Long-distance Pollen Dispersal Research Articles

Historical and contemporary gene dispersal in wild carrot (Daucus carota ssp. carota) populations.

Wild carrot is the ancestor of cultivated carrot and is the most important gene pool for carrot breeding. Transgenic carrot may be released into the environment in the future. The aim of the present study was to determine how far a gene can disperse in wild carrot populations, facilitating risk assessment and management of transgene introgression from cultivated to wild carrots and helping to design sampling strategies for germplasm collections. Wild carrots were sampled from Meijendel and Alkmaar in The Netherlands and genotyped with 12 microsatellite markers. Spatial autocorrelation analyses were used to detect spatial genetic structures (SGSs). Historical gene dispersal estimates were based on an isolation by distance model. Mating system and contemporary pollen dispersal were estimated using 437 offspring of 20 mothers with different spatial distances and a correlated paternity analysis in the Meijendel population. Significant SGSs are found in both populations and they are not significantly different from each other. Combined SGS analysis indicated significant positive genetic correlations up to 27 m. Historical gene dispersal sigma(g) and neighbourhood size N(b) were estimated to be 4-12 m [95 % confidence interval (CI): 3-25] and 42-73 plants (95 % CI: 28-322) in Meijendel and 10-31 m (95 % CI: 7-infinity) and 57-198 plants (95 % CI: 28-infinity) in Alkmaar with longer gene dispersal in lower density populations. Contemporary pollen dispersal follows a fat-tailed exponential-power distribution, implying pollen of wild carrots could be dispersed by insects over long distance. The estimated outcrossing rate was 96 %. SGSs in wild carrots may be the result of high outcrossing, restricted seed dispersal and long-distance pollen dispersal. High outcrossing and long-distance pollen dispersal suggest high frequency of transgene flow might occur from cultivated to wild carrots and that they could easily spread within and between populations.

Plant Parentage, Pollination, and Dispersal: How DNA Microsatellites Have Altered the Landscape

DNA microsatellites provide plant ecologists with molecular markers precise enough to assign parentage to seeds and seedlings. This allows the exact distance and trajectory of successful pollen to be traced to characterize pollination patterns. Parentage assignment of established seedlings also allows researchers to accurately determine how far new recruits have traveled from their seed parent. This paper reviews the history and development of molecular parentage assignment in studies of native plants, as well as the limitations and constraints to this approach. This paper also reviews 53 articles published in the past 15 years that use parentage assignment to study pollination and seed dispersal in native plants. These parentage studies have overturned many common assumptions regarding pollen and seed dispersal patterns. They show that long-distance dispersal of pollen is common in both wind and animal dispersed systems, with average pollination distances commonly being hundreds of meters. The pollination neighborhood is often extremely large, and simple dispersal functions based on distance alone fail to make accurate predictions of pollination. Rather than hindering gene flow, fragmentation and isolation sometimes, and perhaps even commonly, results in increased pollination distances. Studies of seed dispersal using parentage assignment have also yielded some surprises. We now know that it may be erroneous to assume that seeds growing under the crown of a conspecific adult are growing beneath their mother, or that seed dispersal distances are more limited than pollen dispersal distances. Taken together, the studies to date demonstrate that both seed and pollen dispersal are quite complex phenomena influenced by many ecological processes.

Effective Long-Distance Pollen Dispersal in Centaurea jacea

BackgroundAgri-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 FindingsIn 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.ConclusionsWe 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.

