Molecular phylogenetics of Houstonia (Rubiaceae): descending aneuploidy and breeding system evolution in the radiation of the lineage across North America

Nuclear and chloroplast DNA sequence variation was used to infer evolutionary relationships within and among members of Houstonia (Rubiaceae) and other closely related genera in North America. Sequences from the internal transcribed spacer (ITS) of the nrDNA and a cpDNA intron in the trnL gene were used to reconstruct phylogenetic relationships of 30 species of Houstonia and closely related genera. The data suggest that the North American species of Houstonia are not monophyletic, but belong to the same lineage as Stenaria. The radiation of this lineage has been accompanied by changes in the basic chromosome number of the major clades through descending aneuploidy. This loss of chromosomes was also associated with northward colonization of North America. However, other characters, such as an annual versus perennial habit, heterostylous and homostylous breeding systems, and the evolution of self fertilization, seem to be labile throughout the lineage, originating multiple times throughout the evolutionary history of the lineage.     

The genetic legacy of polyploid Bolivian Daphnia: the tropical Andes as a source for the North and South American D. pulicaria complex

We investigated genetic variation in asexual polyploid members of the water flea Daphnia pulex complex from a set of 12 Bolivian high-altitude lakes. We used nuclear microsatellite markers to study genetic relationships among all encountered multilocus genotypes, and combined this with a phylogenetic approach using DNA sequence data of three mitochondrial genes. Analyses of mitochondrial gene sequence divergence showed the presence of three very distinct clades that likely represent cryptic undescribed species. Our phylogenetic results suggest that the Daphnia pulicaria group, a complex of predominantly North American species that has diversified rapidly since the Pleistocene, has its origin in South America, as specific tests of topology indicated that all three South American lineages are ancestral to the North American members of this species group. A comparison between variation of nuclear and mitochondrial markers revealed that closely related polyploid nuclear genotypes sometimes belonged to very divergent mitochondrial lineages, while distantly related nuclear genotypes often belonged to the same mitochondrial lineage. This discrepancy suggests that these South American water fleas originated through reciprocal hybridization between different endemic, sexually reproducing parental lineages. It is also likely that polyploidy of the investigated lineages resulted from this hybridization. Nevertheless, no putative diploid parental lineages were found in the studied region.     

Association of ploidy and sexual system in Lycium californicum (Solanaceae)

In North American Lycium (Solanaceae), the evolution of gender dimorphism has been proposed as a means of restoring outcrossing after polyploidization causes the loss of self-incompatibility. Previous studies of this process in Lycium focused on comparisons between species that differ in ploidy. We examined intraspecific variation in floral morphology and DNA content in populations of L. californicum to determine correlations between sexual system and cytotype. We also used nuclear ITS and GBSSI sequence data to determine whether diploid and polyploid forms represent the same phylogenetic species, and the phylogeographic relationships among populations and ploidy levels. Within populations, no variation in ploidy was found, although among populations there was a perfect correspondence between sexual system and cytotype. Diploid populations were all hermaphroditic, whereas tetraploid populations were all gender dimorphic. There was no clear geographic pattern to the occurrence of diploid and tetraploid forms. Phylogenetic analysis confirms that L. californicum, regardless of ploidy, forms a monophyletic group within the genus Lycium. Sequences from diploid and polyploid individuals did not form reciprocally monophyletic clades, indicating either multiple gains of polyploidy, ongoing gene flow between cytotypes, or lack of lineage sorting since the evolution of polyploidy. The correspondence between ploidy and sex expression is consistent with the hypothesis that polyploidization triggers the evolution of gender dimorphism in this and other Lycium species.     

