Long-distance dispersal of seeds in the fire-tolerant shrub Banksia attenuata

Long-distance dispersal (LDD) of seeds enables alleles, individuals and species of plants to (re)colonize suitable but remote habitats. Banksia attenuata is a long-lived resprouting shrub restricted to dune crests in fire-prone sclerophyll shrublands of the Eneabba sandplain, southwestern Australia. Highly polymorphic microsatellite DNA genetic markers and population assignment tests were employed to identify LDD immigrants among 788 individuals from 27 stands of B. attenuata comprising a metapopulation. Of the 487 (61.8% of the total) individuals unambiguously assigned to a unique source population, 27 (5.5%) were identified as immigrants by assignment to a known population other than that from which they were sampled, while the remaining 460 were assigned to the population from which they were sampled. The distance between source and sink populations for these immigrants ranged from 0.2 to 2.6  km, averaging 1.4  km, and broadly trending in the direction of seasonal winds. These results suggest that B. attenuata has similar long-distance seed dispersal properties as its co-occurring shorter-lived and fire-sensitive congener, B. hookeriana , despite fewer, larger and less mobile seeds. The frequency and distance of LDD for seeds observed in both species (5.5–6.8%) helps explain the persistence of populations on these geographically isolated dunes, where they are subject to local extinction from recurrent fire and severe summer drought, and will remain important under predicted climate change conditions. Analysis also revealed that species richness of the functional group to which B. attenuata belongs was positively correlated with the number of immigrants identified per dune, and such correlation was likely driven by environmental properties of the dunes, particularly water availability.     

Spatial and temporal genetic structure in a hybrid cordgrass invasion

Invasive hybrids and their spread dynamics pose unique opportunities to study evolutionary processes. Invasive hybrids of native Spartina foliosa and introduced S. alterniflora have expanded throughout San Francisco Bay intertidal habitats within the past 35 years by deliberate plantation and seeds floating on the tide. Our goals were to assess spatial and temporal scales of genetic structure in Spartina hybrid populations within the context of colonization history. We genotyped adult and seedling Spartina using 17 microsatellite loci and mapped their locations in three populations. All sampled seedlings were hybrids. Bayesian ordination analysis distinguished hybrid populations from parent species, clearly separated the population that originated by plantation from populations that originated naturally by seed and aligned most seedlings within each population. Population genetic structure estimated by analysis of molecular variance was substantial (F_ST=0.21). Temporal genetic structure among age classes varied highly between populations. At one population, the divergence between adults and 2004 seedlings was low (F_ST=0.02) whereas at another population this divergence was high (F_ST=0.26). This latter result was consistent with local recruitment of self-fertilized seed produced by only a few parental plants. We found fine-scale spatial genetic structure at distances less than ∼200 m, further supporting local seed and/or pollen dispersal. We posit a few self-fertile plants dominating local recruitment created substantial spatial genetic structure despite initial long-distance, human dispersal of hybrid Spartina through San Francisco Bay. Fine-scale genetic structure may more strongly develop when local recruits are dominated by the offspring of a few self-fertile plants.     

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.     

The pollen dispersal kernel and mating system of an insect-pollinated tropical palm,Oenocarpus bataua

Pollen dispersal shapes the local genetic structure of plant populations and determines the opportunity for local selection and genetic drift, but has been well studied in few animal-pollinated plants in tropical rainforests. Here, we characterise pollen movement for an insect-pollinated Neotropical canopy palm, Oenocarpus bataua, and relate these data to adult mating system and population genetic structure. The study covers a 130-ha parcel in which all adult trees (n=185) were mapped and genotyped at 12 microsatellite loci, allowing us to positively identify the source tree for 90% of pollination events (n=287 of 318 events). Mating system analysis showed O. bataua was effectively outcrossed (t_m=1.02) with little biparental inbreeding (t_m−t_s=−0.005) and an average of 5.4 effective pollen donors (N_ep) per female. Dispersal distances were relatively large for an insect-pollinated species (mean=303 m, max=1263 m), and far exceeded nearest-neighbour distances. Dispersal kernel modelling indicated a thin-tailed Weibull distribution offered the best fit to the genetic data, which contrasts with the fat-tailed kernels typically reported for pollen dispersal in trees. Preliminary analyses suggest that our findings may be explained, at least in part, by a relatively diffuse spatial and temporal distribution of flowering trees. Comparison with previously reported estimates of seed movement for O. bataua suggests that pollen and seed dispersal distances may be similar. These findings add to the growing body of information on dispersal in insect-pollinated trees, but underscore the need for continued research on tropical systems in general, and palms in particular.     