Increases in air temperature can promote wind-driven dispersal and spread of plants

Long-distance dispersal (LDD) of seeds and pollen shapes the spatial dynamics of plant genotypes, populations and communities. Quantifying LDD is thus important for predicting the future dynamics of plants exposed to environmental changes. However, environmental changes can also alter the behaviour of LDD vectors: for instance, increasing air temperature may enhance atmospheric instability, thereby altering the turbulent airflow that transports seed and pollen. Here, we investigate temperature effects on wind dispersal in a boreal forest using a 10-year time series of micrometeorological measurements and a Lagrangian stochastic model for particle transport. For a wide range of dispersal and life history types, we found positive relations between air temperature and LDD. This translates into a largely consistent positive effect of +3 degrees C warming on predicted LDD frequencies and spread rates of plants. Relative increases in LDD frequency tend to be higher for heavy-seeded plants, whereas absolute increases in LDD and spread rates are higher for light-seeded plants for which wind is often an important dispersal vector. While these predicted increases are not sufficient to compensate forecasted range losses and environmental changes can alter plant spread in various ways, our results generally suggest that warming can promote wind-driven movements of plant genotypes and populations in boreal forests.

Pollen-mediated gene flow in Chinese commercial fields of glufosinate-resistant canola (Brassica napus)

One of the most pressing issues surrounding transgenic oilseed rape cultivation is the potential impact of GM genes entering neighboring non-GM oilseed rape plants, related weeds or closely related wild relatives. Following the principle of a case-by-case for transgenic crop safety assessment, we designed experiments to study pollen-mediated gene flow from herbicide-resistant transgenic oilseed rape to Chinese commercial varieties. The pollen dispersal was detected as the progeny of recipient plants that were identified as glufosinate resistant. The results indicated that pollen dispersal occurred up to 2000 m and its rate sharply decreased as the distance from the pollen source increased. However, the rate of long-distance pollen dispersal from 33 to 2000 m was less than 0.015% and did not present a gradual decrease, indicating the randomization of dispersal and pollination. Most pollen dispersed within a 4.5 m area around the transgenic pollen source plot, with a maximum of 1.19% at the sampling site of 1.4 m. Wind direction significantly affected the direction and the distance of pollen dispersal, resulting in the more and farther dispersal in southwest direction. However, the number of potentially pollinating bees was not correlated with direction and distance from the pollen source plot, suggesting that bee density and distribution would not contribute to the differences in pollen dispersal among different directions. This paper is the first report on research in a large-scale experiment of pollen-mediated gene flow under the China’s environmental conditions and provided scientific evidence for the effective commercialization of transgenic oilseed rape and its safe management. Our results also provided important data on pollen spatial distribution pattern.

Gene Flow, Risk Assessment and the Environmental Release of Transgenic Plants

The release of genetically modified plants is governed by regulations that aim to provide an assessment of potential impact on the environment. One of the most important components of this risk assessment is an evaluation of the probability of gene flow. In this review, we provide an overview of the current literature on gene flow from transgenic plants, providing a framework of issues for those considering the release of a transgenic plant into the environment. For some plants gene flow from transgenic crops is well documented, and this information is discussed in detail in this review. Mechanisms of gene flow vary from plant species to plant species and range from the possibility of asexual propagation, short- or long-distance pollen dispersal mediated by insects or wind and seed dispersal. Volunteer populations of transgenic plants may occur where seed is inadvertently spread during harvest or commercial distribution. If there are wild populations related to the transgenic crop then hybridization and eventually introgression in the wild may occur, as it has for herbicide resistant transgenic oilseed rape (Brassica napus). Tools to measure the amount of gene flow, experimental data measuring the distance of pollen dispersal, and experiments measuring hybridization and seed survivability are discussed in this review. The various methods that have been proposed to prevent gene flow from genetically modified plants are also described. The current “transgenic traits” in the major crops confer resistance to herbicides and certain insects. Such traits could confer a selective advantage (an increase in fitness) in wild plant populations in some circumstances, were gene flow to occur. However, there is ample evidence that gene flow from crops to related wild species occurred before the development of transgenic crops and this should be taken into account in the risk assessment process.

