Multiple introductions boosted genetic diversity in the invasive range of black cherry (Prunus serotina; Rosaceae)

Background and Aims

Black cherry (Prunus serotina) is a North American tree that is rapidly invading European forests. This species was introduced first as an ornamental plant then it was massively planted by foresters in many countries but its origins and the process of invasion remain poorly documented. Based on a genetic survey of both native and invasive ranges, the invasion history of black cherry was investigated by identifying putative source populations and then assessing the importance of multiple introductions on the maintenance of gene diversity.

Methods

Genetic variability and structure of 23 populations from the invasive range and 22 populations from the native range were analysed using eight nuclear microsatellite loci and five chloroplast DNA regions.

Key Results

Chloroplast DNA diversity suggests there were multiple introductions from a single geographic region (the north-eastern United States). A low reduction of genetic diversity was observed in the invasive range for both nuclear and plastid genomes. High propagule pressure including both the size and number of introductions shaped the genetic structure in Europe and boosted genetic diversity. Populations from Denmark, The Netherlands, Belgium and Germany showed high genetic diversity and low differentiation among populations, supporting the hypothesis that numerous introduction events, including multiple individuals and exchanges between sites, have taken place during two centuries of plantation.

Conclusions

This study postulates that the invasive black cherry has originated from east of the Appalachian Mountains (mainly the Allegheny plateau) and its invasiveness in north-western Europe is mainly due to multiple introductions containing high numbers of individuals.     

Spatial Genetic Heterogeneity in Populations of a Newly Invasive Whitefly in China Revealed by a Nation-Wide Field Survey

Background

Even though introductions of exotic species provide ready-made experiments of rapid evolution, few studies have examined the genetic structure of an exotic species shortly after its initial introduction and subsequent spread. To determine the genetic structure of its populations during the initial introduction, we investigated the invasive sweet potato whitefly (Bemisia tabaci Q, commonly known as B. tabaci biotype Q) in China, which was introduced in approximately 2003. A total of 619 B. tabaci Q individuals in 20 provinces throughout China were collected and analyzed using five microsatellite loci.

Results

The introduced populations of B. tabaci Q in China represent eight genetic clusters with different geographic distributions. The populations in Yunnan Province, where B. tabaci Q was first detected, are genetically different from the other populations in China.

Conclusion

The introduced populations of B. tabaci Q in China have high spatial genetic heterogeneity. Additional research is required to determine whether the heterogeneity results from multiple introductions, rapid evolution following one or few introductions, or some combination of multiple introductions and rapid evolution. The heterogeneity, however, is inconsistent with a single introduction at Yunnan Province, where B. tabaci Q was first detected, followed by spread.     

Reconstructing the introduction history of an invasive fish predator in South Africa

The Largemouth Bass (Micropterus salmoides) is a global invader with demonstrated ecological impacts on native fish communities. Introductions of fishes in freshwater ecosystems are often characterized as complex processes, yet an understanding of the nature of the introduction can inform management and conservation actions. Early in the twentieth century, two introductions of Largemouth Bass were made into South Africa for the establishment of a recreational fishery, and subsequent translocations have expanded their distribution to include much of southern Africa. In this study we quantified neutral genetic variation, modeled potential introduction scenarios, and identified potential source regions from within the native range. We documented limited levels of genetic diversity in nuclear microsatellite genotypes across populations (mean allelic richness = 1.80 and mean observed heterozygosity = 0.16) and observed low levels of genetic differentiation among four of the five focal populations (mean pairwise fixation index = 0.09), with a fifth population displaying greater levels of genetic divergence (mean pairwise fixation index = 0.27). A total of three cytochrome b haplotypes were recovered from South Africa samples and the single most common haplotype (93% of individuals) was identical to a haplotype from a population of Largemouth Bass in Maryland, USA. Using limited available stocking data along with outputs from Principal Component Analysis and approximate Bayesian evaluation of competing introduction scenarios we confirm the presence of multiple introductions. Despite evidence for multiple introductions, Largemouth Bass in South African water bodies harbor extremely low neutral genetic diversity, suggesting that even a very limited number of propagules can experience a high likelihood of success in invading nonnative waters.     

