Patterns of genetic differentiation in two annual bromegrasses,Bromus lanceolatus andB. hordeaceus (Poaceae)

Infraspecific genetic differentiation was analysed in two tetraploid annual bromegrasses,Bromus lanceolatus (from N Africa) andB. hordeaceus (from N Africa and France). Genetic analysis of populations was based on allozyme polymorphisms at 17 loci. Different fixed heterozygous phenotypes were scored in both species, according to their allopolyploid origin. In N Africa, more variation occurred among populations ofB. lanceolatus than ofB. hordeaceus. The variation was not randomly distributed among populations of both species. InB. lanceolatus, differentiation was correlated with climatic variables rather than with geographic distance between populations. Higher correlation of genetic differentiation with geographic distance occurred inB. hordeaceus, particularly at large geographic scale, between French and N African populations. Within each region, the populations appeared weakly genetically differentiated, even when belonging to different subspecies.     

Genetic Evaluation of the Efficacy of In Situ and Ex Situ Conservation of <Emphasis Type="Italic">Parashorea chinensis</Emphasis> (Dipterocarpaceae) in Southwestern China

The majority of research in genetic diversity yields recommendations rather than actual conservation achievements. We assessed the efficacy of actual in situ and ex situ efforts to conserve Parashorea chinensis (Dipterocarpaceae) against the background of the geographic pattern of genetic variation of this species. Samples from seven natural populations, including three in a nature reserve, and one ex situ conservation population were studied. Across the natural populations, 47.8% of RAPD loci were polymorphic; only 20.8% on average varied at the population level. Mean population genetic diversity was 0.787 within natural populations and 1.410 for the whole species. Significant genetic differentiation among regions and isolation by distance were present on larger scales (among regions). AMOVA revealed that the majority of the among-population variation occurred among regions rather than among populations within regions. Regression analysis, Mantel test, principal coordinates analysis, and cluster analysis consistently demonstrated increasing genetic isolation with increasing geographic distance. Genetic differentiation within the region was quite low compared to that among regions. Multilocus spatial autocorrelation analysis of these three populations revealed random distribution of genetic variation in two populations, but genetic clustering was detected in the third population. The ex situ conserved population contained a medium level of genetic variation compared with the seven natural populations; it contained 77.1% of the total genetic variation of this species and 91% of the moderate to high frequency RAPD fragments (f > 0.05). Exclusive bands were detected in natural populations, but none were found in the ex situ conserved population. The populations protected in the nature reserve contained most of the genetic variation of the whole species, with 81.4% of the total genetic variation and 95.7% of the fragments with moderate to high frequency (f > 0.05) of this species conserved. The results show that the ex situ conserved population does not contain enough genetic variation to meet the need of release in the future, and that more extensive ex situ sampling in natural populations TY, NP, HK, and MG is needed. The in situ conserved population contains representative genetic variation to maintain long-term survival and evolutionary processes of P. chinensis.     

A cline of allozyme variation inAbies mariesii

Genetic variation at 22 allozyme loci was examined for 1,003 trees from 11 isolated natural populations ofAbies mariesii covering all except the southernmost region of its geographic range. Genetic diversity within species (H _es=0.063) was low compared to many other long-lived woody species. Most of the genetic variation is found within populations (G _ST=0.144) despite their isolated distribution. Genetic distance between populations was positively correlated with geographic distance. Genetic diversity within populations was generally low (meanH _ep=0.054), but varied across populations in a clinal fashion such that genetic variation decreased with increasing latitude. These genetic characteristics may reflect the distribution history of this species.     

Phylogeographic studies on natural populations of the South American fruit fly, <Emphasis Type="Italic">Anastrepha fraterculus</Emphasis> (Diptera: Tephritidae)

Anastrepha fraterculus is an important pest of commercial fruits in South America. The variability observed for morphological and behavioural traits as well as genetic markers suggests that A. fraterculus represents a complex of synmorphic species rather than a single biological species. We studied the correlation between geographical distribution and genetic variation in natural populations from Argentina and south Brazil. Fragments of the mitochondrial gene COII were sequenced in 28 individuals. The matrix of aligned sequences was phylogenetically analysed by parsimony, yielding a cladogram of haplotypes. Based on Templeton’s nested method, no clade showed any geographic pattern for the gene COII, indicating lack of significant association between haplotypic variability and geographic distribution. The analysis of nucleotide substitution distances by Neighbour-Joining algorithm showed that geographically distant populations exhibit low genetic distances. The corresponding trees clustered the populations without showing any geographic pattern. This result suggests that the populations studied are not reproductively isolated.     

