Phylogeography and population differentiation in terrestrial island populations of Banksia arborea (Proteaceae)

The Banded Iron Formations (BIFs) of south‐western Australia are terrestrial islands characterized by high species richness and endemism. Regional endemics occur across multiple formations without inhabiting the intervening landscape matrix. We investigated whether the occurrence on BIF terrestrial islands has led to genetic differentiation among the eight known populations of the regional endemic, Banksia arborea. Genetic structure was assessed using three chloroplast DNA sequence markers and 11 nuclear microsatellite loci. Phylogenetic relationships were assessed with statistical parsimony and Bayesian methods. Dates of haplotype divergence were estimated using the time to most recent common ancestor of B. arborea and Banksia purdieana, as well as a conservative angiosperm chloroplast (cp)DNA mutation rate. Population genetic diversity and structure was assessed amongst and within populations by genotyping 24 geographically clustered individuals from each BIF and three subpopulations within the Die Hardy Range BIF. Indirect gene flow estimates were determined using a method based on the frequency of private alleles. Banksia arborea showed low genetic diversity in (cp)DNA and a complex structural pattern, with genetic differentiation of some BIF populations and an absence of differentiation amongst others, reflecting either retention of ancestral polymorphism across northern BIF populations or more recent connectivity of these populations. There was little evidence of pollen dispersal both between BIFs and within the large BIF known as Die Hardy Range. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 860–872.     

Does provenance matter? Fitness is not determined by genetic distance or the scale of pollen dispersal in Grevillea sphacelata (Proteaceae)

Understanding how the scale of pollen transfer determines the outcome of matings is important evolutionarily and a key issue in restoration ecology. We tested the effects of pollen transfer distance for the self‐incompatible shrub Grevillea sphacelata using (1) open pollination and transfer among (2) near neighbours, (3) neighbouring subpopulations and (4) populations separated by c. 4 km. We used AFLP markers to test for evidence of genetic differentiation within and among populations. Patterns of seed initiation suggest that open pollinated flowers were pollen limited, although in one subpopulation open seed set was greater than that achieved with pollen from near neighbours or other subpopulations. We detected no other effects of pollen source on seed initiation or seed and seedling development. In contrast, our genetic survey revealed significant spatial autocorrelation to 5 m, moderate differentiation of populations separated by up to 4 km and significant isolation by distance > 16 km. Our data suggest that, although dispersal of pollen may typically be localized, gene flow prevents localized adaptation or co‐adaptation and we detected no effects of inbreeding depression. In a restoration context, our results imply that movement of seed between populations separated by 4 km will not have detrimental consequences, despite significant differentiation at neutral genetic markers, and may be beneficial in maintaining genetic diversity and evolutionary potential. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 290–302.     

A phylogeographic study of the stoneplant Conophytum (Aizoaceae; Ruschioideae; Ruschieae) in the Bushmanland Inselberg Region (South Africa) suggests anemochory

The Bushmanland Inselberg Region (BIR) of South Africa provides an ideal system to study population interactions, as these inselbergs function as islands of Succulent Karoo surrounded by Nama Karoo vegetation. The population genetics of four Conophytum taxa endemic to the quartz-associated habitats of inselbergs in the BIR were investigated using amplified fragment length polymorphisms (AFLP), namely C. marginatum subsp. haramoepense, C. marginatum subsp. marginatum, C. maughanii, and C. ratum. Conophytum marginatum colonizes the quartz outcrops on the summits of the inselbergs, while C. maughanii and C. ratum occupy quartz patches at the summit and base of the inselbergs. A total of 24 populations were sampled to assess genetic differentiation between populations of each species, specifically between summit and base populations of C. ratum, eastern and western populations of C. maughanii and populations of the two subspecies of C. marginatum. Moderate levels of genetic differentiation were recovered between the summit and base populations of C. ratum, with an indication of some genetic connectivity between the populations. Slight differentiation between the eastern and western populations of C. maughanii was recovered, however, this was not reflected in the PCoA and STRUCTURE results. In C. marginatum, no significant genetic differentiation was recovered between populations of the subspecies. These results may reflect evidence of different dispersal mechanisms in the species, with the genetic connectivity between populations of C. ratum possibly indicating dispersal through hygrochastic capsules, while genetic connectivity between populations of C. maughanii and C. marginatum may, for the first time, suggest long-distance dispersal, i.e. anemochory. This study provides the first insights into population interactions across the BIR and highlights the importance of conservation in the region, particularly of the Gamsberg, in light of the recent mining activities.     