Reduced variation among northern deer tick populations at an autosomal microsatellite locus.

To determine whether genes flow freely between populations of the Ixodes ricinus-like ticks of eastern North America, and to determine whether the abundant northerly populations of these vectors of Lyme disease and other zoonotic infections may have arisen recently from a small cohort of ancestral founders, we characterized the nuclear IR27 microsatellite alleles in ticks sampled from the geographic extremes of their ranges. These microsatellite alleles differentiated populations located in the southeastern, northcentral and northeastern United States, respectively. Although evident heterozygous genotypes are about as frequent as would be expected in randomly mating populations, particular microsatellite alleles and diploid genotypes occur more frequently in certain populations than in others. Ticks from the Northeast and upper Midwest are markedly more related to each other than to ticks from the Southeast. Patterns of diversity present in this nuclear microsatellite marker correspond to those evident at a mitochondrial locus and indicate that the deer ticks of the Northeast and upper Midwest are genetically isolated from those in the Southeast. The Ixodes ricinus-like ticks that impose a public health burden in the northeastern and northcentral United States originated recently in a common founder population.     

The diminutive supercolony: the Argentine ants of the southeastern United States

Native to Argentina and Brazil, the Argentine ant (Linepithema humile) is an invasive species that has become established on six continents and many oceanic islands. In several parts of its introduced range, including the western United States, southern Europe and Chile, the Argentine ant is unicolonial, forming extensive supercolonies. We examined population genetic structure and intercolony aggression in two regions of the introduced range of this species in the United States: California and the southeastern United States. Our results show that the southeastern L. humile population has high genotypic variability and strong intercolony aggression relative to the California population. In the California population, intercolony aggression was absent and 23 alleles were found across seven polymorphic microsatellite loci. However, in the Southeast, aggression between colonies was high and 47 alleles were present across the same seven loci in an equal number of colonies. We suggest that distinctly different colonization patterns for California and the Southeast may be responsible for the striking disparity in the genetic diversity of introduced populations. Southeastern colonies may have descended from multiple, independent introductions from the native range, undergoing a bottleneck at each introduction. In contrast, the California supercolony may have originated from one or more colonies inhabiting the southeastern United States, thus experiencing a double bottleneck. The differences in present-day distribution patterns between California and the Southeast may be due to the combined effect of two factors: lower winter temperatures in the Southeast and/or competition with another successful and widely distributed ant invader, the fire ant Solenopsis invicta.     

Coumarin in vanilla extracts: Its detection and significance

Coumarin was a component of some commercial vanilla preparations until banned in the United States by the Food and Drug Administration in 1954 because of evidence of hepatotoxicity in animals. However, in the present investigation coumarin was detected by capillary column gas chromatography/mass spectrometry in Mexican vanilla extracts commercially available or introduced in the United States by tourists or travelers. The significance of the presence of coumarin in Mexican vanilla extracts is discussed in light of recent evidence suggesting a lack of significant coumarin toxicity in humans.     

Haplotype diversity of mt<Emphasis Type="Italic">COI</Emphasis> in the fall webworm <Emphasis Type="Italic">Hyphantria cunea</Emphasis> (Lepidoptera: Arctiidae) in introduced regions in China,Iran, Japan,Korea, and its homeland,the United States

Hyphantria cunea (Drury) has colonized many countries outside its native range of North America and has become a model species for studies of the colonization and subsequent adaptation of agricultural pests. Molecular genetic analyses can clarify the origin and subsequent adaptations to non-native habitats. Using the mitochondrial COI gene, we examined the genetic relationships between invasive populations (China, Iran, Japan, and Korea) and native populations (i.e., the United States). The Jilin (China) and Guilan (Iran) populations showed nine previously unknown haplotypes that differed from those found in the south–central United States, suggesting multiple colonization events and different regions of invasion. A dominant mtDNA haplotype in populations in the United States was shared by all of the populations investigated, suggesting that H. cunea with that haplotype have successfully colonized China, Iran, Japan, and Korea.     