Phylogeographic relationships among worldwide populations of the cosmopolitan marine species, the striped gray mullet (Mugil cephalus), investigated by partial cytochrome b gene sequences

Several factors such as geographical barriers, demographic history, biological and ecological traits may contribute to delineate the evolutionary history of a species. The gray mullet, Mugil cephalus, represents an interesting case of a marine species with a coastal ecology and a cosmopolitan distribution. In this study, partial cytochrome b sequences were resolved for 177 M. cephalus specimens sampled from 14 different geographic sites, in order to investigate the genetic divergence and the phylogeographic relationship among populations at a global scale. Demographic parameters were also assessed. Analysis of partial cyt b sequences showed high levels of differentiation among populations from distant geographic areas as most populations harbored private alleles and showed reciprocal monophyly. Both phylogenetic trees and haplotype network indicate the East Australian haplotypes as the closest to the ancestor sequences, which leads to the hypothesis of an Indo-West Pacific center of origin for M. cephalus. The analyses of the molecular variance showed that the genetic variation in M. cephalus is mainly harbored among populations rather than within populations. The high variability indices (h, π) calculated on M. cephalus individuals pooled together and the very low variability indices detected at some geographic sampling sites suggest that gray mullet populations have undergone a long-term reproductive isolation characterized by events of population abundance reductions which may have favored genetic differentiation among populations.     

基于cpDNA trnL-F序列的胡黄连保护遗传学研究

胡黄连为特产中国-喜马拉雅特有高山植物,作为常用中、藏药材,受到灭绝性采挖,作为濒危和二级保护植物亟待科学的保护。该研究以云南和西藏7个野生居群91个个体为材料,基于 cpDNA trnL-F 非编码序列测序分析胡黄连的遗传多样性和遗传结构,分析显著进化单元,确立优先保护居群并提出科学的保护策略。结果表明：胡黄连 trnL-F 序列长度为871~876 bp,根据序列的核苷酸变异共鉴定出5个单倍型,西藏占有2个单倍型,云南占有3个单倍型,西藏和云南2个地区的所有单倍型均不共享。胡黄连具有较低的单倍型多样性(Hd ＝0.43419)和核苷酸多样性(Dij ＝0.00466)。种群间分化度(Fst ＝0.864520)和基因流(Nm ＝0.04)、居群间的遗传分化水平(GST ＝0.916)、AMOVA 分析(0.78％的遗传变异发生在居群内,60.97％的遗传变异发生在地区内居群间,38.25％的遗传变异发生在地区间)均表明,胡黄连居群间存在明显遗传分化。多数一致性树将胡黄连划分为3个进化分支(Ⅰ、Ⅱ、Ⅲ),这3个分支均与地理相关,分支Ⅰ分布于横断山区的4个居群,分支Ⅱ是分布于东喜马拉雅的一个居群,分支Ⅲ是分布于喜马拉雅中段的2个居群。3个分支分属于3个“进化显著单元(ESU)”。这3个 ESU 中白马雪山、茨中、定日、波密、聂拉木五个居群都需要保护,建议现阶段应优先保护的居群是云南白马雪山和西藏波密居群,以就地保护为主。     

Molecular and phenotypic diversity in Chionactis occipitalis (Western Shovel-nosed Snake), with emphasis on the status of C. o. klauberi (Tucson Shovel-nosed Snake).