Chloroplast population genetics reveals low levels of genetic variation and conformation to the central–marginal hypothesis in Taxus wallichiana var. mairei,an endangered conifer endemic to China

The central–marginal hypothesis predicts that geographically peripheral populations should exhibit reduced genetic diversity and increased genetic differentiation than central populations due to smaller effective population size and stronger geographical isolation. We evaluated these predictions in the endangered conifer Taxus wallichiana var. mairei. Eight plastid simple sequence repeats (cpSSRs) were used to investigate plastid genetic variation in 22 populations of Taxus wallichiana var. mairei, encompassing nearly its entire distribution range. Low levels of plastid genetic variation and differentiation were detected in the populations, and the findings were attributed to low mutation rates, small population sizes, habitat fragmentation and isolation, and effective pollen or seed dispersal. Hunan and Hubei were identified as major refugia based on the number of private haplotypes and species distribution modeling. Trends in plastid genetic diversity and genetic differentiation from central to peripheral populations supported the predictions of the central–marginal hypothesis. In scenarios wherein the future climate becomes warmer, we predict that some peripheral populations will disappear and southern and southeastern regions will become significantly less habitable. Factors that include the levels of precipitation during the driest month, annual precipitation level, and annual temperature range will be decisive in shaping the future distribution of these populations. This study provides a theoretical basis for the conservation of T. wallichiana var. mairei.     

Contrasting patterns of pollen and seed flow influence the spatial genetic structure of sweet vernal grass (Anthoxanthum odoratum) populations

The spatial genetic structure of plant populations is determined by a combination of gene flow, genetic drift, and natural selection. Gene flow in most plants can result from either seed or pollen dispersal, but detailed investigations of pollen and seed flow among populations that have diverged following local adaptation are lacking. In this study, we compared pollen and seed flow among 10 populations of sweet vernal grass (Anthoxanthum odoratum) on the Park Grass Experiment. Overall, estimates of genetic differentiation that were based on chloroplast DNA (cpDNA) and, which therefore resulted primarily from seed flow, were lower (average F(ST) = 0.058) than previously published estimates that were based on nuclear DNA (average F(ST) = 0.095). Unlike nuclear DNA, cpDNA showed no pattern of isolation by adaptation; cpDNA differentiation was, however, inversely correlated with the number of additions (nutrients and lime) that each plot had received. We suggest that natural selection is restricting pollen flow among plots, whereas nutrient additions are increasing seed flow and genetic diversity by facilitating the successful germination and growth of immigrant seeds. This study highlights the importance of considering all potential gene flow mechanisms when investigating determinants of spatial genetic structure, and cautions against the widespread assumption that pollen flow is more important than seed flow for population connectivity in wind-pollinated species.     

PHYLOGEOGRAPHIC STRUCTURE AND OUTBREEDING DEPRESSION REVEAL EARLY STAGES OF REPRODUCTIVE ISOLATION IN THE NEOTROPICAL ORCHID EPIDENDRUM DENTICULATUM