Conservation genetics of redside dace (<Emphasis Type="Italic">Clinostomus elongatus</Emphasis>): phylogeography and contemporary spatial structure

Redside dace Clinostomus elongatus (Teleostei: Cyprinidae) is a species of conservation concern that is declining throughout its range as a result of habitat fragmentation, degradation and loss. We characterized the genetic structure and diversity of redside dace populations across the species range using mitochondrial and microsatellite data to inform conservation efforts and assess how historical and recent events have shaped genetic structure and diversity within and among populations. Phylogeographic structure among 28 redside dace populations throughout southern Ontario (Canada) and the United States was assessed by sequence analysis of the mitochondrial cytochrome b and ATPase 6 and 8 genes. Populations were also genotyped using ten microsatellite loci to examine genetic diversity within and among populations as well as contemporary spatial structuring. Mitochondrial DNA sequence data revealed three geographically distinct lineages, which were highly concordant with groupings identified by microsatellite analysis. The combined genetic data refute published glacial refugia hypotheses of a single Mississippian refugium or of two lineages associated with Mississippian and Atlantic refugia. Secondary contact between the two eastern groups was documented in the Allegheny River drainage and tributaries to Lake Ontario. With the exception of several allopatric populations within the Allegheny watershed, high genetic structuring among populations suggests their isolation, indicating that recovery efforts should be population-based.     

Displacement of native Patagonian freshwater silverside populations (<Emphasis Type="Italic">Odontesthes hatcheri</Emphasis>, Atherinopsidae) by introgressive hybridization with introduced <Emphasis Type="Italic">O. bonariensis</Emphasis>

The Patagonian silverside Odontesthes hatcheri is a native fish restricted to streams and lakes of Patagonia (Argentina and Chile). Stocking programs to enhance recreational fisheries in man-made reservoirs have introduced a nonnative, closely-related species (the pejerrey O. bonariensis) in Patagonia almost a century ago, and yet little is known about the invasiveness of this species. To evaluate the impact of these introductions we analyze genetic data (microsatellite markers and mitochondrial DNA) to quantify the incidence of hybridization between these two species and assess potential effects on native population structure. Phylogeographic analyses reveal weak geographic differentiation among populations of O. hatcheri, in agreement with previous studies for other freshwater fishes in Patagonia strongly influenced by Quaternary glaciations and hydrographic basin changes since the last glaciation. However, many populations have unique genetic pools. In several areas, introductions resulted in extensive hybridization, with high frequencies of F2 and backcrossed hybrids in natural populations, and in some cases O. bonariensis has completely displaced the native populations. The negative impact of these introductions on native populations is correlated to temperature, a critical parameter in the face of global warming, suggesting that invasiveness of O. bonariensis may increase in the future. Our results advise against continuing stocking programs to preserve the integrity of natural populations of the Patagonian silverside.     

Genetic diversity and invasion history of the European subterranean termite <Emphasis Type="Italic">Reticulitermes urbis</Emphasis> (Blattodea,Termitoidae, Rhinotermitidae)