Chionactis occipitalis (Western Shovel-nosed Snake) is a small colubrid snake inhabiting the arid regions of the Mojave, Sonoran, and Colorado deserts. Morphological assessments of taxonomy currently recognize four subspecies. However, these taxonomic proposals were largely based on weak morphological differentiation and inadequate geographic sampling. Our goal was to explore evolutionary relationships and boundaries among subspecies of C. occipitalis, with particular focus on individuals within the known range of C. o. klauberi (Tucson Shovel-nosed snake). Population sizes and range for C. o. klauberi have declined over the last 25 years due to habitat alteration and loss prompting a petition to list this subspecies as endangered. We examined the phylogeography, population structure, and subspecific taxonomy of C. occipitalis across its geographic range with genetic analysis of 1100 bases of mitochondrial DNA sequence and reanalysis of 14 morphological characters from 1543 museum specimens. We estimated the species gene phylogeny from 81 snakes using Bayesian inference and explored possible factors influencing genetic variation using landscape genetic analyses. Phylogenetic and population genetic analyses reveal genetic isolation and independent evolutionary trajectories for two primary clades. Our data indicate that diversification between these clades has developed as a result of both historical vicariance and environmental isolating mechanisms. Thus these two clades likely comprise ‘evolutionary significant units’ (ESUs). Neither molecular nor morphological data are concordant with the traditional C. occipitalis subspecies taxonomy. Mitochondrial sequences suggest specimens recognized as C. o. klauberi are embedded in a larger geographic clade whose range has expanded from western Arizona populations, and these data are concordant with clinal longitudinal variation in morphology.     

Contrasting patterns of geographic variation in the cosmopolitan sibling species Drosophila melanogaster and Drosophila simulans

An electrophoretic study was carried out to compare the geographic pattern of genetic variation in Drosophila simulans with that of its sibling species, Drosophila melanogaster. An identical set of 32 gene-protein loci was studied in four geographically distant populations of D. simulans and two populations of D. melanogaster, all originating from Europe and Africa. The comparison yielded the following results: (1) tropical populations of D. simulans were, in terms of the number of unique alleles, average heterozygosity per locus, and percentage of loci polymorphic, more variable than conspecific-temperate populations; (2) some loci in both species showed interpopulation differences in allele frequencies that suggest latitudinal clines; and (3) temperate-tropical genetic differentiation between populations was much less in D. simulans than in D. melanogaster. Similar differences between these two species have previously been shown for chromosomal, quantitative, physiological, and middle-repetitive DNA variation. Estimates of N _m (number of migrants per generation) from the spatial distribution of rare alleles suggest that both species have similar levels of interpopulation gene flow. These observations lead us to propose two competing hypotheses: the low level of geographic differentiation in D. simulans is due to its evolutionarily recent worldwide colonization and, alternatively, D. simulans has a narrower niche than D. melanogaster. Geographic variation data on different genetic elements (e.g., mitochondrial DNA, two-dimensional proteins, etc.) are required before these hypotheses can be adequately tested.We thank the Natural Science and Engineering Research Council of Canada for financial support (Grant A0235 to R.S.S.).     

Molecular biogeography and host relations of a parasitoid fly

Successful geographic range expansion by parasites and parasitoids may also require host range expansion. Thus, the evolutionary advantages of host specialization may trade off against the ability to exploit new host species encountered in new geographic regions. Here, we use molecular techniques and confirmed host records to examine biogeography, population divergence, and host flexibility of the parasitoid fly, Ormia ochracea (Bigot). Gravid females of this fly find their cricket hosts acoustically by eavesdropping on male cricket calling songs; these songs vary greatly among the known host species of crickets. Using both nuclear and mitochondrial genetic markers, we (a) describe the geographical distribution and subdivision of genetic variation in O. ochracea from across the continental United States, the Mexican states of Sonora and Oaxaca, and populations introduced to Hawaii; (b) demonstrate that the distribution of genetic variation among fly populations is consistent with a single widespread species with regional host specialization, rather than locally differentiated cryptic species; (c) identify the more‐probable source populations for the flies introduced to the Hawaiian islands; (d) examine genetic variation and substructure within Hawaii; (e) show that among‐population geographic, genetic, and host song distances are all correlated; and (f) discuss specialization and lability in host‐finding behavior in light of the diversity of cricket songs serving as host cues in different geographically separate populations.     