Pollen dispersal and genetic structure of the tropical tree Dipteryx panamensis in a fragmented Costa Rican landscape

In the face of widespread deforestation, the conservation of rainforest trees relies increasingly on their ability to maintain reproductive processes in fragmented landscapes. Here, we analysed nine microsatellite loci for 218 adults and 325 progeny of the tree Dipteryx panamensis in Costa Rica. Pollen dispersal distances, genetic diversity, genetic structure and spatial autocorrelation were determined for populations in four habitats: continuous forest, forest fragments, pastures adjacent to fragments and isolated pastures. We predicted longer but less frequent pollen movements among increasingly isolated trees. This pattern would lead to lower outcrossing rates for pasture trees, as well as lower genetic diversity and increased structure and spatial autocorrelation among their progeny. Results generally followed these expectations, with the shortest pollen dispersal among continuous forest trees (240 m), moderate distances for fragment (343 m) and adjacent pasture (317 m) populations, and distances of up to 2.3 km in isolated pastures (mean: 557 m). Variance around pollen dispersal estimates also increased with fragmentation, suggesting altered pollination conditions. Outcrossing rates were lower for pasture trees and we found greater spatial autocorrelation and genetic structure among their progeny, as well as a trend towards lower heterozygosity. Paternal reproductive dominance, the pollen contributions from individual fathers, did not vary among habitats, but we did document asymmetric pollen flow between pasture and adjacent fragment populations. We conclude that long-distance pollen dispersal helps maintain gene flow for D. panamensis in this fragmented landscape, but pasture and isolated pasture populations are still at risk of long-term genetic erosion.

A phylogeographical analysis of the range disjunction for foxtail pine (Pinus balfouriana, Pinaceae): the role of Pleistocene glaciation

Biogeographical patterns within the California Floristic Province have been greatly affected by geological and climatic events. Here, we investigate the phylogeography of foxtail pine (Pinus balfouriana) in an effort to date its range disjunction using molecular data and to further our understanding of phylogeographical patterns for plants within the California Floristic Province. The distribution of foxtail pine is characterized by a 500-km disjunction separating populations located in the Klamath Mountains from those in the southern Sierra Nevada. Previous authors suggested that this disjunction occurred approximately 4000-8000 years ago during the Holocene Xerotherm when western North America became warmer and drier. Those dates, however, are inconsistent with the morphological differences that separate regional populations into formally recognized subspecies. Using the coalescent-based isolation with migration model and DNA sequence data from the chloroplast, mitochondrial, and nuclear genomes, we evaluate several hypotheses addressing the timing of this range disjunction and its effects on subsequent patterns of gene flow. Results from all three genomes are largely consistent with Middle to Early Pleistocene divergence dates. Those dates correspond to the Sherwin glaciation, which was the largest Pleistocene glacial episode in the Sierra Nevada. Gene flow, moreover, was only documented using data from the chloroplast genome, suggesting that low levels of long-distance pollen dispersal (N(e)m < 0.5) have occurred since this divergence event. These results are extended to a discussion of the biogeographical development of subalpine forests in California.

Patterns of pollen and seed dispersal in a small, fragmented population of the wind-pollinated tree Araucaria angustifolia in southern Brazil

Paternity analysis based on eight microsatellite loci was used to investigate pollen and seed dispersal patterns of the dioecious wind-pollinated tree, Araucaria angustifolia. The study sites were a 5.4 ha isolated forest fragment and a small tree group situated 1.7 km away, located in Paranalpha State, Brazil. In the forest fragment, 121 males, 99 females, 66 seedlings and 92 juveniles were mapped and genotyped, together with 210 seeds. In the tree group, nine male and two female adults were mapped and genotyped, together with 20 seeds. Paternity analysis within the forest fragment indicated that at least 4% of the seeds, 3% of the seedlings and 7% of the juveniles were fertilized by pollen from trees in the adjacent group, and 6% of the seeds were fertilized by pollen from trees outside these stands. The average pollination distance within the forest fragment was 83 m; when the tree group was included the pollination distance was 2006 m. The average number of effective pollen donors was estimated as 12.6. Mother-trees within the fragment could be assigned to all seedlings and juveniles, suggesting an absence of seed immigration. The distance of seedlings and juveniles from their assigned mother-trees ranged from 0.35 to 291 m (with an average of 83 m). Significant spatial genetic structure among adult trees, seedlings, and juveniles was detected up to 50 m, indicating seed dispersal over a short distance. The effective pollination neighborhood ranged from 0.4 to 3.3 ha. The results suggest that seed dispersal is restricted but that there is long-distance pollen dispersal between the forest fragment and the tree group; thus, the two stands of trees are not isolated.