中华猕猴桃和美味猕猴桃自然居群遗传结构及其种间杂交渐渗

 利用9对SSR引物对中华猕猴桃（Actinidia chinensis）和美味猕猴桃（A. deliciosa）两近缘种的5个同域分布复合体和各自1个非同域分布居群进行了遗传多样性、居群遗传结构的分析以及种间杂交渐渗的探讨。结果表明：1）两物种共有等位基因比例高达81.13%,物种特有等位基因较少（中华猕猴桃：13.27%,美味猕猴桃：5.61%）,但共享等位基因表型频率在两近缘种间存在差异,而且与各同域复合体中两物种样本的交错程度或间距存在关联;2）两种猕猴桃均具有极高遗传多样性,美味猕猴桃的遗传多样性（H_o＝0 .749, PIC＝0.818）都略高于中华猕猴桃（H_o＝0.686,PIC＝0.799）;3）两猕 猴桃物种均具有较低的Nei’s居群遗传分化度,但AMOVA分析结果揭示种内异域居群间（F_ST=0.091 5）和同域复合体种间（F_ST=0.111 5）均存在一定程度的遗传分化;中华猕猴桃居群遗传分化（G_ST=0.086; F_ST=0.212 1）高于美味猕猴桃（G_ST= 0.080;F_ST=0.142 0）;4）同域分布复合体两物种间的遗传分化（G_ST=0.020）低于物种内异域居群间的遗传分化（中华猕猴桃：G_ST=0.086; 美味猕猴桃：G_ST=0.080）,同域复合体物种间的基因流（N_m＝7.89 -29.75）远远高于 同种异域居群间（中华猕猴桃：N_m =2.663; 美味猕猴桃：N_m=2.880）; 5）居群UPGMA聚类揭示在同一地域的居群优先聚类,个体聚类结果显示多数个体聚在各自居群组内,但各地理居群并不按地理距离的远近聚类,这与Mantel相关性检测所揭示的居群间遗传距离与地理距离没有显著性相关的结果一致。进一步分析表明两种猕猴桃的遗传多样性和居群遗传结构不仅受其广域分布、远交、晚期分化等生活史特性的影响,同时还与猕猴桃的染色体基数高 (x=29)、倍性复杂和种间杂交等因素密切相关,其中两种猕猴桃的共享祖先多态性和同域分布种间杂交基因渗透对两猕猴桃的居群遗传结构产生了重要影响。     

Nuclear-mitochondrial discordance and gene flow in a recent radiation of toads

Natural hybridization among recently diverged species has traditionally been viewed as a homogenizing force, but recent research has revealed a possible role for interspecific gene flow in facilitating species radiations. Natural hybridization can actually contribute to radiations by introducing novel genes or reshuffling existing genetic variation among diverging species. Species that have been affected by natural hybridization often demonstrate patterns of discordance between phylogenies generated using nuclear and mitochondrial markers. We used Amplified Fragment Length Polymorphism (AFLP) data in conjunction with mitochondrial DNA in order to examine patterns of gene flow and nuclear-mitochondrial discordance in the Anaxyrus americanus group, a recent radiation of North American toads. We found high levels of gene flow between putative species, particularly in species pairs sharing similar male advertisement calls that occur in close geographic proximity, suggesting that prezygotic reproductive isolating mechanisms and isolation by distance are the primary determinants of gene flow and genetic differentiation among these species. Additionally, phylogenies generated using AFLP and mitochondrial data were markedly discordant, likely due to recent and/or ongoing natural hybridization events between sympatric populations. Our results indicate that the putative species in the A. americanus group have experienced high levels of gene flow, and suggest that their North American radiation could have been facilitated by the introduction of beneficial genetic variation from admixture between divergent populations coming into secondary contact after glacial retreats.     

Linking genetic and ecological differentiation in an ungulate with a circumpolar distribution