A population genetic evaluation of ecological restoration with the case study on <Emphasis Type="Italic">Cyclobalanopsis myrsinaefolia</Emphasis> (Fagaceae)

The ultimate goal of ecological restoration is to create a self-sustaining ecosystem that is resilient to perturbation without further assistance. Genetic variation is a prerequisite for evolutionary response to environmental changes. However, few studies have evaluated the genetic structure of restored populations of dominant plants. In this study, we compared genetic variation of the restored populations with the natural ones in Cyclobalanopsis myrsinaefolia, a dominant species of evergreen broadleaved forest. Using eight polymorphic microsatellite loci, we analyzed samples collected from restored populations and the donor population as well as two other natural populations. We compared the genetic diversity of restored and natural populations. Differences in genetic composition were evaluated using measurements of genetic differentiation and assignment tests. The mean number of alleles per locus was 4.65. Three parameters (A, A _R, and expected heterozygosity) of genetic variation were found to be lower, but not significantly, in the restored populations than they were in the natural populations, indicating a founder effect during the restoration. Significant but low F _ST (0.061) was observed over all loci, indicating high gene flow among populations, as expected from its wind-pollination. Differentiation between the two restored populations was smallest. However, differences between the donor population and the restored populations were higher than those between other natural populations and the restored populations. Only 13.5% and 25.7% individuals in the two restored populations were assigned to the donor population, but 54.1 and 40% were assigned to another natural population. The genetic variation of the donor population was lowest, and geographic distances from the restoration sites to the donor site were much higher than the other natural populations, indicating that the present donor likely was not the best donor for these ecological restoration efforts. However, no deleterious consequences might be observed in restored populations due to high observed heterozygosity and high gene flow. This study demonstrates that during the restoration process, genetic structures of the restored populations may be biased from the donor population. The results also highlight population genetic knowledge, especially of gene flow-limited species, in ecological restoration.     

Incongruent phylogeographies in spite of similar morphology, ecology, and distribution:Phippsia algida andP. concinna (Poaceae) in the North Atlantic region

Phylogeographic and taxonomic relationships among 54 North Atlantic populations of the snowbed grass genusPhippsia were investigated based on isozymes and genetically based morphological variation. The results support recognition of two distinct species,P. algida andP. concinna, the latter with at least two subspecies. Both of these self-fertilizing arctic pioneer species were genetic allotetraploids almost without intrapopulational variation. The two species showed strikingly different phylogeographies in the North Atlantic region in spite of their similarity in morphology, habitat ecology, mating system, and dispersal ecology, and in spite of their present cooccurrence in many geographic areas, sometimes even in the same snowbeds. The same electrophoretic multilocus phenotype was observed in all populations ofP. algida, and although this species showed considerable morphological variation, the variation was unstructured geographically. Thus,P. algida showed a pattern similar to other arctic species investigated in the North Atlantic region; it has probably dispersed postglacially across the sea barriers among Greenland, Svalbard, Iceland, and Scandinavia. In contrast,P. concinna was virtually fixed for different multilocus phenotypes in the three main geographic areas analyzed (S Norway, Svalbard, Greenland), corresponding to fairly distinct divergence in morphology. This pattern suggests absence of postglacial among-area dispersal ofP. concinna in spite of all of its similarities with its congener; it may have immigrated to the North Atlantic from different source areas and/or survived the last glaciationin situ.     

Studying genetics of adaptive variation in model organisms: flowering time variation in Arabidopsis lyrata

Arabidopsis thaliana has emerged as a model organism for plant developmental genetics, but it is also now being widely used for population genetic studies. Outcrossing relatives of A. thaliana are likely to provide suitable additional or alternative species for studies of evolutionary and population genetics. We have examined patterns of adaptive flowering time variation in the outcrossing, perennial A. lyrata. In addition, we examine the distribution of variation at marker genes in populations form North America and Europe. The probability of flowering in this species differs between southern and northern populations. Northern populations are much less likely to flower in short than in long days. A significant daylength by region interaction shows that the northern and southern populations respond differently to the daylength. The timing of flowering also differs between populations, and is made shorter by long days, and in some populations, by vernalization. North American and European populations show consistent genetic differentiation over microsatellite and isozyme loci and alcohol dehydrogenase sequences. Thus, the patterns of variation are quite different from those in A. thaliana, where flowering time differences show little relationship to latitude of origin and the genealogical trees of accessions vary depending on the genomic region studied. The genetic architecture of adaptation can be compared in these species with different life histories.     

