northern Germany. Cluster assignments provided evidence of long-distance dispersal. The results of an Approximate Bayesian Computation analysis supported a scenario of equal effective population sizes among different pre-defined populations in the large central cluster. Our results are in line with strong gene flow and secondary admixture between neighbouring demes leading to reduced genetic structuring, probably a result of its fairly rapid population expansion after introduction. The results presented here are remarkable in the sense that we identified a homogenous genetic cluster inhabiting an area stretching over more than 1500km. They are also relevant for disease management, as in the event of a significant rabies outbreak, there is a great risk of a rapid virus spread among raccoon dog populations.     

Genetic analysis across different spatial scales reveals multiple dispersal mechanisms for the invasive hydrozoan Cordylophora in the Great Lakes

Discerning patterns of post‐establishment spread by invasive species is critically important for the design of effective management strategies and the development of appropriate theoretical models predicting spatial expansion of introduced populations. The globally invasive colonial hydrozoan Cordylophora produces propagules both sexually and vegetatively and is associated with multiple potential dispersal mechanisms, making it a promising system to investigate complex patterns of population structure generated throughout the course of rapid range expansion. Here, we explore genetic patterns associated with the spread of this taxon within the North American Great Lakes basin. We collected intensively from eight harbours in the Chicago area in order to conduct detailed investigation of local population expansion. In addition, we collected from Lakes Michigan, Erie, and Ontario, as well as Lake Cayuga in the Finger Lakes of upstate New York in order to assess genetic structure on a regional scale. Based on data from eight highly polymorphic microsatellite loci we examined the spatial extent of clonal genotypes, assessed levels of neutral genetic diversity, and explored patterns of migration and dispersal at multiple spatial scales through assessment of population level genetic differentiation (pairwise F_ST and factorial correspondence analysis), Bayesian inference of population structure, and assignment tests on individual genotypes. Results of these analyses indicate that Cordylophora populations in this region spread predominantly through sexually produced propagules, and that while limited natural larval dispersal can drive expansion locally, regional expansion likely relies on anthropogenic dispersal vectors.     

Biological Invasions Across Spatial Scales: Intercontinental, Regional, and Local Dispersal of Cladoceran Zooplankton

The frequency of dispersal of invertebrates among lakes depends upon perspective and spatial scale. Effective passive dispersal requires both the transport of propagules and the establishment of populations large enough to be detected. At a global scale, biogeographic patterns of cladoceran zooplankton species suggest that effective dispersal among continents was originally rare, but greatly increased in the past century with expanded commerce. Genetic analysis allows some reconstruction of past dispersal events. Allozyme and mitochondrial DNA comparisons among New World and Old-World populations of several exotic cladocerans have provided estimates for likely source populations of colonists, their dispersal corridors, and timing of earlier dispersal events. Detecting the Old-World tropical exotic Daphnia lumholtzi early in its invasion of North America has allowed detailed analysis of its spatial spread. Twelve years of collection records indicate a rapid invasion of reservoirs in the United States, by both regional spread and long-distance jumps to new regions. Combining landscape features with zooplankton surveys from south-central US reservoirs revealed higher colonization rates of D. lumholtzi at lower landscape positions, a result which can be explained by either greater propagule load or by higher susceptibility of these downstream reservoirs. Because invaded reservoirs provide a source of propagules for nearby floodplain ponds, the rarity of this species in ponds suggests limitation by local environments. Such analyses of invading species over multiple spatial scales allow a better understanding of ecological processes governing invasion dynamics.     

Spatial extent of analysis influences observed patterns of population genetic structure in a widespread darter species (Percidae)

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Population genetic structure of Bellamya aeruginosa (Mollusca: Gastropoda: Viviparidae) in China: weak divergence across large geographic distances