Genetic differentiation among populations may arise from the disruption of gene flow due to local adaptation to distinct environments and/or neutral accumulation of mutations and genetic drift resulted from geographical isolation. Quantifying the role of these processes in determining the genetic structure of natural populations remains challenging. Here, we analyze the relative contribution of isolation‐by‐resistance (IBR), isolation‐by‐environment (IBE), genetic drift and historical isolation in allopatry during Pleistocene glacial cycles on shaping patterns of genetic differentiation in caribou/reindeer populations Rangifer tarandus across the entire distribution range of the species. Our study integrates analyses at range‐wide and regional scales to partial out the effects of historical and contemporary isolation mechanisms. At the circumpolar scale, our results indicate that genetic differentiation is predominantly explained by IBR and historical isolation. At a regional scale, we found that IBR, IBE and population size significantly explained the spatial distribution of genetic variation among populations belonging to the Euro‐Beringian lineage within North America. In contrast, genetic differentiation among populations within the North American lineage was predominantly explained by IBR and population size, but not IBE. We also found discrepancies between genetic and ecotype designation across the Holarctic species distribution range. Overall, these results indicate that multiple isolating mechanisms have played roles in shaping the spatial distribution of genetic variation across the distribution range of a large mammal with high potential for gene flow. Considering multiple spatial scales and simultaneously testing a comprehensive suite of potential isolating mechanisms, our study contributes to understand the ecological and evolutionary processes underlying organism–landscape interactions.     

Geographic independence and phylogenetic diversity of red shiner introductions

Identifying areas at risk of invasion can be difficult when the distribution of a non-native species encompasses geographically disjunct regions. Understanding genealogical relationships among native and non-native populations can clarify the origins of fragmented distributions, which in turn can clarify how fast and far a non-native species may spread. We evaluated genetic variation across the native and invasive ranges of red shiner (Cyprinella lutrensis), a minnow known to displace and hybridize with native species, to reconstruct invasion pathways across the United States (USA). Examination of mitochondrial cytochrome-b variation found that native range populations of red shiner fall into four highly divergent lineages that likely warrant species recognition. Introduced red shiner populations in the eastern and western USA are derived from only two of these lineages. Western USA populations originate from the mid-western and western genetic lineages, whereas eastern introductions derive only from the mid-western lineage. Western USA invasive populations exhibit fewer, but more diverse haplotypes compared to eastern USA invasive populations. We also recovered an undescribed, divergent lineage of Cyprinella that has been cryptically introduced into the western USA, which raises the possibility that hybridization has proceeded following secondary contact between previously allopatric lineages. Approximate Bayesian Computation modeling suggests that the disjunct distribution of red shiner across North America is an agglomeration of independent regional invasions with distinct origins, rather than stepwise advance of an invasion front or secondary introductions across regions. Thus localized control may be effective in managing non-native red shiner, including further spread to areas of conservation concern.     

Evidence for multiple sources of invasion and intraspecific hybridization in Brachypodium sylvaticum (Hudson) Beauv. in North America

We compared the levels and distribution of genetic diversity in Eurasian and North American populations of Brachypodium sylvaticum (Huds.) Beauv. (false brome), a newly invasive perennial bunchgrass in western North America. Our goals were to identify source regions for invasive populations, determine the number of independent invasion events, and assess the possibility that postinvasion bottlenecks and hybridization have affected patterns of genetic diversity in the invaded range. We tested the hypothesis that this Eurasian grass was accidentally introduced into two areas in Oregon and one site in California by examining nuclear microsatellites and chloroplast haplotype variation in 23 introduced and 25 native populations. In the invaded range, there was significantly lower allelic richness (R(S)), observed heterozygosity (H(O)) and within-population gene diversity (H(S)), although a formal test failed to detect a significant genetic bottleneck. Most of the genetic variation existed among populations in the native range but within populations in the invaded range. All of the allelic variation in the invaded range could be explained based on alleles found in western European populations. The distribution of identified genetic clusters in the North American populations and the unique alleles associated with them is consistent with two historical introductions in Oregon and a separate introduction to California. Further analyses of population structure indicate that intraspecific hybridization among genotypes from geographically distinct regions of western Europe occurred following colonization in Oregon. The California populations, however, are more likely to be derived from one or perhaps several genetically similar regions in the native range. The emergence and spread of novel recombinant genotypes may be facilitating the rapid spread of this invasive species in Oregon.     