Genetic Diversity and Spatial Genetic Structure of the Grassland Perennial Saxifraga granulata along Two River Systems

Due to changes in land use, the natural habitats of an increasing number of plant species have become more and more fragmented. In landscapes that consist of patches of suitable habitat, the frequency and extent of long-distance seed dispersal can be expected to be an important factor determining local genetic diversity and regional population structure of the remaining populations. In plant species that are restricted to riparian habitats, rivers can be expected to have a strong impact on the dynamics and spatial genetic structure of populations as they may enable long-distance seed dispersal and thus maintain gene flow between fragmented populations. In this study, we used polymorphic microsatellite markers to investigate the genetic diversity and the spatial genetic structure of 28 populations of Saxifraga granulata along two rivers in central Belgium. We hypothesized that rivers might be essential for gene flow among increasingly isolated populations of this species. Genetic diversity was high (H_S = 0.68), which to a certain extent can be explained by the octoploid nature of S. granulata in the study area. Populations along the Dijle and Demer rivers were also highly differentiated (G” _ST = 0.269 and 0.164 and D _EST = 0.190 and 0.124, respectively) and showed significant isolation-by-distance, indicating moderate levels of gene flow primarily between populations that are geographically close to each other. Along the river Demer population genetic diversity was higher upstream than downstream, suggesting that seed dispersal via the water was not the primary mode of dispersal. Overall, these results indicate that despite increasing fragmentation populations along both rivers were highly genetically diverse. The high ploidy level and longevity of S. granulata have most likely buffered negative effects of fragmentation on genetic diversity and the spatial genetic structure of populations in riparian grasslands.     

Low levels of realized seed and pollen gene flow and strong spatial genetic structure in a small,isolated and fragmented population of the tropical tree Copaifera langsdorffii Desf

Over the past century, the Brazilian Atlantic forest has been reduced to small, isolated fragments of forest. Reproductive isolation theories predict a loss of genetic diversity and increases in inbreeding and spatial genetic structure (SGS) in such populations. We analysed eight microsatellite loci to investigate the pollen and seed dispersal patterns, genetic diversity, inbreeding and SGS of the tropical tree Copaifera langsdorffii in a small (4.8 ha), isolated population. All 112 adult trees and 128 seedlings found in the stand were sampled, mapped and genotyped. Seedlings had significantly lower levels of genetic diversity (A=16.5±0.45, mean±95% s.e.; H_e=0.838±0.006) than did adult trees (A=23.2±0.81; H_e=0.893±0.030). Parentage analysis did not indicate any seed immigration (m_seeds=0) and the pollen immigration rate was very low (m_pollen=0.047). The average distance of realized pollen dispersal within the stand was 94 m, with 81% of the pollen travelling <150 m. A significant negative correlation was found between the frequency and distance of pollen dispersal (r=−0.79, P<0.01), indicating that short-distance pollinations were more frequent. A significant SGS for both adults (∼50 m) and seedlings (∼20 m) was also found, indicating that most of the seeds were dispersed over short distances. The results suggested that the spatial isolation of populations by habitat fragmentation can restrict seed and pollen gene flow, increase SGS and affect the genetic diversity of future generations.     

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.     

Colonization dynamics of a clonal pioneer plant on a glacier foreland inferred from spatially explicit and size-structured matrix models