Bellamya aeruginosa is a widely distributed Chinese freshwater snail that is heavily harvested, and its natural habitats are under severe threat due to fragmentation and loss. We were interested whether the large geographic distances between populations and habitat fragmentation have led to population differentiation and reduced genetic diversity in the species. To estimate the genetic diversity and population structure of B. aeruginosa, 277 individuals from 12 populations throughout its distribution range across China were sampled: two populations were sampled from the Yellow River system, eight populations from the Yangtze River system, and two populations from isolated plateau lakes. We used seven microsatellite loci and mitochondrial cytochrome oxidase I sequences to estimate population genetic parameters and test for demographic fluctuations. Our results showed that (1) the genetic diversity of B. aeruginosa was high for both markers in most of the studied populations and effective population sizes appear to be large, (2) only very low and mostly nonsignificant levels of genetic differentiation existed among the 12 populations, gene flow was generally high, and (3) relatively weak geographic structure was detected despite large geographic distances between populations. Further, no isolation by linear or stream distance was found among populations within the Yangtze River system and no signs of population bottlenecks were detected. Gene flow occurred even between far distant populations, possibly as a result of passive dispersal during flooding events, zoochoric dispersal, and/or anthropogenic translocations explaining the lack of stronger differentiation across large geographic distances. The high genetic diversity of B. aeruginosa and the weak population differentiation are likely the results of strong gene flow facilitated by passive dispersal and large population sizes suggesting that the species currently is not of conservation concern.     

Age and size of European saltmarshes and the population genetic consequences for ground beetles

Enzyme and dispersal polymorphisms of the saltmarsh carabid beetles Pogonus chalceus and Dicheirotrichus gustavii were studied in European populations varying in size and in isolation in space and time. D. gustavii, a constantly fullwinged species, has a larger genetic diversity and a smaller genetic differentiation between populations than the wing-polymorphic P. chalceus. Clear relationships between population or site characteristics and genetic structure were not observed, except for the special position taken by some small populations in both species. The dispersal power of P. chalceus in small populations is larger than in large populations, suggesting that these populations are unstable and/or young. Small populations, however, do not always show a lower genetic diversity than large populations, as would be expected from genetic drift. Dispersal power in P. chalceus declines with increasing age of the saltmarsh, probably due to continuous emigration of winged individuals. Age and size of saltmarshes, although difficult to study independently, both appear to be important in determining the genetic structure of saltmarsh beetles. Maximum diversity in both parameters is therefore recommended as an optimal nature conservation strategy. Received: 22 September 1997 / Accepted: 29 December 1997     

Dispersal and genetic differentiation of Syntrichia caninervis populations across different desert regions in China

• The moss Syntrichia caninervis is widely distributed in cool temperate and cold deserts where environmental pressures create a dependence on asexual reproduction (fragment reproduction). However, when compared to sporophyte‐producing mosses, there is a lack of evidence to support the capacity of drought‐tolerant mosses that predominantly fragment and produce protonema to disperse over long distances.
• We used 20 microsatellite loci to study genetic variation and structure in six populations (five natural populations and one population from a regeneration site) in three contrasting and widely separated regions of China.
• The genetic diversity and expected heterozygosity were lower in populations from the Tengger Desert than in populations from the other regions. Using PCoA, UPGMA and Structure analysis, the genetic grouping divided the three regions into three distinct groups. This may indicate that in regions where S. caninervis reproduces predominantly asexually, propagules are spread mainly by short‐distance dispersal. The genetic diversity of the population from the regeneration site in the Tengger Desert was slightly higher than that of the nearby, naturally occurring population, and included some input from the Pamir Plateau almost 2,300 km to the west, suggesting long‐distance dispersal of S. caninervis propagules across the region.
• Predominantly asexually reproducing populations of S. caninervis are mainly dependent on short‐distance dispersal. Long‐distance dispersal of S. caninervis propagules across the region is difficult. Establishment of populations with dominant asexual reproduction will eventually result in genetic differentiation.

     

Population size,center–periphery,and seed dispersers’ effects on the genetic diversity and population structure of the Mediterranean relict shrub Cneorum tricoccon

The effect of population size on population genetic diversity and structure has rarely been studied jointly with other factors such as the position of a population within the species’ distribution range or the presence of mutualistic partners influencing dispersal. Understanding these determining factors for genetic variation is critical for conservation of relict plants that are generally suffering from genetic deterioration. Working with 16 populations of the vulnerable relict shrub Cneorum tricoccon throughout the majority of its western Mediterranean distribution range, and using nine polymorphic microsatellite markers, we examined the effects of periphery (peripheral vs. central), population size (large vs. small), and seed disperser (introduced carnivores vs. endemic lizards) on the genetic diversity and population structure of the species. Contrasting genetic variation (H_E: 0.04–0.476) was found across populations. Peripheral populations showed lower genetic diversity, but this was dependent on population size. Large peripheral populations showed high levels of genetic diversity, whereas small central populations were less diverse. Significant isolation by distance was detected, indicating that the effect of long‐distance gene flow is limited relative to that of genetic drift, probably due to high selfing rates (F_IS = 0.155–0.887), restricted pollen flow, and ineffective seed dispersal. Bayesian clustering also supported the strong population differentiation and highly fragmented structure. Contrary to expectations, the type of disperser showed no significant effect on either population genetic diversity or structure. Our results challenge the idea of an effect of periphery per se that can be mainly explained by population size, drawing attention to the need of integrative approaches considering different determinants of genetic variation. Furthermore, the very low genetic diversity observed in several small populations and the strong among‐population differentiation highlight the conservation value of large populations throughout the species’ range, particularly in light of climate change and direct human threats.     