DYNAMICS OF POLYPLOID FORMATION IN TRAGOPOGON (ASTERACEAE): RECURRENT FORMATION,GENE FLOW,AND POPULATION STRUCTURE

Polyploidy is a major feature of angiosperm evolution and diversification. Most polyploid species have formed multiple times, yet we know little about the genetic consequences of recurrent formations. Among the clearest examples of recurrent polyploidy are Tragopogon mirus and T. miscellus (Asteraceae), each of which has formed repeatedly in the last ～80 years from known diploid progenitors in western North America. Here, we apply progenitor‐specific microsatellite markers to examine the genetic contributions to each tetraploid species and to assess gene flow among populations of independent formation. These data provide fine‐scale resolution of independent origins for both polyploid species. Importantly, multiple origins have resulted in considerable genetic variation within both polyploid species; however, the patterns of variation detected in the polyploids contrast with those observed in extant populations of the diploid progenitors. The genotypes detected in the two polyploid species appear to represent a snapshot of historical population structure in the diploid progenitors, rather than modern diploid genotypes. Our data also indicate a lack of gene flow among polyploid plants of independent origin, even when they co‐occur, suggesting potential reproductive barriers among separate lineages in both polyploid species.     

Genetic Variations of 13 Indigenous Chinese Goat Breeds Based on Cytochrome b Gene Sequences

Phylogenetic relationships among and genetic variability within 13 Chinese indigenous goat breeds and Boer goat were analyzed using cytochrome b gene sequences. There were 44 variable sites found in a 642 bp sequence, and 46 Cyt b haplotypes were subsequently defined. The phylogeny analysis of haplotypes in combination with goat Cyt b sequences from GenBank shows that Chinese goats are obviously separated from wild goats and might come from Capra aegagrus. Further analysis indicated that indigenous Chinese goats might descend from at least two lineages; most of the individuals analyzed could be classified into lineage A as defined by Luikart, but five other goats were of uncertain lineage. The Tibet plateau is a possible place of origin for Chinese goats. The neighbor-joining tree based on pairwise differences among populations shows that most Tibetan goats, except the Middle Tibet type, cluster closely with North China goats, and then with South China goats. This result confirms that differences in genetic structure exist among goats in different geographic locations. Nucleotide diversity varied among populations. Tibet and North China goats had higher genetic diversity than South China goats. The fixation index (F _st=87.72%) suggested that most of the total genetic variation was due to variation within populations. In addition, the results indicate that Cyt b gene sequence information alone might not be enough for phylogeny analysis among breeds within species, as shown by fewer polymorphic sites and lower bootstrap values on the neighbor-joining tree.     

Genetic variation and evidence of hybridization in the genus Rhus (Anacardiaceae)

Rhus michauxii, a rare plant species endemic to the southeastern United States, was previously known only from central North Carolina and one site in Georgia. An additional site, which is now believed to represent the largest known concentration of R. michauxii, was recently discovered at Ft. Pickett near Blackstone, Virginia. Morphological characteristics in several of the Ft. Pickett Rhus populations appear to be intermediate between R. michauxii and the widespread R. glabra, a closely related congener that co-occurs at Ft. Pickett. Although morphological evidence of hybridization between R. michauxii and R. glabra in North Carolina has been provided previously, genetic marker data are lacking. In the present study we examined levels of allozyme variation at 11 polymorphic loci within and among seven populations of R. michauxii, one population of R. glabra, and four putative hybrid populations at Ft. Pickett. Overall, R. michauxii had typical levels of within-population genetic variation when compared to other species with similar life-history characteristics. In contrast, the proportion of genetic variation among populations (G(ST)) was considerably lower than expected. Finally, R. michauxii and R. glabra appear to have a fixed allelic difference at the Idh2 locus. This enabled us to confirm hybridization in all four of the putative hybrid populations and one of the R. michauxii populations.     

