Limited seed dispersal shapes fine-scale spatial genetic structure in a Neotropical dioecious large-seeded palm

Seed and pollen dispersal contribute to gene flow and shape the genetic patterns of plants over fine spatial scales. We inferred fine-scale spatial genetic structure (FSGS) and estimated realized dispersal distances in Phytelephas aequatorialis, a Neotropical dioecious large-seeded palm. We aimed to explore how seed and pollen dispersal shape this genetic pattern in a focal population. For this purpose, we genotyped 138 seedlings and 99 adults with 20 newly developed microsatellite markers. We tested if rodent-mediated seed dispersal has a stronger influence than insect-mediated pollen dispersal in shaping FSGS. We also tested if pollen dispersal was influenced by the density of male palms around mother palms in order to further explore this ecological process in large-seeded plants. Rodent-mediated dispersal of these large seeds occurred mostly over short distances (mean 34.76 ± 34.06 m) while pollen dispersal distances were two times higher (mean 67.91 ± 38.29 m). The spatial extent of FSGS up to 35 m and the fact that seed dispersal did not increase the distance at which male alleles disperse suggest that spatially limited seed dispersal is the main factor shaping FSGS and contributes only marginally to gene flow within the population. Pollen dispersal distances depended on the density of male palms, decreasing when individuals show a clumped distribution and increasing when they are scattered. Our results show that limited seed dispersal mediated by rodents shapes FSGS in P. aequatorialis, while more extensive pollen dispersal accounts for a larger contribution to gene flow and may maintain high genetic diversity. Abstract in Spanish is available with online material.     

Habitat loss and fragmentation reduce effective gene flow by disrupting seed dispersal in a neotropical palm

Habitat loss and fragmentation often reduce gene flow and genetic diversity in plants by disrupting the movement of pollen and seed. However, direct comparisons of the contributions of pollen vs. seed dispersal to genetic variation in fragmented landscapes are lacking. To address this knowledge gap, we partitioned the genetic diversity contributed by male gametes from pollen sources and female gametes from seed sources within established seedlings of the palm Oenocarpus bataua in forest fragments and continuous forest in northwest Ecuador. This approach allowed us to quantify the separate contributions of each of these two dispersal processes to genetic variation. Compared to continuous forest, fragments had stronger spatial genetic structure, especially among female gametes, and reduced effective population sizes. We found that within and among fragments, allelic diversity was lower and genetic structure higher for female gametes than for male gametes. Moreover, female gametic allelic diversity in fragments decreased with decreasing surrounding forest cover, while male gametic allelic diversity did not. These results indicate that limited seed dispersal within and among fragments restricts genetic diversity and strengthens genetic structure in this system. Although pollen movement may also be impacted by habitat loss and fragmentation, it nonetheless serves to promote gene flow and diversity within and among fragments. Pollen and seed dispersal play distinctive roles in determining patterns of genetic variation in fragmented landscapes, and maintaining the integrity of both dispersal processes will be critical to managing and conserving genetic variation in the face of continuing habitat loss and fragmentation in tropical landscapes.     

The relative contributions of seed and pollen dispersal to gene flow and genetic diversity in seedlings of a tropical palm

Seed and pollen dispersal shape patterns of gene flow and genetic diversity in plants. Pollen is generally thought to travel longer distances than seeds, but seeds determine the ultimate location of gametes. Resolving how interactions between these two dispersal processes shape microevolutionary processes is a long‐standing research priority. We unambiguously isolated the separate and combined contributions of these two dispersal processes in seedlings of the animal‐dispersed palm Oenocarpus bataua to address two questions. First, what is the spatial extent of pollen versus seed movement in a system characterized by long‐distance seed dispersal? Second, how does seed dispersal mediate seedling genetic diversity? Despite evidence of frequent long‐distance seed dispersal, we found that pollen moves much further than seeds. Nonetheless, seed dispersal ultimately mediates genetic diversity and fine‐scale spatial genetic structure. Compared to undispersed seedlings, seedlings dispersed by vertebrates were characterized by higher female gametic and diploid seedling diversity and weaker fine‐scale spatial genetic structure for female gametes, male gametes and diploid seedlings. Interestingly, the diversity of maternal seed sources at seed deposition sites (N _em) was associated with higher effective number of pollen sources (N _ep), higher effective number of parents (N _e) and weaker spatial genetic structure, whereas seed dispersal distance had little impact on these or other parameters we measured. These findings highlight the importance maternal seed source diversity (N _em) at frugivore seed deposition sites in driving emergent patterns of fine‐scale genetic diversity and structure.     