Biological invasions are among key factors of ecological changes, and social insects appear as highly successful invasive animals. Subterranean termites of the holarctic genus Reticulitermes are present in Europe with six native and one invasive (the nearctic R. flavipes) species. The species R. urbis shows a disjunct distribution in the Western Balkans, Eastern Italy and Southern France. Previous molecular and population genetics data suggested that the taxon originated from the Balkans, and that Italian and French populations are invasive, but it is still unknown how many introduction events occurred and from which Balkan source populations. To address these questions, a population genetics analysis was performed on a larger sampling than previous studies, using mitochondrial cytochrome oxidase II and 6 microsatellite markers on 47 colonies collected across the whole distribution area. Mitochondrial analysis confirmed the presence of two major lineages where colonies from Balkans, Italy, and France intermingle. Similarly, microsatellite loci analysis indicated the presence of two genetic clusters, though not corresponding to the two mitochondrial clades, each including colonies from the three sampled areas and with individuals showing mixed cluster membership. Overall, French and Italian populations showed indications of bottleneck (reduced genetic diversity and change of allele frequencies) and do not appear genetically differentiated from the Balkan population. Results presented here support a history of multiple introductions in Italy and France, in a scenario consistent with continuous exchanges between native and invasive areas, as expected along human trades routes.     

Chromosome numbers,polyploidy, and hybridization in castilleja (scrophulariaceae ) of the Great Basin and Rocky Mountains

Chromosome numbers are reported for 190 collections ofCastilleja of the Great Basin-Rocky Mountains and adjacent regions in western North America. Polyploidy has played a major role in evolu tion of this genus. Of the 35 taxa represented, 14 are exclusively diploid (n =12). The remaining 21 taxa exhibit polyploidy: eight are tetraploid only, two are octopIoid only, while 11 of the more wide-ranging species have two or more levels of polyploidy. BothC. rhexifolia andC. sulphured have diploid, tetraploid, and octoploid races while the widespreadC. miniata has four (2x, 4x, 8x, and 10x). Where appropriate, the role of polyploidy and hybridization inCastilleja evolution and their relevance to taxonomy are discussed for individual species.     

Fine‐scale spatial and temporal population genetics of Aedes japonicus,a new US mosquito,reveal multiple introductions

The newly introduced mosquito Aedes japonicus has expanded from its original range in Northeastern Asia to 29 US states (including Hawaii) plus Canada and northern Europe. Our objectives were to test an earlier hypothesis of multiple introductions of this species to the Northeastern US and evaluate putative temporal changes in genetic makeup. Using a panel of seven microsatellite loci, we confirmed the existence of two abundant genetic forms in specimens originally collected in 1999–2000 (F_ST value based on microsatellite data = 0.26) that matches the disjunctive distribution of mitochondrial haplotypes. To examine the distribution of the two genetic ‘types’ across Pennsylvania we created a fine‐scale genetic map of Ae. japonicus using 439 specimens collected from 54 Pennsylvania counties in 2002–2003. We also made direct comparisons between collections in 1999–2000 and new collections made in 2004–2005 obtained from the same areas in the northeastern US. We observed that the strong association between mtDNA haplotype and microsatellite signature seen in 1999–2000 had weakened significantly by 2002 across Pennsylvania, a trend continued to some extent in 2004–2005 in PA, NJ, and NY, indicating that once easily distinguishable separate introductions are merging. The two expanding genetic forms create a complex correlation between spatial and genetic distances. The existence of multiple introductions would be obscured without sampling early and across time with highly polymorphic molecular markers. Our results provide a high‐resolution analysis of the spatial and temporal dynamics of a newly introduced disease vector and argue that successive introductions may be a common pattern for invasive mosquitoes.     

Invasion Genetics of New World Medflies: Testing Alternative Colonization Scenarios

The Mediterranean fruit fly (Ceratitis capitata) is an invasive agricultural pest with a wide host range and a nearly global distribution. Efforts to forgo the medfly's spread into the United States are dependent on an understanding of population dynamics in newly established populations elsewhere. To explore the potential influence of demographic and historical parameters in six medfly populations distributed from Mexico to Peru, we created population genetic null models using Monte Carlo simulations. Null expectations for genetic differentiation (F _ST) were compared with actual sequence variation from four highly polymorphic nuclear loci. Four colonization scenarios that were modeled led to unique genetic signatures that could be used to interpret empirical data. Unless current gene flow across Latin America was assumed to be very high, we could reject colonizations consisting of multiple introductions, each of low genetic diversity. Further, if simulated populations were small (N _e = 5 × 10² individuals per population), small invasions from a single source consistently produced F _ST values comparable to those currently observed in Latin America. In contrast, only large invasions from diverse sources were compatible with the observed data for large populations (N _e 5 × 10³). This study demonstrates that alternative population genetic hypotheses can be tested empirically even when departures from equilibrium are extreme, and that population genetic theory can be used to explore the processes that underlie biological invasions.     