Mitochondrial DNA diversity in an apomictic Daphnia complex from the Canadian High Arctic

Cyclic parthenogenesis is the ancestral mode of reproduction in the cladoceran crustacean, Daphnia pulex, but some populations have made the transition to obligate parthenogenesis and this is the only mode of reproduction known to occur in arctic populations. Melanism and polyploidy are also common in arctic populations of this species. Prior allozyme studies of arctic D. pulex revealed substantial levels of clonal diversity on a regional scale. Clonal groupings based on cluster analysis of allozyme genotypes do not conform to groupings based on the presence/absence of melanin or on ploidy level. In order to further elucidate genetic relationships among arctic D. pulex clones, mitochondrial DNA (mtDNA) variation was examined in 31 populations from two Canadian high-arctic sites. The data were also compared to a previous study of mtDNA variation in populations from a Canadian low-arctic site. Cladistic analysis of restriction site variation of the entire mitochondrial genome and nucleotide sequence variation of the mitochondrial control region was used to construct genetic relationships among mitochondrial genotypes. Three distinct mitochondrial lineages were detected. One lineage was associated with diploid, nonmelanic clones and is the same as the lineage that is found in temperate populations of D. pulex. The other two lineages (A & B) were associated with polyploid, melanic clones. Sequence divergence between the A and B lineages was 2.4%. Sequence divergence between D. pulex and either of these two lineages exceeded 3%. It is suggested that the melanic, polyploid clones are hybrids between males of D. pulex (and/or a closely related congener, D. pulicaria) and females of either of two ancestral melanic species that have mitochondrial lineages A and B. Geographic patterns of mitochondrial diversity in ‘melanic’ lineage B support the hypothesis of an high-arctic refuge for the ancestral species during the last glacial period.     

表观遗传变异及其在作物改良中的应用

天然植物群体中存在着大量的遗传变异, 包括遗传物质改变和表观遗传变异, 它们是物种赖以生存和进化的源泉。表观遗传变异不涉及DNA序列的改变或者蛋白表达的变化, 但可以通过有丝分裂和(或)减数分裂实现世代间的稳定遗传。文章主要从表观遗传变异的重要来源--植物远缘杂交及多倍体化、环境中各种生物和非生物胁迫两方面, 总结了表观遗传在作物改良中的应用, 分析了它的局限性和存在的问题, 并且提出了相应的解决方法。     

Population genetic structure of two rare tree species (Colubrina oppositifolia and Alphitonia ponderosa,Rhamnaceae) from Hawaiian dry and mesic forests using random amplified polymorphic DNA markers

Hawaiian dry and mesic forests contain an increasingly rare assemblage of species due to habitat destruction, invasive alien weeds and exotic pests. Two rare Rhamnaceae species in these ecosystems, Colubrina oppositifolia and Alphitonia ponderosa, were examined using random amplified polymorphic DNA (RAPD) markers to determine the genetic structure of the populations and the amount of variation relative to other native Hawaiian species. Relative variation is lower than with other Hawaiian species, although this is probably not a consequence of genetic bottleneck. Larger populations of both species contain the highest levels of genetic diversity and smaller populations generally the least as determined by number of polymorphic loci, estimated heterozygosity, and Shannon's index of genetic diversity. Populations on separate islands were readily discernible for both species as were two populations of C. oppositifolia on Hawai'i island (North and South Kona populations). Substructure among Kaua'i subpopulations of A. ponderosa that were ecologically separated was also evident. Although population diversity is thought to have remained at predisturbance levels, population size continues to decline as recruitment is either absent or does not keep pace with senescence of mature plants. Recovery efforts must focus on control of alien species if these and other endemic dry and mesic forest species are to persist.     

