The role of anthropogenic dispersal in shaping the distribution and genetic composition of a widespread North American tree species.

Dispersal and colonization are among the most important ecological processes for species persistence as they allow species to track changing environmental conditions. During the last glacial maximum (LGM), many cold-intolerant Northern Hemisphere plants retreated to southern glacial refugia. During subsequent warming periods, these species expanded their ranges northward. Interestingly, some tree species with limited seed dispersal migrated considerable distances after the LGM ~19,000 years before present (YBP). It has been hypothesized that indigenous peoples may have dispersed valued species, in some cases beyond the southern limits of the Laurentide Ice Sheet. To investigate this question, we employed a molecular genetics approach on a widespread North American understory tree species whose fruit was valued by indigenous peoples. Twenty putative anthropogenic (near pre-Columbian habitations) and 62 wild populations of Asimina triloba (pawpaw), which produces the largest edible fruit of any North American tree, were genetically assayed with nine microsatellite loci. Putative anthropogenic populations were characterized by reduced genetic diversity and greater excess heterozygosity relative to wild populations. Anthropogenic populations in regions that were glaciated during the LGM had profiles consistent with founder effects and reduced gene flow, and shared rare alleles with wild populations hundreds of kilometers away (mean = 723 km). Some of the most compelling evidence for human-mediated dispersal is that putative anthropogenic and wild populations sharing rare alleles were separated by significantly greater distances (mean = 695 km) than wild populations sharing rare alleles (mean = 607 km; p = .014). Collectively, the genetic data suggest that long-distance dispersal played an important role in the distribution of pawpaw and is consistent with the hypothesized role of indigenous peoples.

Selfing rates vary with floral display, pollinator visitation and plant density in natural populations of Mimulus ringens.

Variation in selfing rates within and among populations of hermaphroditic flowering plants can strongly influence the evolution of reproductive strategies and the genetic structure of populations. This intraspecific variation in mating patterns may reflect both genetic and ecological factors, but the relative importance of these factors remains poorly understood. Here, we explore how selfing in 13 natural populations of the perennial wildflower Mimulus ringens is influenced by (a) pollinator visitation, an ecological factor, and (b) floral display, a trait with a genetic component that also responds to environmental variation. We also explore whether genetically based floral traits, including herkogamy, affect selfing. We found substantial variation among populations in selfing rate (0.13-0.55). Selfing increased strongly and significantly with floral display, among as well as within populations. Selfing also increased at sites with lower pollinator visitation and low plant density. However, selfing was not correlated with floral morphology. Overall, these results suggest that pollinator visitation and floral display, two factors that interact to affect geitonogamous pollinator movements, can influence the selfing rate. This study identifies mechanisms that may play a role in maintaining selfing rate variation among populations.

Edge effects and mating patterns in a bumblebee-pollinated plant.

Researchers have long assumed that plant spatial location influences plant reproductive success and pollinator foraging behaviour. For example, many flowering plant populations have small, linear or irregular shapes that increase the proportion of plants on the edge, which may reduce mating opportunities through both male and female function. Additionally, plants that rely on pollinators may be particularly vulnerable to edge effects if those pollinators exhibit restricted foraging and pollen carryover is limited. To explore the effects of spatial location (edge vs. interior) on siring success, seed production, pollinator foraging patterns and pollen-mediated gene dispersal, we established a square experimental array of 49 Mimulus ringens (monkeyflower) plants. We observed foraging patterns of pollinating bumblebees and used paternity analysis to quantify male and female reproductive success and mate diversity for plants on the edge versus interior. We found no significant differences between edge and interior plants in the number of seeds sired, mothered or the number of sires per fruit. However, we found strong differences in pollinator behaviour based on plant location, including 15 % lower per flower visitation rates and substantially longer interplant moves for edge plants. This translated into 40 % greater pollen-mediated gene dispersal for edge than for interior plants. Overall, our results suggest that edge effects are not as strong as is commonly assumed, and that different plant reproduction parameters respond to spatial location independently.

Hermaphroditism promotes mate diversity in flowering plants.