Recipe for disruption: multiple recent arrivals of megachilid bees in Pacific archipelagos

When examining how the ecosystems of remote islands have developed, it is important to know the timing of when various elements arrived and whether they then diversified. Our understanding of the histories behind the biodiverse south west Pacific (SWP) archipelagos is limited, and further impeded by the complex geological histories of this region. Previous studies of the SWP short-tongued halictine bee fauna suggest their presence is much younger than the geological ages of these archipelagos, which is surprising given their critical role as pollinators in other terrestrial ecosystems. The long-tongued megachilid bees represent a considerable proportion of the known bee species for the region, yet little is known of their origin. Here we use genetic diversity within mitochondrial DNA to infer the likely ages and origins of megachilid species from Vanuatu, Fiji, and Samoa. Our results indicate a very recent origin for megachilids in the SWP, with many species exhibiting small intraspecific genetic distances. Three species share almost identical haplotypes with specimens from Southeast Asia, suggesting multiple human-aided introductions. Combined with data from recent studies on other bee groups present in the region, our results have broad implications for how the Pacific island biota developed and how we should approach its management.     

The evolution of Hessian fly from the Old World to the New World: Evidence from molecular markers

Eighteen polymorphic microsatellite loci and 11 single‐nucleotide polymorphisms were genotyped in 1 095 individual Hessian fly specimens representing 23 populations from North America, southern Europe, and southwest Asia. The genotypes were used to assess genetic diversity and interrelationship of Hessian fly populations. While phylogenetic analysis indicates that the American populations most similar to Eurasian populations come from the east coast of the United States, genetic distance is least between (Alabama and California) and (Kazakhstan and Spain). Allelic diversity and frequency vary across North America, but they are not correlated with distance from the historically documented point of introduction in New York City or with temperature or precipitation. Instead, the greatest allelic diversity mostly occurs in areas with Mediterranean climates. The microsatellite data indicate a general deficiency for heterozygotes in Hessian fly. The North American population structure is consistent with multiple introductions, isolation by distance, and human‐abetted dispersal by bulk transport of puparia in infested straw or on harvesting equipment.     

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.     

Invasion genetics of <Emphasis Type="Italic">Senecio vulgaris</Emphasis>: loss of genetic diversity characterizes the invasion of a selfing annual,despite multiple introductions

Genetic variation in invasive populations is affected by a variety of processes including stochastic forces, multiple introductions, population dynamics and mating system. Here, we compare genetic diversity between native and invasive populations of the selfing, annual plant Senecio vulgaris to infer the relative importance of genetic bottlenecks, multiple introductions, post-introduction genetic drift and gene flow to genetic diversity in invasive populations. We scored multilocus genotypes at eight microsatellite loci from nine native European and 19 Chinese introduced populations and compared heterozygosity and number of alleles between continents. We inferred possible source populations for introduced populations by performing assignment analyses and evaluated the relative contributions of gene flow and genetic drift to genetic diversity based on correlations of pairwise genetic and geographic distance. Genetic diversity within Chinese populations was significantly reduced compared to European populations indicating genetic bottlenecks accompanying invasion. Assignment tests provided support for multiple introductions with populations from Central China and southwestern China descended from genotypes matching those from Switzerland and the UK, respectively. Genetic differentiation among populations in China and Europe was not correlated with geographic distance. However, European populations exhibited less variation in the relation between G _ST and geographical distance than populations in China. These results suggest that gene flow probably plays a more significant role in structuring genetic diversity in native populations, whereas genetic drift appears to predominate in introduced populations. High rates of selfing in Chinese populations may restrict opportunities for pollen-mediated gene flow. Repeated colonization-extinction cycles associated with ongoing invasion is likely to maintain low genetic diversity in Chinese populations.     