Population genetic structure and hybridization patterns in the Mediterranean endemics Phlomis lychnitis and P. crinita (Lamiaceae)

BACKGROUND AND AIMS: The historical influence of gene flow and genetic drift after the last glacial phase of the Quaternary Period is reflected in current levels of genetic diversity and population structure of plant species. Moreover, hybridization after secondary contact might also affect population genetic diversity and structure. An assessment was made of the genetic variation and hybrid zone structure in Iberian populations of the Mediterranean Phlomis lychnitis and P. crinita, for which phylogenetic relationships are controversial, and hybridization and introgression are common. METHODS: Allozyme variation at 13 loci was analysed in 1723 individual plants sampled from 35 natural locations of P. lychnitis, P. crinita subsp. malacitana and P. crinita subsp. crinita in southern and eastern Spain. Standard genetic diversity parameters were calculated and patterns of genetic structure in each taxon were tested to fit the equilibrium between gene flow and genetic drift. Individual multilocus genotypes were subjected to Bayesian clustering analysis to estimate hybridization and introgression rates for both geographic regions. KEY RESULTS: Contrasting patterns in the distribution of genetic variation among the three taxa were found. Phlomis lychnitis showed no significant inbreeding, low genetic differentiation among populations and no evidence of isolation by distance. Phlomis crinita subsp. malacitana and P. crinita subsp. crinita showed high levels of genetic structure consistent with a pattern of gene flow-drift equilibrium. Higher instances of hybridization and introgression were detected in locations from southern Spain compared with locations from eastern Spain, matching unimodal and bimodal hybrid zones, respectively. CONCLUSIONS: High instances of historical gene flow, range expansion and altitudinal movement during the Quaternary Period, and lineage sorting can explain the diversity of patterns observed. The results suggest that P. lychnitis is the most differentiated lineage in the group; however, the relationship between the three taxa remains unclear.     

Cryptic intercontinental colonization in water fleas Daphnia pulicaria inferred from phylogenetic analysis of mitochondrial DNA variation

The water fleas of the Daphnia pulex complex play a key role in freshwater ecosystems throughout the northern hemisphere. Despite the fact that they have been the subject of study for numerous biological disciplines, their phylogeny and species delimitation remain controversial. We used DNA sequence variation of the mitochondrial ND5 gene to reconstruct the phylogenetic relationships of D. pulicaria Forbes, a widespread member of this complex from North America and Europe. Populations from the two continents respectively split into two evolutionary lineages, Eastern Nearctic and European, which each belong to another main clade within the D. pulex complex (the pulicaria and tenebrosa groups, respectively). Unexpectedly, melanin and carotenoid pigmented D. pulicaria populations from European high-mountain lakes were not allied with the transparent populations inhabiting the same lakes and the lowland ponds and reservoirs throughout Europe, but were included with the samples from Canada and Greenland in the Eastern Nearctic lineage. Until now populations belonging to this lineage were known only from Canada and North Atlantic islands, but not from mainland Europe. Independent data from microsatellite markers supported the genetic distinctiveness of the sympatric carotenoid pigmented and transparent populations and suggested that they may have undergone transition to obligate parthenogenesis, possibly as a consequence of past introgressive hybridization. Two different taxa are therefore confused under the name D. pulicaria in Europe. The close phylogenetic relationships of European populations with those from Canada and Greenland suggest that the Nearctic lineage is of recent origin in Europe via intercontinental dispersal from the North America. It has evolved melanin and carotenoid pigmentation as adaptations against the UV light stress, which enable it to share habitat occupied by the transparent European species. The Nearctic D. pulicaria thus provides a new model for studying successful intercontinental invasion. In general, our study demonstrates that a considerable part of the diversity among widespread taxa of cladoceran crustaceans has been overlooked in morphological taxonomies.     

Genetic variation and reproductive system among North American species of Nuttallanthus (Plantaginaceae)

We examined the effect of reproductive and life history strategies on the amount and partitioning of genetic variation in three annual species of Nuttallanthus. The North American species N. canadensis, N. floridanus, and N. texanus have regional to widespread ranges that overlap in the southeastern USA, are characterized by homogeneous populations and high fecundity, and possess showy, fragrant flowers seemingly adapted for insect pollination and outbreeding. Field and greenhouse studies on plants from 25 populations indicated that reproductive strategies were similar among species and showed predominant self-fertilization via cleistogamy and self-pollination prior to anthesis in chasmogamous flowers. Species were reproductively isolated and demonstrated complete cross-incompatibility after experimental crosses and no evidence for hybridization in mixed populations. Genetic variation was assessed using starch gel electrophoresis to resolve 15 isozyme loci in 50 populations. Conspecific genetic identity (I) values were high (0.819-0.936), but interspecific comparisons indicated many qualitative allelic differences and correspondingly low I values (0.516-0.623). Low levels of polymorphism and observed heterozygosity within populations and the disproportionate amount of gene diversity distributed among populations were concordant with reproductive data. The pattern of genetic differentiation was most similar to that observed in species with a predominantly inbreeding mating system.     