青藏高原濒危植物唐古特大黄的遗传多样性

唐古特大黄(Rheum tanguticum)是中国传统的中藏药材,近几年由于生境的严重破坏,已濒临灭绝,并被列入濒危植物名单。为了探索唐古特大黄物种濒危的原因并保护其野生资源,本研究采集了9个居群87个个体的唐古特大黄样本,基于该物种的叶绿体基因trn S-G序列对其进行了遗传多样性研究。结果表明,唐古特大黄物种具有较高的遗传多样性水平(Ht=0.694),其中95.97%的遗传分化来自于居群间(G_(ST)=0.960),4.03%的遗传分化来自于居群内(Hs=0.028)。AMOVA分析也显示唐古特大黄居群间基因流较小(N_m=0.01),存在较高的遗传分化(F_(ST)=0.9631)。唐古特大黄较高的遗传多样性水平可能与该物种较长的进化史和生活史有关,居群间较高的遗传分化可能与高山地区特殊的地理环境和人类活动有关。根据研究结果,建议对唐古特大黄所有野生居群进行就地保护,同时收集种质资源开展异地繁殖工作,以保护物种的遗传多样性,维持其进化潜力。     

The impact of species-specific traits and phylogenetic relatedness on allozyme diversity inCarex sect.Phyllostachys (Cyperaceae)

Allozyme variation was examined inCarex sect.Phyllostachys (Cyperaceae) to study the effects of species-specific traits and phylogenetic relatedness on genetic structure. In contrast to the findings of similar studies, genetic variability in thePhyllostachys is poorly correlated with geographic range and putative differences in breeding systems (as inferred from morphology). This suggests that other patterns of evolution, colonization, and gene flow characterize the species found in this section. Fixation indices are negative for all populations suggesting that mechanisms such as disassortative mating and selection are maintaining heterozygous excess within populations. Closely related taxa often exhibit different genetic variability statistics. In some instances, however, clades (e.g.C. jamesii andC. juniperorum) display very similar levels of genetic variability despite marked differences in species-specific traits. Recent speciation coupled with the ability to maintain historical levels of variation within populations may be factors accounting for this phenomenon. Contrary to similar studies, species restricted to known glacial refugia have lower genetic diversity than those species that underwent mass migrations in response to deglaciation. Narrowly endemic species were found to partition their genetic diversity within, as opposed to between populations. The opposite trend was evident in wider ranging congeners.     

Gene flow and spatio-temporal genetic variation among sympatric populations of Tetranychus kanzawai (Acari: Tetranychidae) occurring on different host plants, as estimated by microsatellite gene diversity

We investigated spatio-temporal genetic variation in allele frequency and estimated gene flow among sympatric populations of Tetranychus kanzawai on different host plants by the use of microsatellite markers. In the analysis of spatial genetic variation, no isolation by distance was detected among the populations. Gene flow between populations on Hydrangea macrophylla and those on other host plants was relatively restricted, whereas the populations on Akebia quinata and Clerodendrum trichotomum were almost panmictic. Our study on temporal genetic variation showed (1) that population differentiation was slightly reduced during the period from April to May owing to frequent gene flow among populations; and (2) that population differentiation was greatly enhanced from May to October because of bottleneck effects. Genetic differentiation among T. kanzawai populations was caused by the effect of host plants rather than by the effect of geographic distance among populations, suggesting possibility of sympatric host race formation in this species.This revised version was published online in May 2005 with a corrected cover date.     

