FACTORS INFLUENCING THE REPRODUCTIVE SUCCESS OF IPOMOEA PES-CAPRAE (CONVOLVULACEAE) AROUND THE GULF OF MEXICO

The reproductive biology of Ipomoea pes-caprae, a pantropical beach morning glory, was studied at five sites around the Gulf of Mexico. The primary pollinators were Xylocopa species (carpenter bees) which dispersed pollen up to 90 m. Exclusion experiments demonstrated that ants feeding on extrafloral nectaries increased seed set but did not protect seeds from predation by the bruchid beetle Megacerus. The water-dispersed seed and long-range dispersal of pollen may function to counter the sporophytic incompatability mechanism of the plant; populations exhibit a large neighborhood size. Key reproductive factors in the life cycle of I. pes-caprae are the long-range pollen flow and mass germination of water-dispersed seeds.     

Gene flow and population subdivision in a pantropical plant with sea-drifted seeds Hibiscus tiliaceus and its allied species: evidence from microsatellite analyses

The genetic differentiation and structure of Hibiscus tiliaceus , a pantropical plant with sea-drifted seeds, and four allied species were studied using six microsatellite markers. A low level of genetic differentiation was observed among H. tiliaceus populations in the Pacific and Indian Ocean regions, similar to the results of a previous chloroplast DNA (cpDNA) study. Frequent gene flow by long-distance seed dispersal is responsible for species integration of H. tiliaceus in the wide distribution range. On the other hand, highly differentiated populations of H. tiliaceus were detected in West Africa, as well as of Hibiscus pernambucensis in southern Brazil. In the former populations, the African continent may be a geographical barrier that prevents gene flow by sea-drifted seeds. In the latter populations, although there are no known land barriers, the bifurcating South Equatorial Current at the north-eastern horn of Brazil can be a potential barrier to gene flow and may promote the genetic differentiation of these populations. Our results also suggest clear species segregation between H. tiliaceus and H. pernambucensis , which confirms the introgression scenario between these two species that was suggested by a previous cpDNA study. Our results also provide good evidence for recent transatlantic long-distance seed dispersal by sea current. Despite the distinct geographical structure observed in the cpDNA haplotypes, a low level of genetic differentiation was found between Pacific and Atlantic populations of H. pernambucensis , which could be caused by transisthmian gene flow.     

Pollen flow between flowers of the same morph in invasive populations of Oxalis pes-caprae L. in the western Mediterranean region

Oxalis pes-caprae is a tristylous South African geophyte that is invasive in regions with Mediterranean climate. Given the introduction of the short-styled morph only, vegetative reproduction was, until recently, the reproductive mode known for this invaded area. The detection of new floral morphs, fruit and seed production in natural populations of the invaded range, together with the reported weakening in the heteromorphic incompatibility system, raised the question on whether the short-styled flowers are able to exchange pollen among themselves, and if so, which mechanisms account for it. Thus, flower morphometric analysis and field experiments were conducted to assess pollinator visitation rates and pollen flow in three invasive populations. Flower morphological changes suggesting the evolution from tristyly towards semi-homostyly were observed. Moreover, O. pes-caprae flowers were visited by different functional groups of pollinators, mostly hymenopteran. Dye flow experiments revealed that the short-styled flowers were able to successfully exchange pollen among themselves in natural conditions. All of this, together with the weakening of the incompatibility system, constitutes further data that might help to explain the recently reported occurrence of sexual reproduction in this invaded range, which by itself bears important implications for the invasion process of O. pes-caprae.     

Genetic diversity and dispersal of Phragmites australis in a small river system

Even though the reed, Phragmites australis, is an extensively studied wetland species, little is known about reproduction and dispersal modes within and among reed populations at the scale of small river systems. Using microsatellite analysis of 189 individuals from three adjacent river catchments in the Czech Republic, we elucidated the role of the river corridors in the dispersal of P. australis. Using Bayesian clustering of individuals, we found that 19% of clusters were distributed only along one river, which implied dispersal by water (or by wind) along river corridors, whereas 38% of clusters were widely distributed and were likely the product of wind long-distance dispersal among rivers. Intensive exchange of propagules among river systems is further demonstrated by only 6% of total variance being attributed to the variance among rivers in the AMOVA-analysis. Spatial autocorrelation analysis revealed a decreasing pattern up to 5–10 km and no clear pattern over longer distances. This gives an evidence for pollen and seed dispersal at short distances (up to 1 km), whereas most likely only seed dispersal at longer distances up to 10 km. We found five multilocus genotypes distributed in two different populations. The distances between populations with the same genotype ranged from 0.5 to 10.8 km. This can be interpreted as long-distance vegetative dispersal.     