PREMISE: Genetically diverse sibships are thought to increase parental fitness through a reduction in the intensity of sib competition, and through increased opportunities for seedling establishment in spatially or temporally heterogeneous environments. Nearly all research on mate diversity in flowering plants has focused on the number of fathers siring seeds within a fruit or on a maternal plant. Yet as hermaphrodites, plants can also accrue mate diversity by siring offspring on several pollen recipients in a population. Here we explore whether mate composition overlaps between the dual sex functions, and discuss the implications for plant reproductive success. METHODS: We established an experimental population of 49 Mimulus ringens (monkeyflower) plants, each trimmed to a single flower. Following pollination by wild bees, we quantified mate composition for each flower through both paternal and maternal function. Parentage was successfully assigned to 240 progeny, 98% of the sampled seeds. RESULTS: Comparison of mate composition between male and female function revealed high mate diversity, with almost no outcross mates shared between the two sexual functions of the same flower. CONCLUSIONS: Dual sex roles contribute to a near doubling of mate diversity in our experimental population of Mimulus ringens. This finding may help explain the maintenance of hermaphroditism under conditions that would otherwise favor the evolution of separate sexes.

Phylogeography of the Neotropical epiphytic orchid, Brassavola nodosa: evidence for a secondary contact zone in northwestern Costa Rica.

Spatial patterns of genetic variation can reveal otherwise cryptic evolutionary and landscape processes. In northwestern Costa Rica, an approximately concordant genetic discontinuity occurs among populations of several plant species. We conducted phylogeographic analyses of an epiphytic orchid, Brassavola nodosa, to test for genetic discontinuity and to explore its underlying causes. We genotyped 18 populations with 19 nuclear loci and two non-coding chloroplast sequence regions. We estimated genetic diversity and structure, relative importance of pollen and seed dispersal, and divergence time to understand how genetic diversity was spatially partitioned. Nuclear genetic diversity was high with little differentiation among populations (GSTn = 0.065). In contrast, chloroplast haplotypes were highly structured (GSTc = 0.570) and reveal a discontinuity between northwestern and southeastern populations within Costa Rica. Haplotype differences suggest two formerly isolated lineages that diverged ~10,000-100,000 YBP. Haplotype mixing and greater genetic diversity occur in an intermediate transition zone. Patterns of nuclear and chloroplast data were consistent. Different levels of genetic differentiation for the two genomes reflect the relative effectiveness of biotic versus abiotic dispersers of pollen and seeds, respectively. Isolation of the two lineages likely resulted from the complex environmental and geophysical history of the region. Our results suggest a recent cryptic seed dispersal barrier and/or zone of secondary contact. We hypothesize that powerful northeasterly trade winds hinder movement of wind-borne seeds between the two regions, while the multi-directional dispersal of pollen by strong-flying sphinx moths resulted in lower differentiation of nuclear loci.

Reproductive consequences of variation in flowering phenology in the dry forest tree Enterolobium cyclocarpum in Guanacaste, Costa Rica.

PREMISE OF THE STUDY: Flowering initiation, duration and magnitude, and degree of flowering synchrony within a population can affect the reproductive fitness of individuals. We examined the flowering phenology within a population of the tropical dry forest Guanacaste tree (Enterolobium cyclocarpum) to gauge the impact of phenological variation among trees on fruit production and progeny vigor. METHODS: We monitored the flowering phenology of 93 trees weekly during 2005, 2006, and 2007, using a scale based on the percentage of the crown with open flowers. We also monitored fruit production for each tree in 2005, 2006, 2007, and 2008. Finally, we evaluated the relationship between phenological variation and progeny performance. KEY RESULTS: Ten measures of flowering phenology and synchrony among flowering trees, based on the number of weeks when anthesis of the crown exceeded 50%, were used to develop four phenological profiles. These profiles were correlated with significant differences in fruit production and progeny vigor. Trees with flowers in >50% of their crown for at least 2 weeks produced more fruits and more vigorous progeny than trees with other profiles. Trees also tended to produce the same phenological profile among years than predicted by chance. CONCLUSIONS: Guanacaste trees vary significantly in the initiation of anthesis, duration and magnitude of flowering, and degree of synchrony among trees. Trees also tend to maintain the same flowering profile among years. Finally, the flowering behavior of E. cyclocarpum leads to significant differences in fruit and seed production, germination, and early progeny growth.

Effects of pollination and postpollination processes on selfing rate in Mimulus ringens.