Extensive Long-Distance Pollen Dispersal in a Fragmented Landscape Maintains Genetic Diversity in White Spruce

Conifers are among the most genetically diverse plants but show the lowest levels of genetic differentiation, even among geographically distant populations. High gene flow among populations may be one of the most important factors in maintaining these genetic patterns. Here, we provide empirical evidence for extensive pollen-mediated gene dispersal between natural stands of a widespread northern temperate/boreal conifer, Picea glauca. We used 6 polymorphic allozyme loci to quantify the proportion of seeds sired by pollen originating from different sources in a landscape fragmented by agriculture in North Central Ontario, Canada. In 7 stands, a small proportion of seeds were sired by self-pollen or neighboring trees but 87.1% (+/-1.7% standard error [SE]) of seeds were sired by pollen from at least 250 to 3000 m away. In 4 single isolated trees, self-fertilization rates were low and more than 96% (+/-1.3% SE) of seeds were sired by immigrant pollen. The average minimum pollen dispersal distance in outcrossed matings was 619 m. These results provide strong evidence that extensive long-distance pollen dispersal plays a primary role in maintaining low genetic differentiation among natural populations of P. glauca and helps maintain genetic diversity and minimize inbreeding in small stands in a fragmented landscape.

Extensive pollen dispersal in a bird‐pollinated shrub, Calothamnus quadrifidus, in a fragmented landscape

Pollen dispersal was investigated in six populations of Calothamnus quadrifidus, a bird-pollinated shrub in the fragmented agricultural region of southern Western Australia. Paternity analysis using six microsatellite loci identified a pollen source within populations for 67% of seedlings, and the remainder were assumed to have arisen from pollen sources outside the populations. Outcrossing was variable, ranging from 5% to 82%, and long-distance pollen dispersal was observed in all populations with up to 43% of pollen sourced from outside the populations over distances of up to 5 km. This extensive pollen immigration was positively associated with population size but not isolation. Comparison of two populations of similar size but different density showed greater internal pollination and less selfing in the denser population, suggesting an influence of density on pollinator behaviour. The study revealed extensive long-distance pollen dispersal for C. quadrifidus within this fragmented agricultural landscape and highlighted the interaction between reserve populations and isolated road verge remnants in maintaining genetic connectivity at the landscape scale.

Population-size effects on seeds and seedlings from fragmented eucalypt populations: implications for seed sourcing for ecological restoration