The regional distribution of a plant species is a result of the dynamics of extinctions and colonizations in suitable habitats, especially in strongly fragmented landscapes. Here, we studied the role of spatial dynamics of the long-lived, clonal pioneer plant Geum reptans occurring on glacier forelands in the European Alps. We used demographic data from several years and sites in the Swiss Alps in combination with dispersal data to parametrize a matrix model for G. reptans to simulate extinctions, colonizations and spatial spread of established populations on glacial forelands. We used different scenarios with varying germination rates, wind and animal dispersal capabilities, and modes of spatial spread (seed-only vs clonal spread), resulting in population growth rates (λ) ranging from 1.04 to 1.20. Our results suggest that due to the low germination rate (~1%) and the very limited wind dispersal distances (99.8% of seeds are dispersed < 5 m), G. reptans has a low probability of establishing new populations and a very low spatial spread by seed dispersal alone. In contrast to the low rate of establishment, the persistence of established populations is high and even populations of only a few individuals have an extinction probability of less than 25% within 100 years. This high persistency is partly due to clonal reproduction via aboveground stolons. Clonal reproduction increases the population size and contributes considerably to the spatial spread of established populations. Our simulation results together with the known pattern of molecular diversity of G. reptans indicate that the occurrence of populations of this species in the Alps is unlikely to be a result of recent colonizations by long-distance dispersal, but rather a result of post-glacial colonizations by large migrating populations that were fragmented when glaciers retreated. Additionally, our simulations suggest that the currently observed high rates of glacial retreat might be too fast for pioneer plants, such as G. reptans, to keep up with the retreating ice and therefore might threaten existing populations.     

Genetic variation of liverleaf (Hepatica nobilis Schreb.) in Bavaria against the background of seed transfer guidelines in forestry and restoration

The liver leaf, Hepatica nobilis (Ranunculaceae), is a perennial forest understory herb with specific habitat requirements, which occurs on calcareous soils in beech and oak forests. In Bavaria it can be found, therefore, only in four geographic regions, which are Franconia, the Franconian-Swabian Jura, the prealpine transitions region and the prealpine moraine belt with the Alps. In German forestry, provenance delineations are used since 1987. Similarly for herbs seed provenances and production areas are applied in commercial seed production and restoration. In this study we analyzed whether the genetic variation of H. nobilis reflects the geographic distribution of the species in Bavaria or the provenance delineations used for seed production.We applied AFLPs to study genetic variation within and between 24 populations of H. nobilis.Our analysis revealed high levels of genetic variation within and moderate variation between populations. Variation between seed production areas and seed provenances was low and comparable to the variation between geographic regions. Genetic variation within populations or rare fragments did not differ between production areas, provenances or geographic regions. A significant positive relation of genetic and geographic distances was present within 100 km.Pollination and seed dispersal seem mainly to happen within populations of the myrmecochorous H. nobilis, whereas long-distance dispersal is presumably occasional and random. Our study supports the relevance of dispersal traits for seed collections used in conservation. The protection of historically old beech and oak forests is exceedingly important to preserve the genetic variation of H. nobilis in Bavaria.     

High paternal diversity in the self-incompatible herb Arabidopsis halleri despite clonal reproduction and spatially restricted pollen dispersal

The number of sires fertilizing a given dam is a key parameter of the mating system in species with spatially restricted offspring dispersal, since genetic relatedness among maternal sibs determines the intensity of sib competition. In flowering plants, the extent of multiple paternity is determined by factors such as floral biology, properties of the pollen vector, selfing rate, spatial organization of the population, and genetic compatibility between neighbours. To assess the extent of multiple paternity and identify ecological factors involved, we performed a detailed study of mating patterns in a small population of a self-incompatible clonal herb, Arabidopsis halleri . We mapped and genotyped 364 individuals and 256 of their offspring at 12 microsatellite loci and jointly analysed the level of multiple paternity, pollen and seed dispersal, and spatial genetic structure. We found very low levels of correlated paternity among sibs ( P _full-sib = 3.8%) indicating high multiple paternity. Our estimate of the outcrossing rate was 98.7%, suggesting functional self-incompatibility. The pollen dispersal distribution was significantly restricted (mean effective pollen dispersal distance: 4.42 m) but long-distance successful pollination occurred and immigrating pollen was at most 10% of all pollination events. Patterns of genetic structure indicated little extent of clonal reproduction, and a low but significant spatial genetic structure typical for a self-incompatible species. Overall, in spite of restricted pollen dispersal, the multiple paternity in this self-incompatible species was very high, a result that we interpret as a consequence of high plant density and high pollinator service in this population.     