Phylogeographic studies provide an important framework for investigating the mechanisms operating during the earliest stages of speciation, as reproductive barriers can be examined among divergent lineages in a geographic context. We investigated the evolution of early stages of intrinsic postmating isolation among different populations and lineages of Epidendrum denticulatum, a Neotropical orchid distributed across different biomes in South America. We estimated genetic diversity and structure for both nuclear and plastid markers, using a haplotype network, differentiation tests, Bayesian assignment analysis, and divergence time estimates of the main lineages. Reproductive barriers among divergent lineages were examined by analyzing seed viability following reciprocal crossing experiments. Strong plastid phylogeographic structure was found, indicating that E. denticulatum was restricted to multiple refuges during South American forest expansion events. In contrast, significant phylogeographic structure was not found for nuclear markers, suggesting higher gene flow by pollen than by seeds. Large asymmetries in seed set were observed among different plastid genetic groups, suggesting the presence of polymorphic genic incompatibilities associated with cytonuclear interactions. Our results confirm the importance of phylogeographic studies associated with reproductive isolation experiments and suggest an important role for outbreeding depression during the early stages of lineage diversification.     

Dispersal and habitat fidelity of bog and forest growth forms of Hawaiian Metrosideros (Myrtaceae)

The Hawaiian endemic Metrosideros polymorpha is known for its high levels of morphological diversity and localized adaptation to a range of habitats. At the ecotone between bogs and forests, individuals exhibiting morphological extremes can be found within a few metres of each other. The objective of this study was to examine the genetic diversity and structure of morphologically distinct neighbouring populations of M. polymorpha, growing in bogs and adjacent forests across multiple islands. We explored these relationships using the molecular technique of inter‐simple sequence repeats (ISSRs). The majority (90.79%) of genetic variation was found within populations, 8.53% of the differentiation among populations can be attributed to differences between microhabitat types within islands and very little of the genetic differentiation is explained by the differences among islands (0.68%). These high levels of genetic homogeneity across populations could be the result of extensive gene flow and/or recent isolation of populations. We introduce a nearest genetic neighbour (NGN) analysis to examine detailed relationships of dispersal within and among populations by habitat and island. Using this approach, we provide evidence for habitat fidelity within bog populations and a positive correlation between island age and the proportion of same‐island NGNs. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 558–571.     

Do plant populations on distinct inselbergs talk to each other? A case study of genetic connectivity of a bromeliad species in an Ocbil landscape

Here, we explore the historical and contemporaneous patterns of connectivity among Encholirium horridum populations located on granitic inselbergs in an Ocbil landscape within the Brazilian Atlantic Forest, using both nuclear and chloroplast microsatellite markers. Beyond to assess the E. horridum population genetic structure, we built species distribution models across four periods (current conditions, mid‐Holocene, Last Glacial Maximum [LGM], and Last Interglacial) and inferred putative dispersal corridors using a least‐cost path analysis to elucidate biogeographic patterns. Overall, high and significant genetic divergence was estimated among populations for both nuclear and plastid DNA (Φ_ST(n) = 0.463 and Φ_ST(plastid) = 0.961, respectively, p < .001). For nuclear genome, almost total absence of genetic admixture among populations and very low migration rates were evident, corroborating with the very low estimates of immigration and emigration rates observed among E. horridum populations. Based on the cpDNA results, putative dispersal routes in Sugar Loaf Land across cycles of climatic fluctuations in the Quaternary period revealed that the populations’ connectivity changed little during those events. Genetic analyses highlighted the low genetic connectivity and long‐term persistence of populations, and the founder effect and genetic drift seemed to have been very important processes that shaped the current diversity and genetic structure observed in both genomes. The genetic singularity of each population clearly shows the need for in situ conservation of all of them.     

Hawkmoth and bee pollinators impact pollen dispersal at the landscape but not local scales in two species of Oenothera

Premise

Animal pollinators play an important role in pollen dispersal. Here, we assessed differences in pollen and seed dispersal and the role of pollinator functional groups with different foraging behaviors in generating patterns of genetic diversity over similar geographic ranges for two closely related taxa. We focused on two members of Oenothera section Calylophus (Onagraceae) that co-occur on gypsum outcrops throughout the northern Chihuahuan Desert but differ in floral phenotype and primary pollinator: Oenothera gayleana (bee) and O. hartwegii subsp. filifolia (hawkmoth).