GENETIC CONSEQUENCES OF SEED DISPERSAL IN THREE SYMPATRIC FOREST HERBS. I. HIERARCHICAL POPULATION-GENETIC STRUCTURE

To examine the effects of seed dispersal on spatial genetic structure, we compare three sympatric species of forest herbs in the family Apiaceae whose fruits differ widely in morphological adaptations for animal-attached dispersal. Cryptotaenia canadensis has smooth fruits that are gravity dispersed, whereas Osmorhiza claytonii and Sanicula odorata fruits have appendages that facilitate their attachment to animals. The relative seed-dispersal ability among species, measured as their ability to remain attached to mammal fur, is ranked Sanicula > Osmorhiza > Cryptotaenia. We use a nested hierarchical sampling design to analyze genetic structure at spatial scales ranging from a few meters to hundreds of kilometers. Genetic differentiation among population subdivisions, estimated by average genetic distance and hierarchical F-statistics, has an inverse relationship with dispersal ability such that Cryptotaenia > Osmorhiza > Sanicula. In each species, genetic differentiation increases with distance among population subdivisions. Stochastic variation in gene flow, arising from seed dispersal by attachment to animals, may partly explain the weak relationship between pairwise spatial and genetic distance among populations and heterogeneity in estimates of single locus F-statistics. A hierarchical island model of gene flow is invoked to describe the effects of seed dispersal on population genetic structure. Seed dispersal is the predominant factor affecting variation in gene flow among these ecologically similar, taxonomically related species.     

Species diversity and population density affect genetic structure and gene dispersal in a subtropical understory shrub

Aims The dispersal of pollen and seeds is spatially restricted and may vary among plant populations because of varying biotic interactions, population histories or abiotic conditions. Because gene dispersal is spatially restricted, it will eventually result in the development of spatial genetic structure (SGS), which in turn can allow insights into gene dispersal processes. Here, we assessed the effect of habitat characteristics like population density and community structure on small-scale SGS and estimate historical gene dispersal at different spatial scales.Methods In a set of 12 populations of the subtropical understory shrub Ardisia crenata, we assessed genetic variation at 7 microsatellite loci within and among populations. We investigated small-scale genetic structure with spatial genetic autocorrelation statistics and heterogeneity tests and estimated gene dispersal distances based on population differentiation and on within-population SGS. SGS was related to habitat characteristics by multiple regression.Important findings The populations showed high genetic diversity (H e = 0.64) within populations and rather strong genetic differentiation (F ′ ST = 0.208) among populations, following an isolation-by-distance pattern, which suggests that populations are in gene flow–drift equilibrium. Significant SGS was present within populations (mean Sp = 0.027). Population density and species diversity had a joint effect on SGS with low population density and high species diversity leading to stronger small-scale SGS. Estimates of historical gene dispersal from between-population differentiation and from within-population SGS resulted in similar values between 4.8 and 22.9 m. The results indicate that local-ranged pollen dispersal and inefficient long-distance seed dispersal, both affected by population density and species diversity, contributed to the genetic population structure of the species. We suggest that SGS in shrubs is more similar to that of herbs than to trees and that in communities with high species diversity gene flow is more restricted than at low species diversity. This may represent a process that retards the development of a positive species diversity–genetic diversity relationship.     

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.     