The absence of genotypic diversity in a successful parthenogenetic invader

Invasiveness might depend on the ability of genetically diverse populations of exotic species to adapt to novel environments, which suggests a paradox since exotic species are expected to lose genetic diversity when introduced. The apparent need for genetic diversity is particularly important for exotic species that lack bi-parental reproduction and genetic recombination. We used genetic marker studies to determine the genotypic diversity of invasive US populations of the clonal New Zealand mudsnail (Potamopyrgus antipodarum). We report here on a three-pronged survey of allozyme, microsatellite DNA, and mitochondrial DNA genetic markers of invasive populations with a focus on the western US. Combining the three types of genetic markers, we discovered four distinct genotypes of P. antipodarum. These results show that only one genotype (US 1) occupied the vast majority of the western US range, and a second occurred in the Great Lakes in the eastern US (US 2). Two other genotypes occurred in the western US (US 1a and US 3), but were restricted to populations near the presumed source of invasion in the middle Snake River, ID. These results suggest that P. antipodarum spread across a broad geographic range in the western US from a single introduced source population, and that the populations are comprised of a single clonal lineage.     

High genetic divergence characterizes populations of the endemic plant Lithophragma maximum (Saxifragaceae) on San Clemente Island

Narrowly-ranging species frequently harbor less genetic variability relative to widespread relatives and face graver extinction threats due to the heightened impacts of stochastic events on ecological and genetic diversity. In this study, we examined the impact of historical and current threats to the maintenance of genetic variation in Lithophragma maximum (Saxifragaceae), a perennial herb endemic to San Clemente Island, California. This species exists as small populations confined to canyons along 4 km of the southeast coastline of the island. In 15 populations analyzed with 10 microsatellite markers, we identified an average of 2.05 alleles per locus and 58.7% polymorphic loci. Significant departures from Hardy–Weinberg equilibrium existed in six populations; five of these exhibited heterozygote deficiency. Bayesian inference of genetic structure indicated a significant amount of structure among populations and canyons and infrequent gene flow even over very short distances. We also identified a significant and positive correlation between genetic and geographic distances, indicative of isolation by distance. There was no evidence of recent bottlenecks in any of the sampled populations, but older bottlenecks were detected in two populations. These results suggest that populations of L. maximum have historically been small and isolated, which is likely due to the rugged habitat in which this species occurs and limited pollen and seed dispersal. Given the high degree of structure observed across populations, we suggest that conservation efforts should focus on preserving populations in multiple canyons, maintaining large population sizes to preserve genetic variation, and controlling the spread of invasive species in areas where L. maximum occurs.     

Scale‐dependent post‐establishment spread and genetic diversity in an invading mollusc in South America

Aims Our study aimed to characterize the dispersal dynamics and population genetic structure of the introduced golden mussel Limnoperna fortunei throughout its invaded range in South America and to determine how different dispersal methods, that is, human‐mediated dispersal and downstream natural dispersal, contribute to genetic variation among populations. Location Paraná–Uruguay–Río de la Plata watershed in Argentina, Brazil, Paraguay and Uruguay. Methods We performed genetic analyses based on a comprehensive sampling strategy encompassing 22 populations (N = 712) throughout the invaded range in South America, using the mitochondrial cytochrome c oxidase subunit I (COI) gene and eight polymorphic nuclear microsatellites. We employed both population genetics and phylogenetic analyses to clarify the dispersal dynamics and population genetic structure. Results We detected relatively high genetic differentiation between populations (F_ST = ?0.041 to 0.111 for COI, ?0.060 to 0.108 for microsatellites) at both fine and large geographical scales. Bayesian clustering and three‐dimensional factorial correspondence analyses consistently revealed two genetically distinct clusters, highlighting genetic discontinuities in the invaded range. Results of all genetic analyses suggest ship‐mediated ‘jump’ dispersal as the dominant mode of spread of golden mussels in South America, while downstream natural dispersal has had limited effects on contemporary genetic patterns. Main conclusions Our study provides new evidence that post‐establishment dispersal dynamics and genetic patterns vary across geographical scales. While ship‐mediated ‘jump’ dispersal dominates post‐establishment spread of golden mussels in South America, once colonies become established in upstream locations, larvae produced may be advected downstream to infill patchy distributions. Moreover, genetic structuring at fine geographical scales, especially within the same drainages, suggests a further detailed understanding of dynamics of larval dispersal and settlement in different water systems. Knowledge of the mechanisms by which post‐establishment spread occurs can, in some cases, be used to limit dispersal of golden mussels and other introduced species.     