Genetic structure of the tiger mosquito, Aedes albopictus, in Cameroon (Central Africa)

Background

Aedes albopictus (Skuse, 1884) (Diptera: Culicidae), a mosquito native to Asia, has recently invaded all five continents. In Central Africa it was first reported in the early 2000s, and has since been implicated in the emergence of arboviruses such as dengue and chikungunya in this region. Recent genetic studies of invasive species have shown that multiple introductions are a key factor for successful expansion in new areas. As a result, phenotypic characters such as vector competence and insecticide susceptibility may vary within invasive pest species, potentially affecting vector efficiency and pest management. Here we assessed the genetic variability and population genetics of Ae. albopictus isolates in Cameroon (Central Africa), thereby deducing their likely geographic origin.

Methods and Results

Mosquitoes were sampled in 2007 in 12 localities in southern Cameroon and analyzed for polymorphism at six microsatellite loci and in two mitochondrial DNA regions (ND5 and COI). All the microsatellite markers were successfully amplified and were polymorphic, showing moderate genetic structureamong geographic populations (F_ST = 0.068, P<0.0001). Analysis of mtDNA sequences revealed four haplotypes each for the COI and ND5 genes, with a dominant haplotype shared by all Cameroonian samples. The weak genetic variation estimated from the mtDNA genes is consistent with the recent arrival of Ae. albopictus in Cameroon. Phylogeographic analysis based on COI polymorphism indicated that Ae. albopictus populations from Cameroon are related to tropical rather than temperate or subtropical outgroups.

Conclusion

The moderate genetic diversity observed among Cameroonian Ae. albopictus isolates is in keeping with recent introduction and spread in this country. The genetic structure of natural populations points to multiple introductions from tropical regions.     

Highlighting the occurrence of tetraploidy in Acacia senegal (L.) Willd. and genetic variation patterns in its natural range revealed by DNA microsatellite markers

Acacia senegal (L.) Willd. is the main species producing the internationally traded gum arabic. Genetic studies of this species are rare and until now the chromosome number was thought to be diploid (2n?=?2x?=?26). Here, using chromosome number counting, we demonstrate for the first time that tetraploids (2n?=?4x?=?52) also occur in A. senegal. Nuclear and chloroplast microsatellite markers were used to estimate and compare genetic variation within this newly described polyploidy complex in the Sudano-Sahelian region in Africa. Genetic diversity was higher in diploids, suggesting that the formation of tetraploids is recent and that mutation–drift equilibrium has not yet been reached. The two cytotypes do not have the same genetic structure and are genetically differentiated. Among tetraploids, populations are greatly differentiated and do not share the same chlorotypes. Based on these results, we discuss recurrent formation of tetraploids from different diploid progenitors across the distribution range of A. senegal in the Sudano-Sahelian zone.     

Taxonomic and phylogenetic significance of seed coat microsculpturing in Mentzelia (Loasaceae) in Wyoming and adjacent western states

Six hundred seeds belonging to the genusMentzelia (Loasaceae) were examined using the Scanning Electron Microscope. The seeds represented all six sections of the genus and approximately 76% of the total number of species from northern Mexico and western United States. Emphasis was placed on seed material from Wyoming and adjoining states. Ovule serial sectioning was done to determine testa ontogeny. In all cases seed surface structures could be related to excrescences developing from the integument cells. Six basic seed coat relief features could be distinguished, corresponding to the six sections in the genus. Minor variations in the basic types are constant and characteristic for species or, in very few instances, for species groups. A dendrogram was constructed from the seed testa data which supports the phylogeny established from chromosomal and morphological data. Practical application of seed coat microcharacter specificity to identification ofMentzelia seeds from prehistoric sites is suggested.