Contemporary pollen and seed dispersal in a Prunus mahaleb population: patterns in distance and direction

Pollination and seed dispersal determine the spatial pattern of gene flow in plant populations and, for those species relying on pollinators and frugivores as dispersal vectors, animal activity plays a key role in determining this spatial pattern. For these plant species, reported dispersal patterns are dominated by short-distance movements with a significant amount of immigration. However, the contribution of seed and pollen to the overall contemporary gene immigration is still poorly documented for most plant populations. In this study we investigated pollination and seed dispersal at two spatial scales in a local population of Prunus mahaleb (L.), a species pollinated by insects and dispersed by frugivorous vertebrates. First, we dissected the relative contribution of pollen and seed dispersal to gene immigration from other parts of the metapopulation. We found high levels of gene immigration (18.50%), due to frequent long distance seed dispersal events. Second, we assessed the distance and directionality for pollen and seed dispersal events within the local population. Pollen and seed movement patterns were non-random, with skewed distance distributions: pollen tended moved up to 548 m along an axis approaching the N-S direction, and seeds were dispersed up to 990 m, frequently along the SW and SE axes. Animal-mediated dispersal contributed significantly towards gene immigration into the local population and had a markedly nonrandom pattern within the local population. Our data suggest that animals can impose distinct spatial signatures in contemporary gene flow, with the potential to induce significant genetic structure at a local level.     

Population structure on breeding grounds of Lake Malawi’s ‘twilight zone’ cichlid fishes

Aim Free‐ranging benthopelagic fishes often have large population sizes and high rates of dispersal. These traits can act to homogenize population structure across the distributional range of a species and to reduce the likelihood of allopatric speciation. The apparent absence of any barriers to gene flow among populations, together with prior molecular evidence for panmixia across the ranges of three species, has resulted in Diplotaxodon, a genus of benthopelagic cichlid fishes of Lake Malawi, being proposed as a candidate case of sympatric speciation. Our aim was to further investigate this possibility by testing for intraspecific genetic subdivision among breeding populations, and intraspecific differences in breeding habitat. Location Lake Malawi, central‐east Africa. Methods We analysed eight microsatellite DNA loci to test for spatial genetic differences among populations on breeding grounds of eight Diplotaxodon species. We also tested for temporal population genetic differences within breeding grounds of three species. Records of ripe Diplotaxodon encountered during sampling were analysed to test if spatial variation in assemblage structure was linked to nearshore water depth and geographic proximity of sampling sites. Results Consistent with previous molecular evidence, within four of the eight species tested we found no evidence of spatial genetic structuring among breeding populations. However, within the other four species we found slight yet significant spatial genetic differences, indicating restricted gene flow among breeding grounds. There was no evidence of temporal genetic differences within sites. Analyses of the distributions of ripe Diplotaxodon revealed differences in assemblage structure linked to nearshore water depth. Main conclusions Together, these results demonstrate both the evolution of fidelity to deep‐water breeding locations in some Diplotaxodon species, and differences in breeding habitat among species. These findings are consistent with a role for divergence of breeding habitat in speciation of these cichlids, possibly promoted by dispersal limitation among geographically segregated spawning aggregations.     

Fine‐scale genetic structure and flowering output of the seagrass Enhalus acoroides undergoing disturbance

Seagrass are under great stress in the tropical coast of Asia, where Enhalus acoroides is frequently the dominant species with a large food web. Here, we investigate the question of the fine‐scale genetic structure of this ecologically important foundation species, subject to severe anthropogenic disturbance in China. The genetic structure will illuminate potential mechanisms for population dynamics and sustainability, which are critical for preservation of biodiversity and for decision‐making in management and restoration. We evaluated the fine‐scale spatial genetic structure (SGS) and flowering output of E. acoroides, and indirectly estimated the relative importance of sexual versus asexual reproduction for population persistence using spatial autocorrelation analysis. Results reveal high clonal diversity for this species, as predicted from its high sexual reproduction output. The stronger Sp statistic at the ramet‐level compared with genet‐level indicates that clonality increases the SGS pattern for E. acoroides. Significant SGS at the genet‐level may be explained by the aggregated dispersal of seed/pollen cohorts. The estimated gene dispersal variance suggests that dispersal mediated by sexual reproduction is more important than clonal growth in this study area. The ongoing anthropogenic disturbance will negatively affect the mating pattern and the SGS patterns in the future due to massive death of shoots, and less frequency of sexual reproduction.     