GENETIC VARIATION IN LATHYRUS LATIFOLIUS (LEGUMINOSAE)

Lathyrus latifolius (everlasting pea) is a perennial vine native to Europe. Naturalized populations of L. latifolius occur in fields and on roadsides over large areas of the United States. Widely cultivated as a garden flower, L. latifolius produces abundant racemes of showy flowers that are bumblebee-pollinated. The seeds are heavy, large, and round, and exhibit no specialized means of dispersal. Allozyme diversity and population structure were determined for 32 populations of L. latifolius—30 from the southeastern United States and two from Oregon. Results from 21 allozyme loci indicate that genetic diversity is higher and population divergence is lower than expected based on the life history characteristics of the species. No association was found between genetic identity statistics and geographic distance between populations. Although the range in genetic diversity statistics among populations was unusually large, genetic drift did not appear to play a major role in structuring genetic variation. We conclude that the level of genetic diversity maintained within L. latifolius populations, and the level of population divergence found, is strongly influenced by its status as a cultivated garden flower and its human-associated mode of gene flow via seed dispersal.     

Patterns of population genetic diversity in riparian and aquatic plant species along rivers

Aim The downstream hydrochoric spread of seeds of aquatic and riparian plant species, without upstream compensation, can be expected to result in downstream accumulation of population genetic diversity. This idea has been termed the ‘unidirectional dispersal hypothesis’ and is the genetic equivalent of the more generally known ‘drift paradox’. Our aim was to test this unidirectional diversity hypothesis, and to present a general synthesis of the patterns of population genetic variation across different riparian and aquatic plant species along rivers. Location The Meuse River (Belgium) and rivers world‐wide. Methods First, we used amplified fragment length polymorphism markers to compare patterns of within‐ and between‐population genetic diversity among three riparian plant species (Sisymbrium austriacum, Erysimum cheiranthoides and Rorippa sylvestris), typically occurring in different habitats along a gradient perpendicular to the Meuse River. Second, we performed a meta‐analysis on studies reporting on the population genetic structure of riparian and aquatic plant species along rivers. Results Along the Meuse River, we found significant genetic differentiation among populations of all three riparian species, and significant isolation by distance for one of them (R. sylvestris). There was no clear association between the typical habitat of a species and its population genetic structure. None of the three species provided evidence for the unidirectional dispersal hypothesis. The meta‐analysis, based on 21 data records, did not support the unidirectional dispersal hypothesis either. Average weighted population genetic differentiation across species was significant. Main conclusions Important mechanisms of upstream seed dispersal, probably through zoochory, together with higher seed recruitment opportunities in upstream habitats due to density dependence of recruitment, may explain the absence of downstream accumulation of genetic diversity. Also, it seems difficult to find consistent patterns in genetic variation in species from aquatic and riparian habitats. We argue that this is due to the recurrent extinctions and colonizations characteristic of these habitats, resulting in complex genetic patterns. Our results strongly support previous suggestions that stream ecology should consistently embrace metapopulation theory to be able to understand patterns of genetic diversity, as well as species diversity.     

Multiple introduction events and artificial long-distance dispersal of the exotic slug Ambigolimax valentianus in Japan

ABSTRACT

The three-band garden slug Ambigolimax valentianus, native to the Iberian Peninsula, is an exotic species distributed worldwide and was introduced to Japan into the 1950s. For control of this slug, it is necessary to elucidate its introduction sources and dispersal patterns since the introduction event occurred. However, knowledge about this slug in Japan remains poor. Thus, we attempted to estimate the origin and dispersal patterns of this slug using genetic approaches. We investigated the genetic diversity of the mitochondrial cytochrome c oxidase subunit I gene across 11 populations of A. valentianus in three prefectures in central Japan. Ten haplotypes were found in the Japanese populations, five of which were haplotypes found in foreign individuals. The same haplotypes were found in all three prefectures, whereas the average haplotype diversity was high across all populations. Significant genetic differentiation was detected among 22 of 55 population pairs, but there was no isolation by distance. These molecular results suggest that multiple introductions to Japan may have occurred in the past such that the founder effect in the early stage of introduction was diluted. Furthermore, long-distance dispersal of A. valentianus would have occurred after introduction into Japan because of human activities rather than natural dispersal.     