Genetic biogeography of the rare “copper moss,” Mielichhoferia elongata (Bryaceae)

Mielichhoferia elongata, one of the so-called “copper mosses,” has a broad but highly disjunctive geographic distribution and is rare throughout its range. A genetic analysis of 30 populations based on a survey of 21 allozyme loci reveals the following. 1) Total gene diversity at the specific level is high (0.41). 2) Within-population diversity is low, and over 90% of all genetic variation is among rather than within populations (mean G_ST = 0.93). 3) There is little differentiation in allele frequencies between North American and European populations. 4) Populations consist of one to six multilocus genotypes; 13 of the populations appear to consist of a single clone. 5) Colorado populations contain a tremendous reservoir of genetic variation (88% of all alleles found in the species in North America and Europe occur in one or more Colorado populations). 6) Populations in the eastern and western United States, and in Europe, contain subsets of the allelic diversity found in Colorado. The genetic structure of M. elongata suggests repeated dispersal and founding of populations.     

Dispersal Pathways and Genetic Differentiation among Worldwide Populations of the Invasive Weed Centaurea solstitialis L. (Asteraceae)

The natural history of introduced species is often unclear due to a lack of historical records. Even when historical information is readily available, important factors of the invasions such as genetic bottlenecks, hybridization, historical relationships among populations and adaptive changes are left unknown. In this study, we developed a set of nuclear, simple sequence repeat markers and used these to characterize the genetic diversity and population structure among native (Eurasian) and non-native (North and South American) populations of Centaurea solstitialis L., (yellow starthistle). We used these data to test hypotheses about the invasion pathways of the species that were based on historical and geographical records, and we make inferences about historical relationships among populations and demographic processes following invasion. We confirm that the center of diversity and the native range of the species is likely the eastern Mediterranean region in the vicinity of Turkey. From this region, the species likely proceeded to colonize other parts of Europe and Asia via a slow, stepwise range expansion. Spanish populations were the primary source of seed to invade South America via human-mediated events, as was evident from historical records, but populations from the eastern Mediterranean region were also important. North American populations were largely derived from South America, but had secondary contributors. We suggest that the introduction history of non-native populations from disparate parts of the native range have allowed not just one, but multiple opportunities first in South America then again in North America for the creation of novel genotypes via intraspecific hybridization. We propose that multiple intraspecific hybridization events may have created especially potent conditions for the selection of a noxious invader, and may explain differences in genetic patterns among North and South America populations, inferred differences in demographic processes, as well as morphological differences previously reported from common garden experiments.     

MORPHOLOGICAL AND GENETIC VARIABILITY IN PLANTAGO CORDATA (PLANTAGINACEAE), A THREATENED AQUATIC PLANT

Morphological and genetic variation were studied in Plantago cordata Lam., an imperiled aquatic plant. The existence of distinct seasonal morphs was confirmed by numerical morphological analyses of specimens from 53 populations. Plants from two North Carolina populations possessed leaves that were significantly shorter, narrower, and fewer-veined than populations in other portions of the species range. Genetic variation within and among ten populations in seven states/provinces (Illinois, Missouri, New York, North Carolina, Ohio, Ontario, and Wisconsin) was examined electrophoretically. The species is an apparent allopolyploid with fixed heterozygosity observed at 67% of the loci surveyed. Electrophoretic variation was mostly partitioned among populations. The genetic identity among populations was high except for North Carolina populations which differed at 27% of the loci surveyed.