Microsatellite diversity and genetic structure of fragmented populations of the rare,fire-dependent shrub Grevillea macleayana

Recent habitat loss and fragmentation superimposed upon ancient patterns of population subdivision are likely to have produced low levels of neutral genetic diversity and marked genetic structure in many plant species. The genetic effects of habitat fragmentation may be most pronounced in species that form small populations, are fully self-compatible and have limited seed dispersal. However, long-lived seed banks, mobile pollinators and long adult lifespans may prevent or delay the accumulation of genetic effects. We studied a rare Australian shrub species, Grevillea macleayana (Proteaceae), that occurs in many small populations, is self-compatible and has restricted seed dispersal. However, it has a relatively long adult lifespan (c. 30 years), a long-lived seed bank that germinates after fire and is pollinated by birds that are numerous and highly mobile. These latter characteristics raise the possibility that populations in the past may have been effectively large and genetically homogeneous. Using six microsatellites, we found that G. macleayana may have relatively low within-population diversity (3.2-4.2 alleles/locus; Hexp = 0.420-0.530), significant population differentiation and moderate genetic structure (FST = 0.218) showing isolation by distance, consistent with historically low gene flow. The frequency distribution of allele sizes suggest that this geographical differentiation is being driven by mutation. We found a lack mutation-drift equilibrium in some populations that is indicative of population bottlenecks. Combined with evidence for large spatiotemporal variation of selfing rates, this suggests that fluctuating population sizes characterize the demography in this species, promoting genetic drift. We argue that natural patterns of pollen and seed dispersal, coupled with the patchy, fire-shaped distribution, may have restricted long-distance gene flow in the past.     

Multiscale spatial genetic structure within and between populations of wild cherry trees in nuclear genotypes and chloroplast haplotypes

Spatial genetic structure (SGS) of plants mainly depends on the effective population size and gene dispersal. Maternally inherited loci are expected to have higher genetic differentiation between populations and more intensive SGS within populations than biparentally inherited loci because of smaller effective population sizes and fewer opportunities of gene dispersal in the maternally inherited loci. We investigated biparentally inherited nuclear genotypes and maternally inherited chloroplast haplotypes of microsatellites in 17 tree populations of three wild cherry species under different conditions of tree distribution and seed dispersal. As expected, interpopulation genetic differentiation was 6–9 times higher in chloroplast haplotypes than in nuclear genotypes. This difference indicated that pollen flow 4–7 times exceeded seed flow between populations. However, no difference between nuclear and chloroplast loci was detected in within‐population SGS intensity due to their substantial variation among the populations. The SGS intensity tended to increase as trees became more aggregated, suggesting that tree aggregation biased pollen and seed dispersal distances toward shorter. The loss of effective seed dispersers, Asian black bears, did not affect the SGS intensity probably because of mitigation of the bear loss by other vertebrate dispersers and too few tree generations after the bear loss to alter SGS. The findings suggest that SGS is more variable in smaller spatial scales due to various ecological factors in local populations.     