PREMISE OF THE STUDY: Selfing rates vary widely within and among populations of self-compatible flowering plants. This variation is often attributed to differences in the amount and timing of self and outcross pollen deposition on stigmas, as well as to the influence of postpollination mechanisms that control fertilization success. This study explores the relative importance of pollination and postpollination processes in determining selfing rates in monkeyflower, Mimulus ringens. METHODS: We hand-pollinated flowers on 17 unrelated mothers with pollen from one of three experimental treatments intended to replicate field conditions: (1) simultaneous deposition of 50% self pollen and 50% outcross pollen from 5 unrelated donors; (2) self pollen followed 15 min later by application of an equal amount of outcross pollen from five unrelated donors; and (3) outcross pollen from 5 unrelated donors followed 15 min later by application of an equal amount of self pollen. We genotyped 757 progeny at 8 polymorphic microsatellite loci and used paternity exclusion to determine whether each seedling was selfed or outcrossed. KEY RESULTS: When self and outcross pollen arrived simultaneously, and when self pollen arrived 15 min prior to outcross pollen, the observed proportions of self and outcross progeny did not deviate from the expected 1:1 ratio. However, when outcross pollen was applied 15 min prior to self pollen, there was a significant excess of outcross progeny. CONCLUSIONS: Selfing rate in Mimulus ringens is influenced by small differences in the timing of pollen arrival, but not by nonrandom postpollination sorting.

Characterization of microsatellite loci for an Australian epiphytic orchid, Dendrobium calamiforme, using Illumina sequencing.

PREMISE OF THE STUDY: Microsatellite loci were developed for the epiphytic pencil orchid Dendrobium calamiforme for population genetic and phylogeographic investigation of this Australian taxon. METHODS AND RESULTS: Nineteen microsatellite loci were identified from an Illumina paired-end shotgun library of D. calamiforme. Polymorphism and genetic diversity were assessed in 24 individuals from five populations separated by a maximum distance of ∼80 km. All loci were polymorphic with two to 14 alleles per locus, expected heterozygosity ranging from 0.486 to 0.902, and probability of identity values ranging from 0.018 to 0.380. CONCLUSIONS: These novel markers will serve as valuable tools for investigation of levels of genetic diversity as well as patterns of gene flow, genetic structure, and phylogeographic history.

Genetic and morphological contrasts between wild and anthropogenic populations of Agave parryi var. huachucensis in south-eastern Arizona.

BACKGROUND AND AIMS: At least seven species of Agave, including A. parryi, were cultivated prehistorically in Arizona, serving as important sources of food and fibre. Many relict populations from ancient cultivation remain in the modern landscape, offering a unique opportunity to study pre-Columbian plant manipulation practices. This study examined genetic and morphological variation in six A. p. var. huachucensis populations of unknown origin to compare them with previous work on A. parryi populations of known origin, to infer their cultivation history and to determine whether artificial selection is evident in populations potentially managed by early agriculturalists. METHODS: Six A. p. var. huachucensis and 17 A. parryi populations were sampled, and morphometric, allozyme and microsatellite data were used to compare morphology and genetic structure in purportedly anthropogenic and wild populations, as well as in the two taxa. Analysis of molecular variance and Bayesian clustering were performed to partition variation associated with taxonomic identity and hypothesized evolutionary history, to highlight patterns of similarity among populations and to identify potential wild sources for the planting stock. KEY RESULTS: A p. var. huachucensis and A. parryi populations differed significantly both morphologically and genetically. Like A. parryi, wild A. p. var. huachucensis populations were more genetically diverse than the inferred anthropogenic populations, with greater expected heterozygosity, percentage of polymorphic loci and number of alleles. Inferred anthropogenic populations exhibited many traits indicative of past active cultivation: greater morphological uniformity, fixed heterozygosity for several loci (non-existent in wild populations), fewer multilocus genotypes and strong differentiation among populations. CONCLUSIONS: Where archaeological information is lacking, the genetic signature of many Agave populations in Arizona can be used to infer their evolutionary history and to identify potentially fruitful sites for archaeological investigation of ancient settlements and cultivation practices. The same approach can clearly be adopted for other species in similar situations.

Relative importance of pollen and seed dispersal across a Neotropical mountain landscape for an epiphytic orchid.