Post-glacial history of Trillium grandiflorum (Melanthiaceae) in eastern North America: inferences from phylogeography

Dispersal and migration are important processes affecting the evolutionary history and genetics of species. Here we investigate post-glacial migration and gene flow in Trillium grandiflorum (Melanthiaceae), a wide-ranging, forest herb from eastern North America. Using phylogeographic approaches, we examined cpDNA and allozyme diversity in 35 populations of T. grandiflorum sampled from throughout the geographic range of the species. Nested clade analysis (NCA) of cpDNA haplotypes indicated that T. grandiflorum likely survived in two refugia in the southeastern US during the last glaciation and that long-distance dispersal characterized the post-glacial recolonization of northern areas. There was no evidence for reduced allozyme diversity in populations from glaciated compared to ice-free regions, probably because of the greater abundance and larger effective size of populations in the north. An analysis of isolation-by-distance based on the allozyme data suggested a pattern of population differentiation consistent with restricted gene flow. Notwithstanding the significance of rare seed dispersal events for migration, a comparison of allozyme and cpDNA genetic structure indicates that pollen flow between populations is more likely than seed dispersal. These results for T. grandiflorum represent the first phylogeographic analysis of a temperate woodland herb in eastern North America and support the importance of occasional long-distance dispersal events in the post-glacial migration of plants.     

Effective Long-Distance Pollen Dispersal in Centaurea jacea

Background

Agri-environment schemes play an increasingly important role for the conservation of rare plants in intensively managed agricultural landscapes. However, little is known about their effects on gene flow via pollen dispersal between populations of these species.

Methodology/Principal Findings

In a 2-year experiment, we observed effective pollen dispersal from source populations of Centaurea jacea in restored meadows, the most widespread Swiss agri-environment scheme, to potted plants in adjacent intensively managed meadows without other individuals of this species. Potted plants were put in replicated source populations at 25, 50, 100 m and where possible 200 m distance from these source populations. Pollen transfer among isolated plants was prevented by temporary bagging, such that only one isolated plant was accessible for flower visitors at any one time. Because C. jacea is self-incompatible, seed set in single-plant isolates indicated insect mediated effective pollen dispersal from the source population. Seed set was higher in source populations (35.7±4.4) than in isolates (4.8±1.0). Seed set declined from 18.9% of that in source populations at a distance of 25 m to 7.4% at 200 m. At a distance of 200 m seed set was still significantly higher in selfed plants, indicating long-distance effective pollen dispersal up to 200 m. Analyses of covariance suggested that bees contributed more than flies to this long-distance pollen dispersal. We found evidence that pollen dispersal to single-plant isolates was positively affected by the diversity and flower abundance of neighboring plant species in the intensively managed meadow. Furthermore, the decline of the dispersal was less steep when the source population of C. jacea was large.

Conclusions

We conclude that insect pollinators can effectively transfer pollen from source populations of C. jacea over at least 200 m, even when “recipient populations” consisted of single-plant isolates, suggesting that gene flow by pollen over this distance is very likely. Source population size and flowering environment surrounding recipient plants appear to be important factors affecting pollen dispersal in C. jacea. It is conceivable that most insect-pollinated plants in a network of restored sites within intensively managed grassland can form metapopulations, if distances between sites are of similar magnitude as tested here.     

Pollen dispersal and breeding structure in a hawkmoth-pollinated Pampa grasslands species Petunia axillaris (Solanaceae)