Partitioning nuclear and chloroplast variation at multiple spatial scales in the neotropical epiphytic orchid, Laelia rubescens

Insights into processes that lead to the distribution of genetic variation within plant species require recognition of the importance of both pollen and seed movement. Here we investigate the contributions of pollen and seed movement to overall gene flow in the Central American epiphytic orchid, Laelia rubescens. Genetic diversity and structure were examined at multiple spatial scales in the tropical dry forest of Costa Rica using nuclear (allozymes) and chloroplast restriction fragment length polymorphism (RFLP) markers, which were found to be diverse (allozymes, P = 73.3%; HE = 0.174; cpDNA, HE = 0.741). Nuclear genetic structure (FSTn) was low at every spatial scale (0.005-0.091). Chloroplast markers displayed more structure (0.073-0.254) but relatively similar patterns. Neither genome displayed significant isolation-by-distance. Pollen and seed dispersal rates did not differ significantly from one another (mp/ms = 1.40) at the broadest geographical scale, among sites throughout Costa Rica. However, relative contributions of pollen and seeds to gene flow were scale-dependent, with different mechanisms determining the dominant mode of gene flow at different spatial scales. Much seed dispersal is highly localized within the maternal population, while some seeds enter the air column and are dispersed over considerable distances. At the intermediate scale (10s to 100s of metres) pollinators are responsible for substantial pollen flow. This species appears capable of distributing its genes across the anthropogenically altered landscape that now characterizes its Costa Rican dry forest habitat.     

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.

     

Revisiting the provenance delineation of a widespread shrub, <Emphasis Type="Italic">Frangula alnus</Emphasis>—the role of spatial,temporal and environmental patterns

Including population genetic aspects into the selection of planting material within the framework of conservation and restoration measures is of vital importance for the long-term persistence of populations. This is especially true facing climate change since genetic diversity and the spread of potentially beneficial alleles are important for the adaptability of populations. Therefore, knowledge about genetic variability within and between populations is a critical aspect when determining provenance regions. In our study, we investigated the population genetic structure of a widespread, insect-pollinated and mainly bird-dispersed shrub species, Frangula alnus, on the basis of seven microsatellites and two chloroplast DNA markers throughout Germany. The aim was to determine the spatial, temporal and ecological processes genetically structuring populations to critically revise existing provenance regions. Therefore, we conducted analyses on different spatial scales (country-wide, regional and local) using the two different marker sets in addition to environmental variables. We detected distinct patterns on all spatial scales which indicated influences of historic recolonization processes, regional differences of seed dispersal across the landscape, as well as small-scale spatial genetic structures attributable to local dispersal processes. No relation of underlying environmental gradients such as temperature or precipitation and genetic patterns was found. We conclude that different aspects of historic and more recent gene flow shape population genetic structures and that a thorough analysis on a variety of spatial, temporal and environmental scales is necessary to appropriately select planting material for conservation and restoration measures. Correspondingly, management advice regarding provenance delineations will be provided.     

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.     

Pollen and seed flow under different predominant winds in wind-pollinated and wind-dispersed species <Emphasis Type="Italic">Engelhardia roxburghiana</Emphasis>

In most plants, the contributions of pollen and seed flow to their genetic structures are generally difficult to disentangle. For typical wind-pollinated and wind-dispersed species Engelhardia roxburghiana in a 20-ha natural forest plot in lower subtropic China, because the prevailing wind directions change during its pollen release and seed dispersal seasons, we could compare its genetic structures in different directions, which could result primarily from pollen or seed flow. Furthermore, because the plot has undergone from an open to a closed canopy stage historically, we also examined forest canopy effects on gene flow in different generations and different directions. Using 522 E. roxburghiana individuals mapped in the plot, our results revealed that greater pollen flow led to biased gene flow in the pollen dispersal-predominant direction (pollen direction), while greater seed flow generated less spatial genetic structure in the seed dispersal-predominant direction (seed direction). The results predicted from generalized additive models indicated that canopy closure enhanced resistance to gene flow from the old generation to the new generation. Analyses by landscape genetic models for the new generation revealed that gene flow associated with pollen direction was more strongly affected by canopy than with seed direction. Our study is new by proposing an alternative way to separate effects of the pollen and seed flow on spatial variation patterns in E. roxburghiana. To our knowledge, our study is also the first attempt to use landscape genetic models to represent canopy effects for different dispersal vectors in spatial scales only up to a few hundred meters.     