Methods

We measured breeding system and floral traits and studied gene flow and population differentiation at the local (<13 km; four populations) and landscape (60–440 km; five populations) scales using 10–11 nuclear (pollen dispersal) and three plastid (seed dispersal) microsatellite markers.

Results

Both taxa were self-incompatible and floral traits were consistent with expectations for different pollinators. Seed and pollen dispersal patterns were distinctly different for both species. We found no evidence of genetic structure at the local scale but did at the landscape scale; O. gayleana showed greater differentiation and significant isolation by distance than in O. hartwegii subsp. filifolia. The plastid data were consistent with gravity dispersal of seeds and suggest that pollen dispersal is the principal driver of genetic structure in both species.

Conclusions

We demonstrated that pollinator functional groups can impact genetic differentiation in different and predictable ways. Hawkmoths, with larger foraging distances, can maintain gene flow across greater spatial scales than bees.

     

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.     

Genetic structure of the Canarian palm tree (Phoenix canariensis) at the island scale: does the ‘island within islands’ concept apply to species with high colonisation ability?

• Oceanic islands are dynamic settings that often promote within‐island patterns of strong population differentiation. Species with high colonisation abilities, however, are less likely to be affected by genetic barriers, but island size may impact on species genetic structure regardless of dispersal ability.
• The aim of the present study was to identify the patterns and factors responsible for the structure of genetic diversity at the island scale in Phoenix canariensis, a palm species with high dispersal potential. To this end, we conducted extensive population sampling on the three Canary Islands where the species is more abundant and assessed patterns of genetic variation at eight microsatellite loci, considering different within‐island scales.
• Our analyses revealed significant genetic structure on each of the three islands analysed, but the patterns and level of structure differed greatly among islands. Thus, genetic differentiation fitted an isolation‐by‐distance pattern on islands with high population densities (La Gomera and Gran Canaria), but such a pattern was not found on Tenerife due to strong isolation between colonised areas. In addition, we found a positive correlation between population geographic isolation and fine‐scale genetic structure.
• This study highlights that island size is not necessarily a factor causing strong population differentiation on large islands, whereas high colonisation ability does not always promote genetic connectivity among neighbouring populations. The spatial distribution of populations (i.e. landscape occupancy) can thus be a more important driver of plant genetic structure than other island, or species′ life‐history attributes.

     

The interplay of dispersal limitation,rivers, and historical events shapes the genetic structure of an Amazonian frog

Disentangling the impact of landscape features such as rivers and historical events on dispersal is a challenging but necessary task to gain a comprehensive picture of the evolution of diverse biota such as that found in Amazonia. Adenomera andreae, a small, territorial, terrestrial frog species of the Amazonian forest represents a good model for such studies. We combined cytochrome b sequences with 12 microsatellites to investigate the genetic structure at two contrasted spatial scales in French Guiana: along a ～6‐km transect, to evaluate dispersal ability, and between paired bank populations along a ～65‐km stretch of the Approuague river, to test the effect of rivers as barriers to dispersal. We observed significant spatial genetic structure between individuals at a remarkably small geographical scale, and conclude that the species has a restricted dispersal ability that is probably tied to its life‐history traits. Mitochondrial and microsatellite data also indicate a high level of differentiation among populations on opposite banks of the river, and, in some cases, among populations on the same riverbank. These results suggest that the observed population structure in A. andreae is the result of restricted dispersal abilities combined with the action of rivers and Quaternary population isolation. Given that Amazonia hosts a great portion of anurans, as well as other small vertebrates, that display life‐history traits comparable with A. andreae, we argue that our analyses provide new insights into the complex interactions among evolutionary processes shaping Amazonian biodiversity. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 356–373.     