Background and Aims The evolution of selfing is one of the most common transitions in flowering plants, and this change in mating pattern has important systematic and ecological consequences because it often initiates reproductive isolation and speciation. Petunia axillaris (Solanaceae) includes three allopatric subspecies widely distributed in temperate South America that present different degrees of self-compatibity and incompatibility. One of these subspecies is co-distributed with P. exserta in a restricted area and presents a complex, not well-understood mating system. Artificial crossing experiments suggest a complex system of mating in this sympatric area. The main aims of this study were to estimate the pollen dispersal distance and to evaluate the breeding structure of P. axillaris subsp. axillaris, a hawkmoth-pollinated taxon from this sympatric zone.Methods Pollen dispersal distance was compared with nearest-neighbours distance, and the differentiation in the pollen pool among mother plants was estimated. In addition, the correlation between genetic differentiation and spatial distance among plants was tested. All adult individuals (252) within a space of 2800 m² and 15 open-pollinated progeny (285 seedlings) were analysed. Genetic analyses were based on 12 polymorphic microsatellite loci.Key Results A high proportion of self-pollination was found, indicating a mixed-mating system. The maximum pollen dispersal distance was 1013 m, but most pollination events (96 %) occurred at a distance of 0 m, predominantly in an inbreeding system. Both parents among sampled individuals could be identifed in 60–85 % of the progeny.Conclusions The results show that most pollen dispersal in the hawkmoth-pollinated P. axillaris subsp. axillaris occurs within populations and there is a high proportion of inbreeding. This mating system appears to favour species integrity in a secondary contact zone with the congener species P. exserta.     

Limited pollen-mediated dispersal and partial self-incompatibility in the rare ironstone endemic <Emphasis Type="Italic">Tetratheca paynterae</Emphasis> subsp. <Emphasis Type="Italic">paynterae</Emphasis> increase the risks associated with habitat loss

Patterns of mating and dispersal are key factors affecting the dynamics, viability and evolution of plant populations. Changes in mating system parameters can provide evidence of anthropogenic impacts on populations of rare plants. Tetratheca paynterae subsp. paynterae is a critically endangered perennial shrub confined to a single ironstone range in Western Australia. Mining of the range removed 25% of plants in 2004 and further plants may be removed if the viability of the remaining populations is not compromised. To provide baseline genetic data for monitoring mining impacts, we characterised the mating system and pollen dispersal over two seasons in T. paynterae subsp. paynterae and compared mating system parameters with two other ironstone endemics, T. paynterae subsp. cremnobata and T. aphylla subsp. aphylla that were not impacted by mining. T. paynterae subsp. paynterae was the only taxon showing evidence of inbreeding (t _m = 0.89), although hand pollination revealed pre-zygotic self-incompatibility limits the production of seed from self-pollen. In a year of lower fruit set (2005), the estimate of correlated paternity increased from 20 to 35%. Direct estimates of realised pollen dispersal, made by paternity assignment in two small populations where all adult plants were genotyped, revealed a leptokurtic distribution with 30% of pollen dispersed less than 3 m and 90% less than 15 m. Restricted pollen dispersal maintains the strong genetic structuring of the adult populations in succeeding generations. As a consequence of preferential outcrossing, any reduction in effective population size, flowering plant density and/or the abundance and activity of pollinators may impact negatively on population viability through reduced seed set, increased inbreeding and increased correlated paternity.     

Fine-scale spatial genetic structure and gene dispersal in Silene latifolia

Plants are sessile organisms, often characterized by limited dispersal. Seeds and pollen are the critical stages for gene flow. Here we investigate spatial genetic structure, gene dispersal and the relative contribution of pollen vs seed in the movement of genes in a stable metapopulation of the white campion Silene latifolia within its native range. This short-lived perennial plant is dioecious, has gravity-dispersed seeds and moth-mediated pollination. Direct measures of pollen dispersal suggested that large populations receive more pollen than small isolated populations and that most gene flow occurs within tens of meters. However, these studies were performed in the newly colonized range (North America) where the specialist pollinator is absent. In the native range (Europe), gene dispersal could fall on a different spatial scale. We genotyped 258 individuals from large and small (15) subpopulations along a 60 km, elongated metapopulation in Europe using six highly variable microsatellite markers, two X-linked and four autosomal. We found substantial genetic differentiation among subpopulations (global F_ST=0.11) and a general pattern of isolation by distance over the whole sampled area. Spatial autocorrelation revealed high relatedness among neighboring individuals over hundreds of meters. Estimates of gene dispersal revealed gene flow at the scale of tens of meters (5–30 m), similar to the newly colonized range. Contrary to expectations, estimates of dispersal based on X and autosomal markers showed very similar ranges, suggesting similar levels of pollen and seed dispersal. This may be explained by stochastic events of extensive seed dispersal in this area and limited pollen dispersal.     