Populations of many species are isolated within narrow elevation bands of Neotropical mountain habitat, and how well dispersal maintains genetic connectivity is unknown. We asked whether genetic structure of an epiphytic orchid, Epidendrum firmum, corresponds to gaps between Costa Rican mountain ranges, and how these gaps influence pollen and seed flow. We predicted that significant genetic structure exists among mountain ranges due to different colonization histories and limited gene flow. Furthermore, we predicted that pollen movement contributes more to gene flow than seeds because seeds are released into strong winds perpendicular to the narrow northwest-southeast species distribution, while the likely pollinators are strong fliers. Individuals from 12 populations and three mountain ranges were genotyped with nuclear microsatellites (nDNA) and chloroplast sequences (cpDNA). Genetic diversity was high for both markers, while nDNA genetic structure was low (FSTn  = 0.020) and cpDNA structure was moderate (FSTc  = 0.443). Significant cpDNA barriers occurred within and among mountain ranges, but nDNA barriers were not significant after accounting for geographic distance. Consistent with these contrasting patterns of genetic structure, pollen contributes substantially more to gene flow among populations than seed (mp /ms  = 46). Pollinators mediated extensive gene flow, eroding nDNA colonization footprints, while seed flow was comparatively limited, possibly due to directional prevailing winds across linearly distributed populations. Dispersal traits alone may not accurately inform predictions about gene flow or genetic structure, supporting the need for research into the potentially crucial role of pollinators and landscape context in gene flow among isolated populations.

Highly diverse and spatially heterogeneous mycorrhizal symbiosis in a rare epiphyte is unrelated to broad biogeographic or environmental features.

Symbiotic interactions are common in nature. In dynamic or degraded environments, the ability to associate with multiple partners (i.e. broad specificity) may enable species to persist through fluctuations in the availability of any particular partner. Understanding how species interactions vary across landscapes is necessary to anticipate direct and indirect consequences of environmental degradation on species conservation. We asked whether mycorrhizal symbiosis by populations of a rare epiphytic orchid (Epidendrum firmum) is related to geographic or environmental heterogeneity. The latter would suggest that interactions are governed by environmental conditions rather than historic isolation of populations and/or mycorrhizal fungi. We used DNA-based methods to identify mycorrhizal fungi from eleven E. firmum populations in Costa Rica. We used molecular and phylogenetic analyses to compare associations. Epidendrum firmum exhibited broad specificity, associating with diverse mycorrhizal fungi, including six Tulasnellaceae molecular operational taxonomic units (MOTUs), five Sebacinales MOTUs and others. Notably, diverse mycorrhizal symbioses formed in disturbed pasture and roadside habitats. Mycorrhizal fungi exhibited significant similarity within populations (spatial and phylogenetic autocorrelation) and significant differences among populations (phylogenetic community dissimilarity). However, mycorrhizal symbioses were not significantly associated with biogeographic or environmental features. Such unexpected heterogeneity among populations may result from complex combinations of fine-scale environmental factors and macro-evolutionary patterns of change in mycorrhizal specificity. Thus, E. firmum exhibits broad specificity and the potential for opportunistic associations with diverse fungi. We suggest that these characteristics could confer symbiotic assurance when mycorrhizal fungi are stochastically available, which may be crucial in dynamic or disturbed habitats such as tropical forest canopies.

Genetic inference of epiphytic orchid colonization; it may only take one.

Colonization of vacant habitat is a fundamental ecological process that affects the ability of species to persist and undergo range modifications in continually shifting landscapes. Thus, understanding factors that affect and limit colonization has important ecological and conservation implications. Epiphytic orchids are increasingly threatened by various factors, including anthropogenic habitat disturbance. As cleared areas (e.g. pastures) are recolonized by suitable host trees, the establishment and genetic composition of epiphytic orchid populations are likely a function of their colonization patterns. We used genetic analyses to infer the prevailing colonization pattern of the epiphytic orchid, Brassavola nodosa. Samples from three populations (i.e. individuals within a tree) from each of five pastures in the dry forest of Costa Rica were genotyped with neutral nuclear and chloroplast markers. Spatial autocorrelation and hierarchical genetic structure analyses were used to assess the relatedness of individuals within populations, among populations within pastures and among populations in different pastures. The results showed significant relatedness within populations (mean r=0.166) and significant but lower relatedness among populations within a pasture (mean r=0.058). Our data suggest that colonization of available habitats is by few individuals with subsequent population expansion resulting from in situ reproduction, and that individuals within a tree are not a random sample of the regional seed pool. Furthermore, populations within a pasture were likely colonized by seeds produced by founders of a neighbouring population within that pasture. These results have important ramifications for understanding conservation measures needed for this species and other epiphytic orchids.