High gene flow in two thrips‐pollinated South‐East Asian pioneer trees: genetic diversity and population structure of Macaranga hypoleuca and M. beccariana (Euphorbiaceae)

As a result of intensive exploitation, disturbed forests now dominate large areas of lowland tropical rainforest in South‐East Asia. The genus Macaranga comprises some of the most important pioneer tree species of the region, among them M. beccariana and M. hypoleuca, two closely related obligate ant‐plants pollinated by thrips. We used nuclear and plastid DNA markers to address questions of genetic diversity and population structure. Twelve plastid haplotypes were detected among 281 samples, three of which were shared between the two study species. Hybrids between the two species appear to be rare. Overall, genetic diversity in both species was moderate to high, with low levels of population differentiation, consistent with other tropical pioneer trees. Genetic structure was generally more pronounced in plastid than in nuclear data, indicating that gene flow via pollen may be more efficient than via seeds. Thrips apparently also serve as efficient pollinators over long distances, perhaps through a combination of passive dispersal by wind and active search for inflorescences in the target area. Our results indicate that M. beccariana and M. hypoleuca populations from recently disturbed habitats do not yet suffer from reduced genetic diversity or increased inbreeding. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 606–621.     

The role of seed dispersal,pollination and historical effects on genetic patterns of an insular plant that has lost its only seed disperser

Aim The genetic structure of many plant species is heavily dependent on their pollinators and seed dispersers, and can thus be altered if either of the associated mutualistic interactions is disrupted. In this study we assess the genetic diversity and structure and infer the seed/pollen gene‐flow patterns among insular populations of Daphne rodriguezii, a shrub pollinated and dispersed by animals that has lost its only disperser (the lizard Podarcis lilfordi) in most of its populations. Location The island of Menorca and the islet of Colom (Balearic Islands, Western Mediterranean). Methods To assess the contribution of gene flow via pollen and seeds to the genetic structure of D. rodriguezii we used amplified fragment length polymorphisms (AFLPs; seeds and pollen) and plastid DNA sequences (cpDNA; seeds). We sampled individuals from all population nuclei of the species (12–19 adults per population): one population in Colom, where the plant–lizard interaction persists, and four in Menorca, where the seed dispersal mutualism disappeared with the extinction of the lizard. Results The highest heterozygosity values were found in Colom and in its closest population (Favàritx), whereas values were lower in the smallest Menorcan populations, which also had higher relatedness among individuals. We found distinct genetic signals between AFLP and cpDNA analyses. While AFLP markers showed low differentiation between populations, cpDNA showed a clear differentiation between them. Main conclusions Our results point to negative impacts of the disperser loss on genetic diversity and relatedness in the smaller and more isolated populations. They also suggest an old isolation by seeds, probably occurring well before the extinction of the lizard (c. 2000 years ago). Gene flow was maintained via pollination; however, the seed disperser loss may ultimately hinder pollinator‐mediated gene flow, as a result of reduced probabilities of effective pollination among increasingly distant and scarce individuals.     

Contrasting patterns of genetic differentiation in Macaronesian lineages of Ilex (Aquifoliaceae)

Islands offer an interesting framework in which to study the effect of geographical isolation on population genetic differentiation. For plant species with high dispersal abilities, however, oceanic barriers may not represent a factor promoting strong population structure. In this work, we analysed seven nuclear microsatellite loci in Ilex (Aquifoliaceae), a bird‐dispersed plant group, to infer patterns of genetic differentiation among Macaronesian taxa: I. canariensis, I. perado ssp. lopezlilloi, I. perado ssp. platyphylla (Canary Islands) and I. perado ssp. azorica (Azores). In agreement with current taxonomic classification, our results revealed a high genetic differentiation between Ilex lineages (I. canariensis and the I. perado complex), and also supported previous hypotheses that these are the result of independent dispersal events to the islands. In contrast, genetic differentiation between I. perado ssp. azorica and the two subspecies from the Canaries was high, suggesting that taxonomic revision may be necessary. Levels of genetic variation at microsatellite loci in ssp. azorica were, in addition, the lowest reported among Macaronesian bird‐dispersed taxa. Lastly, low genetic differentiation was observed between subspecies occurring on the same island (sspp. platyphylla and lopezlilloi). In summary, our results revealed contrasting patterns between Macaronesian Ilex lineages: I. canariensis displayed moderate population structure across islands, whereas the I. perado complex showed strong differentiation among populations sampled on different islands. Thus, the Macaronesian Ilex taxa show that long‐distance dispersal syndromes (ornithochory) do not always ensure genetic connectivity across large areas in island systems. Plant groups that successfully colonized the islands on multiple occasions may have found barriers to gene flow within certain lineages. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2013, 173 , 258–268.     