Ecological restoration of degraded habitats is a major conservation activity requiring the collection of large amounts of native seed. Seed production and the genetic quality of seed may be influenced by properties of the source population, such as population size and fragmentation, potentially having an impact on restoration goals. We assessed the population-size effects on seed production and seedling performance in two Western Australian wheatbelt eucalypts, Eucalyptus salmonophloia F.Muell. and E. salubris F.Muell. Both species were historically widespread and dominant, but, as a consequence of land-clearing for agriculture, now exist as small, highly fragmented populations throughout the western half of their range. Given their former importance in the landscape, these species will be critical in ecological restoration of the region. We assessed small (n = 6–12) and large (n &gt; 200) remnant populations in a highly fragmented landscape and compared these to large unfragmented populations. Seed number per capsule was dependent on population size and fragmentation for E. salubris, but not for E. salmonophloia. Large, unfragmented populations of E. salubris produced more than twice the number of seeds per capsule (mean = 2.95) than small and/or fragmented populations. However, seed germination, seed weight, seedling survival and seedling vigour to 1 year were independent of population size or fragmentation in both species. Our results suggest that reduced population size and increased fragmentation can negatively affect pollen quantity and/or quality, thereby limiting seed production, although no fitness effects were observed post-seed maturation. We suggest that the relative absence of post-seed maturation fitness effects in these small fragmented populations are a consequence of (1) wide outcrossing resulting from long-distance dispersal of pollen by highly mobile birds among fragmented populations and/or (2) efficient pre- or post-zygotic selection against more homozygous zygotes within fruits so that only relatively outbred seeds mature. The consequences on seed collection for ecological restoration of reduced population size and increased fragmentation for these eucalypts may be fewer seeds for the same collecting effort, but no apparent fitness effects of mature seeds. However, caution should be exercised when harvesting seed from these smaller populations, as over-harvesting may have an impact on recruitment and hence long-term persistence.

Using SSR markers to determine the population genetic structure of wild apricot (Prunus armeniaca L.) in the Ily Valley of West China

Genetic structure of three wild populations (Xinyuan, Gongliu and Daxigou) of apricot in the Ily Valley, Xinjiang Uygur Autonomous Region of China, was investigated with microsatellite (simple sequence repeat, SSR) markers. The higher polymorphism and greater transportability of these markers between Prunus species proved SSR markers were much efficient for conducting genetic diversity studies in wild apricot. Nei's gene diversity (He) and Shannon's index of diversity (I) were 0.287 and 0.458, respectively. This indicated that the wild apricot in the Ily Valley still maintained a relatively high level of diversity. The Gst of 0.137 and Fst of 0.164 revealed that genetic variation mainly resided among individuals within populations (83.6–86.3%). Population differentiation could also be found according to the distribution of SSR alleles between the populations. Mantel test showed the genetic distance between populations was significantly correlated to the geographical distance. The modest amount of gene flow (2.684) would reduce the disjunction between wild apricots. The long-distance dispersal of pollen by insects was probably the main way of gene flow between populations. Based on the study of population genetic structure, an effective conservation strategy of the species was discussed.

Ancestral chloroplast polymorphism and historical secondary contact in a broad hybrid zone ofAesculus(Sapindaceae)

Knowledge regarding the origin and maintenance of hybrid zones is critical for understanding the evolutionary outcomes of natural hybridization. To evaluate the contribution of historical contact vs. long-distance gene flow in the formation of a broad hybrid zone in central and northern Georgia that involves Aesculus pavia, A. sylvatica, and A. flava, three cpDNA regions (matK, trnD-trnT, and trnH-trnK) were analyzed. The maternal inheritance of cpDNA in Aesculus was confirmed via sequencing of matK from progeny of controlled crosses. Restriction site analyses identified 21 unique haplotypes among 248 individuals representing 29 populations from parental species and hybrids. Haplotypes were sequenced for all cpDNA regions. Restriction site and sequence data were subjected to phylogeographic and population genetic analyses. Considerable cpDNA variation was detected in the hybrid zone, as well as ancestral cpDNA polymorphism; furthermore, the distribution of haplotypes indicates limited interpopulation gene flow via seeds. The genealogy and structure of genetic variation further support the historical presence of A. pavia in the Piedmont, although they are at present locally extinct. In conjunction with previous allozyme studies, the cpDNA data suggest that the hybrid zone originated through historical local gene flow, yet is maintained by periodic long-distance pollen dispersal.