Limited structure and widespread diversity suggest potential buffers to genetic erosion in a threatened grassland shrub Pimelea spinescens (Thymelaeaceae)

Pimelea spinescens is a critically endangered species of the temperate grasslands of southeastern Australia. Two subspecies are recognised. Subspecies, P. spinescens subsp. spinescens, formerly common and widespread, is found in isolated remnants of previously extensive grasslands. The second subspecies, subsp. pubiflora, is thought to have been historically rare with only two geographically-isolated extant populations. The grassland communities exist now as fragmented remnants representing <1 % of their extent prior to European settlement in the early 1800s. Conservation management strategies for species in these critically endangered ecosystems rely on an understanding of genetic diversity and population structure to ensure long-term evolutionary potential. We used chloroplast DNA (cpDNA) and microsatellite markers to examine the population genetic structure of both subspecies. Analysis of cpDNA revealed 14 haplotypes with high divergence between the single haplotype found in subsp. pubiflora and most remaining haplotypes restricted to subsp. spinescens. Microsatellites also indicated high genetic differentiation between subspecies but little evidence of sub-structuring within either subspecies. Results suggest that seed dispersal has not been as limited as previously thought. In this fragmented habitat, a lack of genetic structure suggests buffers to genetic erosion, with current patterns reflecting genetic diversity prior to fragmentation. Plant longevity and the presence of seed banks may contribute to the maintenance of these patterns, resulting in a lag between fragmentation and genetic erosion. Whilst factors such as longevity and seed banks may be preserving historic genetic diversity, management is required to ensure the maintenance of this diversity into the future.     

Genetic diversity and spatial genetic structure of Pinus strobus (Pinaceae) across an island landscape inferred from allozyme and cpDNA markers

The Beaver Island Archipelago (BIA) provides a model system to address the impact of long-term isolation on genetic diversity and gene flow. Low lake levels are assumed to have caused the BIA to be attached to mainland Michigan for at least 4000 years (10000 yr B.P.- 6000 yr B.P.), eventually, rising lake levels would have kept the islands isolated since 6000 yr B.P. If the island populations of a plant species in the BIA were indeed once continuous with the mainland of Michigan, then we would expect similar levels of genetic diversity in populations of such a species on the islands vs. the mainland. We compared levels of allozyme genetic diversity of 20 plots of Pinus strobus in the BIA with two mainland populations in northern Michigan. In addition, if pollen is a primary agent of gene flow across islands, a low degree of allozyme differentiation among the island populations of P. strobus in the BIA would be evident. Furthermore, since seed dispersal is more limited than pollen dispersal in P. strobus, a more pronounced spatial genetic structure (SGS) is expected in allozymes than in cpDNA markers. To gain insights on the pattern of seed and pollen dispersal among the 20 plots, we further analyzed spatial autocorrelation using Moran's I-statistics for both data sets [biparentally inherited, allozymes and paternally inherited, cpDNA microsatellites (cpDNA SSR)]. We found a similar level of allozyme variability in both the BIA (mean H _e = 0.080) and the two mainland populations (mean H _e = 0.078). As predicted, we observed a low but significant degree of genetic divergence among populations for allozymes (mean F _ST = 0.033 across 20 plots). Our allozyme-based SGS analysis revealed significant evidence of SGS (i.e. isolation-by-distance; slope β = ?0.194 from regression analysis of observed averaged Moran's I values against the logarithm of the upper bound of six distance classes). In contrast, little evidence of SGS was found in cpDNA SSR data across the BIA (β = 0.013). These results suggest that although gene flow via seed dispersal is somewhat limited, pollen flow has been sufficient to maintain genetic diversity and prevent differentiation across the island landscape over several thousand years of isolation.     

Morphological differentiation supports the genetic pattern of the geographic structure of Juniperus thurifera (Cupressaceae)

Juniperus thurifera is an important component of woodland communities of dry sites within the West Mediterranean region and is characterised by a strongly disjunctive geographic range. Two subspecies were recognised, subsp. thurifera in Europe and subsp. africana in Africa. The aim of the study was the comparison of phenetic diversity to the pattern of AFLP geographic differentiation of the species described in the literature. The examination of phenetic diversity was based on the biometrical analysis of 17 populations using 12 morphological characters of cone and seed. The differences among populations were analysed using Student’s t test, analysis of discrimination, UPGMA agglomeration and hierarchical analysis of variance. The majority of morphological characters differentiated at a statistically significant level between populations and between J. thurifera subsp. thurifera and subsp. africana. Three groups of populations were detected using multivariate statistical analyses. The first, well separated, is subsp. africana, while the following two concern subsp. thurifera. The morphological differentiation of populations appeared similar to that described on the AFLP. The Gibraltar Straight appeared to be the most important barrier.     