Life in the desert: The impact of geographic and environmental gradients on genetic diversity and population structure of Ivesia webberi

For range‐restricted species with disjunct populations, it is critical to characterize population genetic structure, gene flow, and factors that influence functional connectivity among populations in order to design effective conservation programs. In this study, we genotyped 314 individuals from 16 extant populations of Ivesia webberi, a United States federally threatened Great Basin Desert using six microsatellite loci. We assessed the effects of Euclidean distance, landscape features, and ecological dissimilarity on the pairwise genetic distance of the sampled populations, while also testing for a potential relationship between I. webberi genetic diversity and diversity in the vegetative communities. The results show low levels of genetic diversity overall (H _e = 0.200–0.441; H _o = 0.192–0.605) and high genetic differentiation among populations. Genetic diversity was structured along a geographic gradient, congruent with patterns of isolation by distance. Populations near the species’ range core have relatively high genetic diversity, supporting in part a central‐marginal pattern, while also showing some evidence for a metapopulation dynamic. Peripheral populations have lower genetic diversity, significantly higher genetic distances, and higher relatedness. Genotype cluster admixture results suggest a complex dispersal pattern among populations with dispersal direction and distance varying on the landscape. Pairwise genetic distance strongly correlates with elevation, actual evapotranspiration, and summer seasonal precipitation, indicating a role for isolation by environment, which the observed phenological mismatches among the populations also support. The significant correlation between pairwise genetic distance and floristic dissimilarity in the germinated soil seed bank suggests that annual regeneration in the plant communities contribute to the maintenance of genetic diversity in I. webberi.     

Long-Distance Dispersal by Sea-Drifted Seeds Has Maintained the Global Distribution of Ipomoea pes-caprae subsp. brasiliensis (Convolvulaceae)

Ipomoea pes-caprae (Convolvulaceae), a pantropical plant with sea-drifted seeds, is found globally in the littoral areas of tropical and subtropical regions. Unusual long-distance seed dispersal has been believed to be responsible for its extraordinarily wide distribution; however, the actual level of inter-population migration has never been studied. To clarify the level of migration among populations of I. pes-caprae across its range, we investigated nucleotide sequence variations by using seven low-copy nuclear markers and 272 samples collected from 34 populations that cover the range of the species. We applied coalescent-based approaches using Bayesian and maximum likelihood methods to assess migration rates, direction of migration, and genetic diversity among five regional populations. Our results showed a high number of migrants among the regional populations of I. pes-caprae subsp. brasiliensis, which suggests that migration among distant populations was maintained by long-distance seed dispersal across its global range. These results also provide strong evidence for recent trans-oceanic seed dispersal by ocean currents in all three oceanic regions. We also found migration crossing the American continents. Although this is an apparent land barrier for sea-dispersal, migration between populations of the East Pacific and West Atlantic regions was high, perhaps because of trans-isthmus migration via pollen dispersal. Therefore, the migration and gene flow among populations across the vast range of I. pes-caprae is maintained not only by seed dispersal by sea-drifted seeds, but also by pollen flow over the American continents. On the other hand, populations of subsp. pes-caprae that are restricted to only the northern part of the Indian Ocean region were highly differentiated from subsp. brasiliensis. Cryptic barriers that prevented migration by sea dispersal between the ranges of the two subspecies and/or historical differentiation that caused local adaptation to different environmental factors in each region could explain the genetic differentiation between the subspecies.     

Limitations to reproductive output and genetic rescue in populations of the rare shrub Grevillea repens (Proteaceae)

Background and Aims

When conserving rare plant species, managers are often faced with small and/or isolated populations displaying low levels of sexual reproduction and genetic variation. One option for reinvigorating these populations is the introduction of genetic material from other sites, but in some cases fitness may be reduced as a result of outbreeding depression. Here the pollination biology of the rare shrub Grevillea repens is studied across its natural range and reproductive responses following cross-pollination among populations are examined to determine factors that may be limiting sexual reproduction and the potential for genetic rescue.

Methods

Pollen manipulation treatments (self-, autogamous self-, cross- and open pollination) were applied to flowers to examine the breeding system and fruit and seed production in five populations of G. repens. Pollen production, presentation and viability were investigated and interpopulation crosses of increasing genetic distance performed among the populations.

Key Results

The study species is self-incompatible and displayed very low natural seed set over two seasons, due partly to low pollen viability in one of the populations. Within-population crossing increased fruit and seed production at some sites, indicating pollinator limitation. Interpopulation crosses further increased reproductive output in one population, suggesting mate limitation, and for this site there was a positive relationship between genetic distance among populations and the size of genetic rescue benefits. However, in other populations there was a decrease in fruit and seed set with increasing genetic distance.