Characterization of 42 polymorphic microsatellite loci in Mimulus ringens (Phrymaceae) using Illumina sequencing.

PREMISE OF THE STUDY: Microsatellite markers were isolated and characterized in Mimulus ringens (Phrymaceae), a herbaceous wetland perennial, to facilitate studies of mating patterns and population genetic structure. • METHODS AND RESULTS: A total of 42 polymorphic loci were identified from a sample of 24 individuals from a single population in Ohio, USA. The number of alleles per locus ranged from two to nine, and median observed heterozygosity was 0.435. • CONCLUSIONS: This large number of polymorphic loci will enable researchers to quantify male fitness, patterns of multiple paternity, selfing, and biparental inbreeding in large natural populations of this species. These markers will also permit detailed study of fine-scale patterns of genetic structure.

Microsatellite primers for the neotropical epiphyte Epidendrum firmum (Orchidaceae).

PREMISE OF THE STUDY: Ten microsatellite loci were isolated and characterized for the neotropical epiphytic orchid Epidendrum firmum to examine levels of genetic diversity and genetic structure at multiple spatial scales. METHODS AND RESULTS: We screened loci in 12-25 individuals from each of two populations in Costa Rica and identified 10 polymorphic loci. The number of alleles per locus ranged from one to 15 while observed heterozygosity for polymorphic loci ranged from 0.360 to 0.960. CONCLUSIONS: Primers for these informative genetic markers will be useful for quantifying genetic diversity, spatial genetic structure, and gene flow in E. firmum.

Evaluating the utility of microsatellites for investigations of autopolyploid taxa.

Autopolyploid taxa present numerous challenges for population genetic analyses due to difficulties determining allele dosage. Dosage ambiguity hinders accurate assessment of allele frequencies, multilocus genotypes (MLGTs), as well as levels and patterns of clonality. The pervasiveness of polyploidy in the evolutionary history of plant taxa makes this a recurring problem. Whereas diploidization of loci may occur over time, duplication of at least some loci is still frequently evident. Fortunately, with high-quality allozyme gels, it is possible to accurately infer allele dosage and, thus, determine exact MLGTs. However, accurately assessing dosage of microsatellite peaks is nearly impossible when studying wild populations with a large number of alleles per locus. Even if precise knowledge of genotypes is not required, for comparable numbers of alleles per locus and loci, the number of "phenotypes" is always lower with microsatellites than allozymes due to the inability to assess allele dosage. Microsatellite loci typically have more alleles per locus relative to allozymes although fewer loci are generally employed. Here, we present a mathematical model for comparing the relative utility of simple sequence repeat (SSR) versus allozyme markers to discriminate MLGTs. For example, the average plant allozyme study (2.6 alleles per locus, 10 polymorphic loci) has better discriminating power than SSR markers with 10 alleles at each of 3 loci, 9 alleles at 4 loci, 6 alleles at 5 loci, 5 alleles at 6 loci, and 4 alleles at 8 loci, demonstrating the value of assessing the relative discriminating power of these markers.

Inferring ancient Agave cultivation practices from contemporary genetic patterns.

Several Agave species have played an important ethnobotanical role since prehistory in Mesoamerica and semiarid areas to the north, including central Arizona. We examined genetic variation in relict Agave parryi populations northeast of the Mogollon Rim in Arizona, remnants from anthropogenic manipulation over 600 years ago. We used both allozymes and microsatellites to compare genetic variability and structure in anthropogenically manipulated populations with putative wild populations, to assess whether they were actively cultivated or the result of inadvertent manipulation, and to determine probable source locations for anthropogenic populations. Wild populations were more genetically diverse than anthropogenic populations, with greater expected heterozygosity, polymorphic loci, effective number of alleles and allelic richness. Anthropogenic populations exhibited many traits indicative of past active cultivation: fixed heterozygosity for several loci in all populations (nonexistent in wild populations); fewer multilocus genotypes, which differed by fewer alleles; and greater differentiation among populations than was characteristic of wild populations. Furthermore, manipulated populations date from a period when changes in the cultural context may have favoured active cultivation near dwellings. Patterns of genetic similarity among populations suggest a complex anthropogenic history. Anthropogenic populations were not simply derived from the closest wild A. parryi stock; instead they evidently came from more distant, often more diverse, wild populations, perhaps obtained through trade networks in existence at the time of cultivation.