Extensive long-distance pollen dispersal and highly outcrossed mating in historically small and disjunct populations of Acacia woodmaniorum (Fabaceae), a rare banded iron formation endemic

Background and Aims

Understanding patterns of pollen dispersal and variation in mating systems provides insights into the evolutionary potential of plant species and how historically rare species with small disjunct populations persist over long time frames. This study aims to quantify the role of pollen dispersal and the mating system in maintaining contemporary levels of connectivity and facilitating persistence of small populations of the historically rare Acacia woodmaniorum.

Methods

Progeny arrays of A. woodmaniorum were genotyped with nine polymorphic microsatellite markers. A low number of fathers contributed to seed within single pods; therefore, sampling to remove bias of correlated paternity was implemented for further analysis. Pollen immigration and mating system parameters were then assessed in eight populations of varying size and degree of isolation.

Key Results

Pollen immigration into small disjunct populations was extensive (mean minimum estimate 40 % and mean maximum estimate 57 % of progeny) and dispersal occurred over large distances (≤1870m). Pollen immigration resulted in large effective population sizes and was sufficient to ensure adaptive and inbreeding connectivity in small disjunct populations. High outcrossing (mean t_m = 0·975) and a lack of apparent inbreeding suggested that a self-incompatibility mechanism is operating. Population parameters, including size and degree of geographic disjunction, were not useful predictors of pollen dispersal or components of the mating system.

Conclusions

Extensive long-distance pollen dispersal and a highly outcrossed mating system are likely to play a key role in maintaining genetic diversity and limiting negative genetic effects of inbreeding and drift in small disjunct populations of A. woodmaniorum. It is proposed that maintenance of genetic connectivity through habitat and pollinator conservation will be a key factor in the persistence of this and other historically rare species with similar extensive long-distance pollen dispersal and highly outcrossed mating systems.     

Pollen Dispersal in Fragmented Populations of the Dioecious Wind-Pollinated Tree,Allocasuarina verticillata (Drooping Sheoak,Drooping She-Oak; Allocasuarinaceae)

Vegetation clearing, land modification and agricultural intensification have impacted on many ecological communities around the world. Understanding how species respond to fragmentation and the scales over which functionality is retained, can be critical for managing biodiversity in agricultural landscapes. Allocasuarina verticillata (drooping sheoak, drooping she-oak) is a dioecious, wind-pollinated and -dispersed species with key conservation values across southeastern Australia. But vegetation clearing associated with agricultural expansion has reduced the abundance and spatial distribution of this species in many regions. Spatial genetic structure, relatedness among trees, pollen dispersal and mating patterns were examined in fragmented A. verticillata populations selected to represent the types of remnants that now characterise this species. Short scale spatial genetic structure (5–25 m) and relatedness among trees were observed in most populations. Unexpectedly, the two male trees closest to each female did not have a reproductive advantage accounting for only 4–15% of the seed produced in larger populations. Biparental inbreeding was also generally low (<4%) with limited evidence of seed crop domination by some male trees. More male trees contributed to seed crops in linear remnants (mean 17) compared to those from patch remnants (mean 11.3) which may reflect differences in pollen dispersal within the two remnant types. On average, pollen travels ~100 m irrespective of remnant type but was also detected to have dispersed as far as 1 km in open landscapes. Low biparental inbreeding, limited reproductive assurance for near-neighbour and probably related males and variability in the distances over which females sample pollen pools suggest that some mechanism to prevent matings between relatives exists in this species.     