Conclusions

The results highlight that management strategies involving interpopulation crosses can improve reproductive output in small, isolated populations of rare plants, but guidelines need to be developed on a population by population basis.Key words: Grevillea repens, Proteaceae, genetic rescue, pollination ecology, self-incompatibility, breeding system, interpopulation cross, outbreeding depression, pollinator limitation, mate limitation, resource limitation     

Microsatellite Analysis of Museum Specimens Reveals Historical Differences in Genetic Diversity between Declining and More Stable Bombus Species

Worldwide most pollinators, e.g. bumblebees, are undergoing global declines. Loss of genetic diversity can play an essential role in these observed declines. In this paper, we investigated the level of genetic diversity of seven declining Bombus species and four more stable species with the use of microsatellite loci. Hereto we genotyped a unique collection of museum specimens. Specimens were collected between 1918 and 1926, in 6 provinces of the Netherlands which allowed us to make interspecific comparisons of genetic diversity. For the stable species B. pascuorum, we also selected populations from two additional time periods: 1949–1955 and 1975–1990. The genetic diversity and population structure in B. pascuorum remained constant over the three time periods. However, populations of declining bumblebee species showed a significantly lower genetic diversity than co-occurring stable species before their major declines. This historical difference indicates that the repeatedly observed reduced genetic diversity in recent populations of declining bumblebee species is not caused solely by the decline itself. The historically low genetic diversity in the declined species may be due to the fact that these species were already rare, making them more vulnerable to the major drivers of bumblebee decline.     

Conservation genetics of Heritiera littoralis (Sterculiaceae), a threatened mangrove in China,based on AFLP and ISSR markers

AFLP and ISSR markers were used to determine the genetic variations in eight mangrove and non-mangrove populations of Heritiera littoralis (Sterculiaceae), a threatened species in China. Our results showed a moderate to high level of genetic variation in this species (P = 63.69%, H_T = 0.20 for AFLP; P = 76.07%, H_T = 0.22 for ISSR), and a relatively high level of genetic differentiation among populations (G_ST = 0.24 for AFLP and 0.27 for ISSR). Life history traits, long-distance dispersal of floating seeds, and local environments may play important roles in shaping the genetic diversity and genetic structure of this species. Investigation of the plant’s reproductive capacity showed that the natural seed production of H. littoralis was low, as was the germination rate and the transformation rate from juvenile to adult. H. littoralis is seriously threatened and is in urgent need of conservation in China.     

Spatial genetic structure in wild cherry (Prunus avium L.): I. variation among natural populations of different density

Conservation of forest genetic resources requires intensive knowledge of the spatial arrangement of genetic diversity. In this study, we used four natural Prunus avium stands in Germany with contrasting for densities to understand patterns of spatial genetic structure. To this end, we genotyped adults and saplings at eight microsatellite markers, 54 AFLP loci and at the gametophytic incompatibility locus. We estimated levels of clonal propagation, spatial genetic structure and gene dispersal. High mortality occurred among young clonal individuals, as depicted by the lower clonal diversity in saplings. Contrasting levels of spatial genetic structure were observed among markers, ontogenic stages and populations. AFLP were more efficient for detecting spatial autocorrelation but did not allow us to differentiate low and high density populations, while high density populations showed substantially stronger spatial genetic structure at microsatellite loci. Furthermore, kinship decreased with tree age only in low density stands. We discuss the present results in terms of population history, pollen and seed dispersal and population density. Although conspecific density seems to be an interesting indicator of genetic diversity for conservation programmes, we still need to disentangle the relative influence of clonal propagation and density on the strength of spatial genetic structure. Simulation studies are needed to further address this question.     

Contrasting patterns of clonality and fine-scale genetic structure in two rare sedges with differing geographic distributions