Pollen flow in the wildservice tree, Sorbus torminalis (L.) Crantz. II. Pollen dispersal and heterogeneity in mating success inferred from parent–offspring analysis

Knowing the extent of gene movements from parents to offspring is essential to understand the potential of a species to adapt rapidly to a changing environment, and to design appropriate conservation strategies. In this study, we develop a nonlinear statistical model to jointly estimate the pollen dispersal kernel and the heterogeneity in fecundity among phenotypically or environmentally defined groups of males. This model uses genotype data from a sample of fruiting plants, a sample of seeds harvested on each of these plants, and all males within a circumscribed area. We apply this model to a scattered, entomophilous woody species, Sorbus torminalis (L.) Crantz, within a natural population covering more than 470 ha. We estimate a high heterogeneity in male fecundity among ecological groups, both due to phenotype (size of the trees and flowering intensity) and landscape factors (stand density within the neighbourhood). We also show that fat-tailed kernels are the most appropriate to depict the important abilities of long-distance pollen dispersal for this species. Finally, our results reveal that the spatial position of a male with respect to females affects as much its mating success as ecological determinants of male fecundity. Our study thus stresses the interest to account for the dispersal kernel when estimating heterogeneity in male fecundity, and reciprocally.

To self, or not to self… A review of outcrossing and pollen-mediated gene flow in neotropical trees

Despite the typically low population densities and animal-mediated pollination of tropical forest trees, outcrossing and long-distance pollen dispersal are the norm. We reviewed the genetic literature on mating systems and pollen dispersal for neotropical trees to identify the ecological and phylogenetic correlates. The 36 studies surveyed found >90% outcrossed mating for 45 hermaphroditic or monoecious species. Self-fertilization rates varied inversely with population density and showed phylogenetic and geographic trends. The few direct measures of pollen flow (N=11 studies) suggest that pollen dispersal is widespread among low-density tropical trees, ranging from a mean of 200 m to over 19 km for species pollinated by small insects or bats. Future research needs to examine (1) the effect of inbreeding depression on observed outcrossing rates, (2) pollen dispersal in a wide range of pollination syndromes and ecological classes, (3) and the range of variation of mating system expression at different hierarchical levels, including individual, seasonal, population, ecological, landscape and range wide.

Parentage versus two‐generation analyses for estimating pollen‐mediated gene flow in plant populations

Assessment of contemporary pollen-mediated gene flow in plants is important for various aspects of plant population biology, genetic conservation and breeding. Here, through simulations we compare the two alternative approaches for measuring pollen-mediated gene flow: (i) the NEIGHBORHOOD model--a representative of parentage analyses, and (ii) the recently developed TWOGENER analysis of pollen pool structure. We investigate their properties in estimating the effective number of pollen parents (N(ep)) and the mean pollen dispersal distance (delta). We demonstrate that both methods provide very congruent estimates of N(ep) and delta, when the methods' assumptions considering the shape of pollen dispersal curve and the mating system follow those used in data simulations, although the NEIGHBORHOOD model exhibits generally lower variances of the estimates. The violations of the assumptions, especially increased selfing or long-distance pollen dispersal, affect the two methods to a different degree; however, they are still capable to provide comparable estimates of N(ep). The NEIGHBORHOOD model inherently allows to estimate both self-fertilization and outcrossing due to the long-distance pollen dispersal; however, the TWOGENER method is particularly sensitive to inflated selfing levels, which in turn may confound and suppress the effects of distant pollen movement. As a solution we demonstrate that in case of TWOGENER it is possible to extract the fraction of intraclass correlation that results from outcrossing only, which seems to be very relevant for measuring pollen-mediated gene flow. The two approaches differ in estimation precision and experimental efforts but they seem to be complementary depending on the main research focus and type of a population studied.