Inferring demographic processes from the genetic structure of a metapopulation of <Emphasis Type="Italic">Boltonia decurrens</Emphasis>(Asteraceae)

Boltonia decurrens(Asteraceae), a federally listed, threatened floodplain species, requires regular flooding for suitable habitat and seed dispersal. Flood suppression and habitat destruction have resulted in fewer than 25 populations remaining throughout its 400 km range. Because individual populations are widely interspaced (>10 km) and subject to frequent extinction and colonization, seed dispersal along the river, not pollen flow, is likely the primary determinant of population genetic structure. We used neutral genetic markers (isozymes) assayed for fourteen populations to determine which demographic processes contribute to the genetic structure of B. decurrens. Significant genetic differentiation was detected among populations (F _ST=0.098, P< 0.05) but not among regions (F _RT=0.013, P> 0.05), suggesting that long-distance dispersal events occur and involve seed from a small number of populations. Correspondingly, we found no evidence of isolation by distance, and admixture analyses indicate that colonization events involve seed from 3 to 5 source populations. Individual populations exhibited high levels of fixation (mean F _IS=0.192, P< 0.05), yet mean population outcrossing rates were high (t _m=0.87–0.95) and spatial autocorrelation analyses revealed no fine-scale within population structure, indicating that inbreeding alone cannot explain the observed fixation. Rather, genetic bottlenecks, detected for 12 of 14 populations, and admixture at population founding may be important sources of fixation. These observations are consistent with a metapopulation model and confirm the importance of regular flooding events, capable of producing suitable habitat and dispersing seed long distances, to the long-term persistence of B. decurrens.     

Diverse spore rains and limited local exchange shape fern genetic diversity in a recently created habitat colonized by long-distance dispersal

Background and Aims

Populations established by long-distance colonization are expected to show low levels of genetic variation per population, but strong genetic differentiation among populations. Whether isolated populations indeed show this genetic signature of isolation depends on the amount and diversity of diaspores arriving by long-distance dispersal, and time since colonization. For ferns, however, reliable estimates of long-distance dispersal rates remain largely unknown, and previous studies on fern population genetics often sampled older or non-isolated populations. Young populations in recent, disjunct habitats form a useful study system to improve our understanding of the genetic impact of long-distance dispersal.

Methods

Microsatellite markers were used to analyse the amount and distribution of genetic diversity in young populations of four widespread calcicole ferns (Asplenium scolopendrium, diploid; Asplenium trichomanes subsp. quadrivalens, tetraploid; Polystichum setiferum, diploid; and Polystichum aculeatum, tetraploid), which are rare in The Netherlands but established multiple populations in a forest (the Kuinderbos) on recently reclaimed Dutch polder land following long-distance dispersal. Reference samples from populations throughout Europe were used to assess how much of the existing variation was already present in the Kuinderbos.

Key Results

A large part of the Dutch and European genetic diversity in all four species was already found in the Kuinderbos. This diversity was strongly partitioned among populations. Most populations showed low genetic variation and high inbreeding coefficients, and were assigned to single, unique gene pools in cluster analyses. Evidence for interpopulational gene flow was low, except for the most abundant species.

Conclusions

The results show that all four species, diploids as well as polyploids, were capable of frequent long-distance colonization via single-spore establishment. This indicates that even isolated habitats receive dense and diverse spore rains, including genotypes capable of self-fertilization. Limited gene flow may conserve the genetic signature of multiple long-distance colonization events for several decades.     

Origin,Migration Routes and Worldwide Population Genetic Structure of the Wheat Yellow Rust Pathogen Puccinia striiformis f.sp. tritici

Analyses of large-scale population structure of pathogens enable the identification of migration patterns, diversity reservoirs or longevity of populations, the understanding of current evolutionary trajectories and the anticipation of future ones. This is particularly important for long-distance migrating fungal pathogens such as Puccinia striiformis f.sp. tritici (PST), capable of rapid spread to new regions and crop varieties. Although a range of recent PST invasions at continental scales are well documented, the worldwide population structure and the center of origin of the pathogen were still unknown. In this study, we used multilocus microsatellite genotyping to infer worldwide population structure of PST and the origin of new invasions based on 409 isolates representative of distribution of the fungus on six continents. Bayesian and multivariate clustering methods partitioned the set of multilocus genotypes into six distinct genetic groups associated with their geographical origin. Analyses of linkage disequilibrium and genotypic diversity indicated a strong regional heterogeneity in levels of recombination, with clear signatures of recombination in the Himalayan (Nepal and Pakistan) and near-Himalayan regions (China) and a predominant clonal population structure in other regions. The higher genotypic diversity, recombinant population structure and high sexual reproduction ability in the Himalayan and neighboring regions suggests this area as the putative center of origin of PST. We used clustering methods and approximate Bayesian computation (ABC) to compare different competing scenarios describing ancestral relationship among ancestral populations and more recently founded populations. Our analyses confirmed the Middle East-East Africa as the most likely source of newly spreading, high-temperature-adapted strains; Europe as the source of South American, North American and Australian populations; and Mediterranean-Central Asian populations as the origin of South African populations. Although most geographic populations are not markedly affected by recent dispersal events, this study emphasizes the influence of human activities on recent long-distance spread of the pathogen.     