For plants with mixed reproductive capabilities, asexual reproduction is more frequent in rare species and is considered a strategy for persistence when sexual recruitment is limited. We investigate whether asexual reproduction contributes to the persistence of two co-occurring, rare sedges that both experience irregular seed set and if their differing geographic distributions have a role in the relative contribution of clonality. Genotypic richness was high (R=0.889±0.02) across the clustered populations of Lepidosperma sp. Mt Caudan and, where detected, clonal patches were small, both in ramet numbers (⩽3 ramets/genet) and physical size (1.3±0.1 m). In contrast, genotypic richness was lower in the isolated L. sp. Parker Range populations, albeit more variable (R=0.437±0.13), with genets as large as 17 ramets and up to 5.8 m in size. Aggregated clonal growth generated significant fine-scale genetic structure in both species but to a greater spatial extent and with additional genet-level structure in L. sp. Parker Range that is likely due to restricted seed dispersal. Despite both species being rare, asexual reproduction clearly has a more important role in the persistence of L. sp. Parker Range than L. sp. Mt Caudan. This is consistent with our prediction that limitations to sexual reproduction, via geographic isolation to effective gene exchange, can lead to greater contributions of asexual reproduction. These results demonstrate the role of population isolation in affecting the balance of alternate reproductive modes and the contextual nature of asexual reproduction in rare species.     

The colonization history of the Mediterranean dwarf palm (<Emphasis Type="Italic">Chamaerops humilis</Emphasis> L., Palmae)

Chamaerops humilis L. (Mediterranean dwarf palm) is an important floristic element of the western Mediterranean region because it is the only palm species naturally distributed in both Europe and Africa. The combination of a time-calibrated phylogeny, a haplotype network and genetic diversity analyses based on plastid sequences, together with previous nuclear DNA fingerprint results, helped reconstruct the colonization history of the dwarf palm. Based on a sample of 218 individuals taken from 29 geographical areas that cover the current distribution of Chamaerops, we detected four plastid DNA (petA-psbJ) haplotypes distributed in two haplotype groups (lineage 1: haplotypes A/B; lineage 2: haplotypes C/D). Haplotypes A, B, and C showed a widespread geographical distribution in both Africa and Europe, whereas haplotype D was restricted to two African localities. Paleobotanical data, species distribution modeling and divergence time estimates suggest that Chamaerops diverged from Trachycarpus in the Miocene (27.05–6.05 Ma), followed by a split of the two C. humilis lineages that remained isolated during the Miocene-Pliocene. Divergence estimates also support a derived split into two haplotypes (A/B) in the Pleistocene, when the Mediterranean Sea barrier was in existence. This, together with geographical distribution of haplotypes A and B, strongly suggests that the disjunct distribution of C. humilis haplotypes in Europe and Africa is the result of long-distance dispersal (LDD) events rather than vicariance. In agreement with recurrent gene flow (events of LDD colonization), AMOVA revealed that most of the genetic variance was found among populations (61.52%). Irrespective of predominant plant translocations by humans or seed dispersal by natural means, our results support that current populations are the result of relatively recent contacts between Africa and Europe in the Quaternary.     

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.     

Bayesian approach reveals confounding effects of population size and seasonality on outcrossing rates in a fragmented subalpine conifer

Mating systems have long been recognized as key factors determining genetic structure within and between populations. Outcrossing promotes genetic diversity and gene flow between populations, while inbreeding, on the other hand, decreases recombination rates, facilitating fixation of co-adapted genes. In small populations, selfing moderates pollen limitation because of low mate availability, but at the cost of increased inbreeding depression. These conflicts are of more than theoretical interest; they are critical for the management of endangered species. In order to help designing conservation strategies for the management of the gene pool of fragmented populations of Pinus cembra, a protected species in Poland, we have characterized pollen flow and mating structure using nuclear microsatellite markers. We demonstrated that P. cembra in the studied stands of the Tatra Mts. is characterized by an average outcrossing rate (t) of 0.72. Unlike with the existing approaches, using the newly developed Bayesian method, we found that population size and seasonal variation had confounding effects on outcrossing rates. In concordance with predictions, large populations showed significantly higher outcrossing rates (t?=?0.89) than smaller ones (t?=?0.51). Temporal variation revealed in the outcrossing rate might be linked with masting behavior of the species. On the other hand, we showed that outcrossing rates were not associated with a trunk diameter of a mother tree. Our study also demonstrated that biparental inbreeding is a significant component of mating system. However, we further show that pollen dispersal follows a fat-tailed distribution (with the average dispersal distance of 1,267 m) so that at least some long-distance pollen dispersal must be occurring. Overall, we conclude that the high inbreeding (both selfing and mating between relatives) found in P. cembra buffers for pollen limitation. We argue that small, isolated stands can be at risk of gene pool erosion, despite the potential for long-distance pollen and seed dispersal.