Genetic differentiation among populations of an oceanic island: The case of Metrosideros boninensis, an endangered endemic tree species in the Bonin Islands

Conservation of endemic species on oceanic islands is an essential issue for biodiversity conservation. Metrosideros boninensis (Myrtaceae) is an endangered tree species endemic to the Bonin Islands of the western North Pacific Ocean. This species is considered to be extremely rare with less than 400 adult individuals, a number that has fluctuated between the 1880s and 1980s through human influence. We analyzed the genetic diversity and genetic structure of this species using amplified fragment length polymorphism markers and microsatellite markers. Genetic diversity of M. boninensis was extremely low compared to related taxa and similar endemic species from small islands. This low genetic diversity might be attributed to a stepwise colonization process with repeated founder bottlenecks in the dispersal pathway to the Bonin Islands. Populations of M. boninensis showed significant genetic differentiation and isolation by distance over a small geographical scale, despite the fact that this species should have extensive gene dispersal ability. This genetic differentiation might be caused by limited gene flow via pollen and seed among populations and genetic drift amid a small number of remnant individuals. Taken together, these findings suggest that the genetic diversity and connectivity of tree populations on islands are more vulnerable to habitat fragmentation than previously thought. We offer some recommendations for management to ameliorate habitat fragmentation and biological invasion.     

Multiple-scaled spatial genetic structures of <Emphasis Type="Italic">Fraxinus mandshurica</Emphasis> over a riparian–mountain landscape in Northeast China

Landscape genetics increasingly focuses on the way in which landscape features cause the fragmentation of lineages of terrestrial organisms. However, landscape features can also provide functional connectivity or corridors, enhancing the dispersal of plant populations, particularly the case in riparian habitat. Unfortunately, recent research in tree genetics has paid little attention to this role. To examine the possible effects of landscape connectivity on the current population genetic distribution of Fraxinus mandshurica and to provide insights into conserving the local genetic diversity for this endangered tree species, we used nine nuclear microsatellite loci to examine the spatial genetic structure of F. mandshurica at multiple-scales over a riparian–mountain landscape in Northeast China. F-statistics indicated that the magnitude of among-population genetic differentiation was significantly higher between the riparian and mountain habitats than within the riparian habitat. Spatial analysis of molecular variance and principal coordinate analysis consistently revealed that this species exhibited a clear landscape genetic structure between the riparian and mountain habitats, despite no significant isolation by distance pattern being identified by the Mantel test. Spatial autocorrelation analysis further demonstrated significant, positive fine-scale spatial genetic structure among individuals over short distances (<80 m) in each mountain population. Conversely, no spatial genetic structures were identified within and among the riparian populations. Overall, the results suggest that seed dispersal is very low among mountain populations; however seed transport is probably enhanced by a secondary phase of hydrochory (water-dispersal) among riparian populations during flooding. Despite this, there was no significant accumulation of genetic diversity in downstream populations along the main channel. This result suggests that hydrochory is not sufficient to produce a clear unidirectional gene flow along the water course, although it may impede the development of spatial genetic structuring within and among riparian populations.     