Serial founder effects and genetic differentiation during worldwide range expansion of monarch butterflies

Range expansions can result in founder effects, increasing genetic differentiation between expanding populations and reducing genetic diversity along the expansion front. However, few studies have addressed these effects in long-distance migratory species, for which high dispersal ability might counter the effects of genetic drift. Monarchs (Danaus plexippus) are best known for undertaking a long-distance annual migration in North America, but have also dispersed around the world to form populations that do not migrate or travel only short distances. Here, we used microsatellite markers to assess genetic differentiation among 18 monarch populations and to determine worldwide colonization routes. Our results indicate that North American monarch populations connected by land show limited differentiation, probably because of the monarch''s ability to migrate long distances. Conversely, we found high genetic differentiation between populations separated by large bodies of water. Moreover, we show evidence for serial founder effects across the Pacific, suggesting stepwise dispersal from a North American origin. These findings demonstrate that genetic drift played a major role in shaping allele frequencies and created genetic differentiation among newly formed populations. Thus, range expansion can give rise to genetic differentiation and declines in genetic diversity, even in highly mobile species.     

Effect of gene flow on spatial genetic structure in the riparian canopy tree Cercidiphyllum japonicum revealed by microsatellite analysis

Few studies have analyzed pollen and seed movements at local scale, and genetic differentiation among populations covering the geographic distribution range of a species. We carried out such a study on Cercidiphyllum japonicum; a dioecious broad-leaved tree of cool-temperate riparian forest in Japan. We made direct measurement of pollen and seed movements in a site, genetic structure at the local scale, and genetic differentiation between populations covering the Japanese Archipelago. Parentage analysis of seedlings within a 20-ha study site indicated that at least 28.8% of seedlings were fertilized by pollen from trees outside the study site. The average pollination distance within the study site was 129 m, with a maximum of 666 m. The genotypes of 30% of seedlings were incompatible with those of the nearest female tree, and the maximum seed dispersal distance within the study site was over 300 m. Thus, long-distance gene dispersal is common in this species. The correlation between genetic relatedness and spatial distance among adult trees within the population was not significant, indicating an absence of fine-scale genetic structure perhaps caused by high levels of pollen flow and overlapping seed shadows. Six populations sampled throughout the distribution of C. japonicum in Japan showed significant isolation-by-distance but low levels of genetic differentiation (F(ST) = 0.043), also indicating long-distance gene flow in C. japonicum. Long-distance gene flow had a strong influence on the genetic structure at different spatial scales, and contributes to the maintenance of genetic diversity in C. japonicum.     

Slatkin’s Paradox: when direct observation and realized gene flow disagree. A case study in Ficus

In flying insects, there is frequently a lack of congruence between empirical estimates of local demographic parameters and the prediction that differentiation between populations should decrease with increasing dispersal, a puzzling phenomenon known as Slatkin’s Paradox. Here, we generalize Slatkin’s Paradox to other taxa, drawing from available information on dispersal to predict the relative importance of pollen vs. seed migration in structuring broad‐scale patterns of genetic variation in Ficus hirta, a dioecious fig whose pollen is dispersed by minute, species‐specific fig wasps and whose seeds are disseminated by a variety of vagile vertebrates (especially bats and birds). Local‐scale observational and genetic studies of dioecious understory figs suggest comparable rates of pollen and seed migration. In contrast, we found unusually low nuclear differentiation (F_ST = 0.037, R_ST = 0.074) and high chloroplast differentiation (G_ST = 0.729, N_ST = 0.798) among populations separated by up to 2850 km, leading us to reject the hypothesis of equal pollen and seed migration rates and to obtain an equilibrium estimate of the ratio of pollen to seed migration of r = 16.2–36.3. We reconcile this example of Slatkin’s Paradox with previously published data for dioecious figs and relate it to the picture of exceptionally long‐distance wasp‐mediated pollen dispersal that is emerging for large monoecious fig trees. More generally, we argue that Slatkin’s Paradox is a general phenomenon and suggest it may be common in plants and animals.