Isolation and characterization of highly polymorphic microsatellite markers from the marine invasive species Crepidula fornicata (Gastropoda: Calyptraeidae)

Many marine organisms have pelagic larvae and these are often important agents of dispersal and establishment of exotic species. The larval phase and the hermaphroditic reproductive system of the invasive marine gastropod Crepidula fornicata are crucial in determining the success of this species along the European coasts. To analyse these factors, we developed seven microsatellite loci specific to C. fornicata. On the same set of individuals, the microsatellite loci exhibited a greater level of polymorphism than previously studied allozyme markers and, thus, will permit fine‐scale analysis and larval pool studies.     

Patterns of seed dispersal and pollen flow in Quercus garryana (Fagaceae) following post-glacial climatic changes

Aim We examined the genetic structure of Quercus garryana to infer post‐glacial patterns of seed dispersal and pollen flow to test the hypotheses that (1) peripheral populations are genetically distinct from core populations and from one another; (2) genetic diversity declines towards the poleward edge of the species’ range; and (3) genetic diversity in the chloroplast genome, a direct measure of seed dispersal patterns, declines more sharply with increasing latitude than diversity in the nuclear genome. We address our findings in the context of known historical oak distribution from pollen core data derived from previously published research. Location Oak–savanna ecosystems from southern Oregon, USA (core populations/non‐glaciated range) northward to Vancouver Island, British Columbia, Canada (peripheral populations/glaciated range). Methods We genotyped 378 trees from 22 sites with five chloroplast and seven nuclear microsatellite loci. For both sets of markers, we estimated genetic diversity and differentiation using an analysis of molecular variance and generated Mantel correlograms to detect genetic and geographical distance correlations. For the nuclear markers, we also used a Bayesian approach to infer population substructure. Results There was a large degree of population differentiation revealed by six chloroplast haplotypes, with little (≤ 3) or no haplotype diversity within sites. Peripheral island locations shared the same, maternally inherited chloroplast haplotype, whereas locations in mainland Washington had greater haplotype diversity. In contrast, genetic diversity of the nuclear markers was high at all locations sampled. Populations clustered into two groups and were significantly positively correlated over large spatial scales (≤ 200 km), although allele richness decreased significantly with latitude. Population substructure was observed between core and peripheral populations because rare alleles were absent in peripheral localities and common allele frequencies differed. Main conclusions The observed pattern of chloroplast haplotype loss at the northern periphery suggests restricted seed dispersal events from mainland sites to peripheral islands. This pattern was unexpected, however, as refugial oak populations remained near the current post‐glacial range even during the Last Glacial Maximum. Using nuclear markers, we found high within‐population diversity and population differentiation only over large spatial scales, suggesting that pollen flow is relatively high among populations.     

Population structure and the impact of regional and local habitat isolation upon levels of genetic diversity of the endangered damselfly Coenagrion mercuriale (Odonata: Zygoptera)

1. Coenagrion mercuriale is one of Europe's most threatened damselflies. There is concern for the long‐term persistence of many of its U.K. colonies because adult lifetime movement is limited, making isolated populations susceptible to extinction. 2. Using 14 microsatellite loci we characterised levels of genetic diversity, evidence for a recent decline and the spatial genetic structure for C. mercuriale population in Wales, U.K. 3. Spatial isolation is not an absolute predictor of low genetic diversity at either local or regional scales. 4. One population inhabiting a remote, edge of range site is genetically impoverished with levels of variability (at microsatellite loci) among the lowest reported for any insect species. 5. Agricultural land and high ground are physical barriers to dispersal by adults. 6. Consistent with work from elsewhere, movement by mature C. mercuriale in Pembrokeshire is sufficient to prevent significant genetic differentiation throughout a habitat matrix of some 3–4 km if the suitable habitat sites are <2 km apart and lack barriers to movement. Even within a good habitat matrix, however, genetic isolation by distance develops within 10 km.