Fragmentation can increase spatial genetic structure without decreasing pollen-mediated gene flow in a wind-pollinated tree

Fragmentation reduces population sizes, increases isolation between habitats and can result in restricted dispersal of pollen and seeds. Given that diploid seed dispersal contributes more to shaping fine-scale spatial genetic structure (SGS) than haploid pollen flow, we tested whether fine-scale SGS can be sensitive to fragmentation even if extensive pollen dispersal is maintained. Castanopsis sclerophylla (Lindley & Paxton) Schottky (Fagaceae), a wind-pollinated and gravity seed-dispersed tree, was studied in an area of southeast China where its populations have been fragmented to varying extents by human activity. Using different age classes of trees in areas subject to varying extents of fragmentation, we found no significant difference in genetic diversity between prefragmentation vs. postfragmentation C. sclerophylla subpopulations. Genetic differentiation among postfragmentation subpopulations was also only slightly lower than among prefragmentation subpopulations. In the most fragmented habitat, selfing rates were significantly higher than zero in prefragmentation, but not postfragmentation, cohorts. These results suggest that fragmentation had not decreased gene flow among these populations and that pollen flow remains extensive. However, significantly greater fine-scale SGS was found in postfragmentation subpopulations in the most fragmented habitat, but not in less fragmented habitats. This alteration in SGS reflected more restricted seed dispersal, induced by changes in the physical environments and the prevention of secondary seed dispersal by rodents. An increase in SGS can therefore result from more restricted seed dispersal, even in the face of extensive pollen flow, making it a sensitive indicator of the negative consequences of population fragmentation.     

Southern isolation and northern long‐distance dispersal shaped the phylogeography of the widespread,but highly disjunct,European high mountain plant Artemisia eriantha (Asteraceae)

We investigated the range dynamics of Artemisia eriantha, a widespread, but rare, mountain plant with a highly disjunct distribution in the European Alpine System. We focused on testing the roles of vicariance and long‐distance dispersal in shaping the current distribution of the species. To this end, we collected AFLP and plastid DNA sequence data for 17 populations covering the entire distributional range of the species. Strong phylogeographical structure was found in both datasets. AFLP data suggested that almost all populations were genetically strongly differentiated, with 58% of the overall genetic variation partitioned among populations. Bayesian clustering identified five groups of populations: Balkans, Pyrenees, Central Apennines, one southwestern Alpine population and a Widespread cluster (eastern Pyrenees, Alps, Carpathians). Major groups were supported by neighbor‐joining and NeighbourNet analyses. Fourteen plastid haplotypes were found constituting five strongly distinct lineages: Alps plus Pyrenees, Apennines, Balkans, southern Carpathians, and a Widespread group (eastern Pyrenees, northern Carpathians, Mt. Olympus). Plastid DNA data suggested that A. eriantha colonized the European Alpine System in a westward direction. Although, in southern Europe, vicariant differentiation among the Iberian, Italian and Balkan Peninsulas predominated, thus highlighting their importance as glacial refugia for alpine species, in temperate mountain ranges, long‐distance dispersal prevailed. This study emphasizes that currently highly disjunct distributions can be shaped by both vicariance and long‐distance dispersal, although their relative importance may be geographically structured along, for instance, latitude, as in A. eriantha. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 214–226.     

Directed dispersal by rotational shepherding supports landscape genetic connectivity in a calcareous grassland plant

Directed dispersal by animal vectors has been found to have large effects on the structure and dynamics of plant populations adapted to frugivory. Yet, empirical data are lacking on the potential of directed dispersal by rotational grazing of domestic animals to mediate gene flow across the landscape. Here, we investigated the potential effect of large‐flock shepherding on landscape‐scale genetic structure in the calcareous grassland plant Dianthus carthusianorum, whose seeds lack morphological adaptations to dispersal to animals or wind. We found a significant pattern of genetic structure differentiating population within grazed patches of three nonoverlapping shepherding systems and populations of ungrazed patches. Among ungrazed patches, we found a strong and significant effect of isolation by distance (r = 0.56). In contrast, genetic distance between grazed patches within the same herding system was unrelated to geographical distance but significantly related to distance along shepherding routes (r = 0.44). This latter effect of connectivity along shepherding routes suggests that gene flow is spatially restricted occurring mostly between adjacent populations. While this study used nuclear markers that integrate gene flow by pollen and seed, the significant difference in the genetic structure between ungrazed patches and patches connected by large‐flock shepherding indicates the potential of directed seed dispersal by sheep across the landscape.