The impact of landscape disturbance on spatial genetic structure in the Guanacaste tree, Enterolobium cyclocarpum (Fabaceae).

We examined spatial genetic structure (SGS) in Enterolobium cyclocarpum (the Guanacaste tree), a dominant tree of Central American dry forests in 4 sites in Guanacaste Province, Costa Rica. In disturbed dry forest sites (e.g., pastures), E. cyclocarpum is primarily dispersed by cattle and horses, whose movements are restricted by pasture boundaries. The study sites varied in tree densities and disturbance. Allozyme analyses of adult trees demonstrated significant levels of SGS in 3 of 4 sites. SGS was primarily due to clusters of young adults located along seasonal streams, rocky areas, and in abandoned pastures. SGS was highest in the first distance class in the least disturbed population, which also had the lowest density of large adults. Low, but significant SGS characterized the site with the highest number of large adults located in individual pastures. The semiurban site, had no clusters of young adults and, probably as a result, failed to exhibit SGS. Our results demonstrate that disturbance can strongly influence SGS patterns and are consistent with a landscape model in which the location of potential recruitment sites, restricted seed disperser movements, and the number and location of maternal individuals dictate the level and pattern of SGS.

Spatial genetic structure of the southeastern North American endemic, Ceratiola ericoides (Empetraceae).

Understanding the spatial distribution of genetic diversity (i.e., spatial genetic structure [SGS]) within plant populations can elucidate mechanisms of seed dispersal and patterns of recruitment that may play an important role in shaping the demography and spatial distribution of individuals in subsequent generations. Here we investigate the SGS of allozyme diversity in 2 populations of the southeastern North American endemic shrub, Ceratiola ericoides. The data suggest that the 2 populations have similar patterns of SGS at distances of 0-45 m that likely reflect the isolation by distance (IBD) model of seed dispersal. However, at distances >or=50 m, the pattern of SGS differs substantially between the 2 populations. Whereas one population continues to reflect the classical IBD pattern, the second population shows a marked increase in autocorrelation coefficient (r) values at 50-75 m. Furthermore, r values at these distances are as much as 33% higher than at 0-5 m where the highest r value would be predicted by IBD. A likely explanation is the differing frequencies of 2 fruit morphologies in these populations and the greater role that birds play in seed dispersal in the second population.

Isolation and characterization of microsatellite loci in the Guanacaste tree, Enterolobium cyclocarpum.

We isolated nine microsatellite loci from the Guanacaste tree (Enterolobium cyclocarpum) and optimized them for future research on breeding populations of this species. Loci were screened across 53 individuals from one population and were shown to be variable with the number of alleles per locus ranging from five to 15. Polymorphic information content ranged from 0.420 to 0.900 and observed heterozygosity ranged from 0.547 to 0.906.

Genetic insights into the biogeography of the southeastern North American endemic, Ceratiola ericoides (Empetraceae).

The southeastern United States harbors an unusually large number of endemic plant taxa, which may reflect the refugial nature of the region during Pleistocene glacial maxima. Understanding the genetic diversity and structure of extant plant taxa can provide insights into the biogeographical processes that shaped them genetically. Here, we investigate the levels and partitioning of allozyme diversity in the southeastern North American endemic, Ceratiola ericoides, which displayed greater genetic variation and structure than other endemics. Central Florida populations represent a center of genetic diversity, whereas South Carolina and Georgia Fall Line sandhill populations have a subset of the Central Florida genetic diversity and may be relicts of a once continuous distribution. This much broader, continuous distribution throughout the southeastern United States occurred during glacial maxima when the scrub habitat, dominated by C. ericoides, expanded considerably owing to drier climatic conditions. Georgia Coastal Plain populations appear to have been independently founded more recently by propagules from Central Florida and the Fall Line sandhills because they have an even more limited subset of genetic diversity and greater genetic heterogeneity among populations. Since their establishment, coastal plain populations appear to have had little, if any, gene exchange among each other or with the relatively proximate Fall Line sandhill populations. These data underscore the importance of understanding the genetic composition and historical biogeography of species before intelligent management or restoration decisions can be made regarding their preservation.