Viability and sunlight sensitivity of oak pollen and its implications for pollen-mediated gene flow

Pollen-mediated gene flow and the male reproductive success of wind-pollinated trees depend on the initial viability of the pollen and the changes that occur in its viability during transport in the atmosphere. The viability of Quercus robur pollen was determined before and during exposure to sunlight by in vitro germination and the fluorescein diacetate reaction (FCR) in 2002 and 2003, respectively. These experiments allowed us to calculate initial pollen viability and pollen sensitivity to sunlight. The germination test revealed a lower initial pollen viability (25–65%) than the FCR (53–92%). Following 9.5 h of irradiation the viability was reduced to 75–100% as determined by the in vitro germination test or to 40–70% as determined by the FCR. The actual values of initial pollen viability and pollen sensitivity to sunlight were used to define a range of values for modelling pollen dispersal using the mesoscale meteorological model METRAS. The deposition patterns of viable pollen varied by as much as a factor of 14 by changing the viability parameters in the range of the observed values. This suggests significant differences in male reproductive success. Variations in initial pollen viability have stronger effects on the gene-flow pattern than do variations in pollen sensitivity to sunlight. In particular, pollen distribution throughout the local environment is shaped by the initial pollen viability, while pollen sensitivity to sunlight mainly influences long-distance pollen dispersal.

SELECTION AND DISPERSAL IN A MULTISPECIES OAK HYBRID ZONE

The four western North American red oak species (Quercus wislizeni, Q. parvula, Q. agrifolia, and Q. kelloggii) are known to produce hybrid products in all interspecific combinations. However, it is unknown whether hybrids are transitory resulting from interspecific gene flow or whether they are maintained through extrinsic selection. Here, we examine cryptic hybrid structure in Q. wislizeni through a broad region including contact and isolation from three other western North American red oaks using amplified fragment length polymorphism molecular markers. All four species were simultaneously detected in the genetic background of individuals morphologically assigned to Q. wislizeni, although the contribution of Q. kelloggii was minor. In some cases, introgression was detected well outside the region of sympatry with one of the parental species. Molecular structure at the individual level indicated this was due to long-distance pollen dispersal and not to local extinction of parental species. Species admixture proportions were correlated with climatic variables and greater proportions of Q. agrifolia and Q. parvula were present in the genetic background of Q. wislizeni in sites with cooler and more humid summers, corresponding with habitat preferences of the parental species. Partial Mantel tests indicated that climate was more important than distance from pollen source in this association. Despite high levels of introgression, species integrity was maintained in some populations in close proximity to the other species, providing further support to environmental selection in determining population genetic structure. Thus, the contribution of species mixtures to population genetic structure varies across the landscape according to availability of pollen, but more importantly to varying environmental selection pressures that produce a complex pattern of hybrid and pure gene pools.

SELECTION AND DISPERSAL IN A MULTISPECIES OAK HYBRID ZONE

The four western North American red oak species (Quercus wislizeni, Q. parvula, Q. agrifolia, and Q. kelloggii) are known to produce hybrid products in all interspecific combinations. However, it is unknown whether hybrids are transitory resulting from interspecific gene flow or whether they are maintained through extrinsic selection. Here, we examine cryptic hybrid structure in Q. wislizeni through a broad region including contact and isolation from three other western North American red oaks using amplified fragment length polymorphism molecular markers. All four species were simultaneously detected in the genetic background of individuals morphologically assigned to Q. wislizeni, although the contribution of Q. kelloggii was minor. In some cases, introgression was detected well outside the region of sympatry with one of the parental species. Molecular structure at the individual level indicated this was due to long-distance pollen dispersal and not to local extinction of parental species. Species admixture proportions were correlated with climatic variables and greater proportions of Q. agrifolia and Q. parvula were present in the genetic background of Q. wislizeni in sites with cooler and more humid summers, corresponding with habitat preferences of the parental species. Partial Mantel tests indicated that climate was more important than distance from pollen source in this association. Despite high levels of introgression, species integrity was maintained in some populations in close proximity to the other species, providing further support to environmental selection in determining population genetic structure. Thus, the contribution of species mixtures to population genetic structure varies across the landscape according to availability of pollen, but more importantly to varying environmental selection pressures that produce a complex pattern of hybrid and pure gene pools.