Effects of Competition and Temporal Variation on the Evolutionary Potential of Two Native Bunchgrass Species

The capacity of restored plant populations to adapt to new environmental challenges depends on within‐population genetic variation. We examined how much genetic and environmentally based variation for fitness‐associated traits exists within populations of two native grasses commonly used for restoration in California. We were also interested in understanding how phenotypic expression of genetic variation for these traits varies with growth environment. Thirty maternal families of Elymus glaucus (Blue wild rye) and Nassella pulchra (Purple needlegrass) were sampled from both coastal and interior populations and reciprocally transplanted into three replicated common gardens with and without interspecific competition at each site. Reproductive output of families differed both among years and with competition treatments. Phenotypic expression of genetic variation in culm production differed among populations and was very low when families were grown with interspecific competition. Without interspecific competition, the degree of genetic determination peaked in year two in both species (8.4 and 15.1% in E. glaucus and N. pulchra, respectively). Significant genetic differences in reproduction and phenotypic plasticity were found among N. pulchra subpopulations sampled less than 3 km apart, further highlighting the importance of thoroughly sampling available genetic variation in populations used for restoration. The variable and generally low expression of genetic variation indicates that rates of adaptation in restored populations of these native grasses may vary temporally and may be especially slow within competitive environments.     

Limiting inbreeding in disjunct and isolated populations of a woody shrub

Pollen movements and mating patterns are key features that influence population genetic structure. When gene flow is low, small populations are prone to increased genetic drift and inbreeding, but naturally disjunct species may have features that reduce inbreeding and contribute to their persistence despite genetic isolation. Using microsatellite loci, we investigated outcrossing levels, family mating parameters, pollen dispersal, and spatial genetic structure in three populations of Hakea oldfieldii, a fire‐sensitive shrub with naturally disjunct, isolated populations prone to reduction in size and extinction following fires. We mapped and genotyped a sample of 102 plants from a large population, and all plants from two smaller populations (28 and 20 individuals), and genotyped 158–210 progeny from each population. We found high outcrossing despite the possibility of geitonogamous pollination, small amounts of biparental inbreeding, a limited number of successful pollen parents within populations, and significant correlated paternity. The number of pollen parents for each seed parent was moderate. There was low but significant spatial genetic structure up to 10 m around plants, but the majority of successful pollen came from outside this area including substantial proportions from distant plants within populations. Seed production varied among seven populations investigated but was not correlated with census population size. We suggest there may be a mechanism to prevent self‐pollination in H. oldfieldii and that high outcrossing and pollen dispersal within populations would promote genetic diversity among the relatively small amount of seed stored in the canopy. These features of the mating system would contribute to the persistence of genetically isolated populations prone to fluctuations in size.     

DIRECT AND INDIRECT ESTIMATES OF SEED VERSUS POLLEN MOVEMENT WITHIN A POPULATION OF PONDEROSA PINE

We examined the spatial distribution of maternally inherited mitochondrial DNA and paternally inherited chloroplast DNA polymorphisms in a permanently marked stand of ponderosa pine (Pinus ponderosa Laws). Movement of maternally inherited mtDNA occurs only via seed dispersal, and mtDNA haplotypes showed significant patch structure. Moreover, individuals within patches identified by mtDNA haplotypes were related approximately as half-sibs based upon analysis of allozyme genotypes. Thus, seed dispersal is limited within the population, and creates matrilineal clusters in space. By contrast, paternally inherited cpDNA is dispersed by movement of both seed and pollen. Chloroplast DNA polymorphisms showed no evidence of patch structure, but rather a weak (and nonsignificant) trend toward hyperdispersion, suggesting nearly unlimited movement of pollen among trees within this stand. Two of the trees had unique allozyme alleles, which were used to directly measure pollen movement away from those trees. Marked pollen was as likely to disperse across the population as it was to fertilize near neighbors.     

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

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