Variation in nectar volume and composition of Impatiens capensis at the individual,plant, and population levels

Although the volume and chemical composition of nectars are known to vary among plant species and to affect pollinator response to plants, relatively little is known of the variation in volume, and sugar and amino acid composition within species. We collected nectar from Impatiens capensis in a nested design: three flowers from each of three plants from each of three populations. This design enabled us to quantify variation within individual plants, among plants within populations, and among populations. Using high performance liquid chromatography, we analyzed the sugar and amino composition of the 27 flowers. Analysis of variance showed that none of the parameters (volume, concentrations of three sugars and 24 amino compounds) varied within individuals. Variation in nectar volume was not significant among plants but was nearly significant among populations. Of the three sugars detected (sucrose, glucose, and fructose), the only significant variation was that of sucrose among populations. Concentrations of 12 amino compounds varied significantly at the plant level while 7 amino compounds varied among populations. The results indicate that: (1) pooling of nectar samples from flowers of individual plants can be an acceptable methodology for those seeking to understand within species variation; (2) amino compounds appear to vary more than either volumes or sugar concentrations; (3) future studies should assess how much of the observed variation is due to genetic versus environmental differences; (4) additional studies should examine the geographic variation in nectar parameters and pollinators of I. capensis in order to assess the role different pollinators play in shaping nectar composition.     

Extent and structure of genetic variation in two colonising Diplotaxis species (Brassicaceae) with contrasting breeding systems

Genetic variation within two closely related Diplotaxis species was studied as indicated by isozymes and RAPDs. These species differ in their mating systems, their life forms, and in their evolutionary history, but both are successful colonisers. The diploid perennial D. tenuifolia is an outbreeder, the allotetraploid annual to biennial D. muralis is predominantly selfing. D. muralis was nearly devoid of genetic variation due to a young phylogenetic age and/or population history. Estimations of genetic variation within D. tenuifolia and F-statistics indicated random mating at the species and population level and confirms obligate outbreeding. However, influence of genetic drift relative to gene flow was high and mirrors colonisation processes as indicated by considerable heterogeneity across populations and the lack of correlation between population divergence and geographic distance.     

Large-scale geographic patterns of genetic variation in Melica nutans, a widespread Eurasian woodland grass

 The geographic distribution of allozyme variation within the Eurasian boreo-nemoreal woodland grass Melica nutans L. has been investigated together with a minor subset of other Melica species. Twenty alleles were found at nine polymorphic loci in M. nutans. Allelic richness was highest in areas central in the species' European distribution, i.e. in southern Fennoscandia. High population densities, reducing the effects of genetic drift, as well as accumulation of variation through long-distance gene-flow from different marginal populations, is proposed to explain high allelic richness in this area. Several alleles showed geographic patterns in distribution and frequency variation. However, these patterns were not congruent, e.g. some alleles appear to have migrated to northern Europe from the south-west whereas others may have spread from the east. Genetic distances between geographic regions, each consisting of 2–6 populations, were generally low between all Fennoscandian, Russian and Siberian regions, but much higher between western and continental European regions. On the population level, cluster analysis grouped populations from Siberia, Russia, coastal and lowland areas in Fennoscandia and British Cumbria into one subcluster whereas other subclusters contained mainly south-west European populations or populations from almost throughout the distribution range. A scenario with several independent glacial refugia in central Europe, south-western Siberia and possibly western Norway, and subsequent colonisation of Fennoscandia mainly from the east, but with some long-distance gene-flow from central Europe, is proposed. Received April 3, 2002; accepted September 17, 2002 Published online: December 11, 2002     

GEOGRAPHIC VARIATION AND PHENOTYPIC PLASTICITY OF NUMBER OF TRUNK VERTEBRAE IN SLENDER SALAMANDERS,BATRACHOSEPS (CAUDATA: PLETHODONTIDAE)

To understand the evolutionary significance of geographic variation, one must identify the factors that generate phenotypic differences among populations. I examined the causes of geographic variation in and evolutionary history of number of trunk vertebrae in slender salamanders, Batrachoseps (Caudata: Plethodontidae). Number of trunk vertebrae varies at many taxonomic levels within Batrachoseps. Parallel clines in number occur along an environmental gradient in three lineages in the Coast Ranges of California. These parallel clines may signal either adaptation or a shared phenotypically plastic response to the environmental gradient. By raising eggs from 10 populations representing four species of Batrachoseps, I demonstrated that number of trunk vertebrae can be altered by the developmental temperature; however, the degree of plasticity is insufficient to account for geographic variation. Thus, the geographic variation results largely from genetic variation. Number of trunk vertebrae covaries with body size and shape in diverse vertebrate taxa, including Batrachoseps. I hypothesize that selection for different degrees of elongation, possibly related to fossoriality, has led to the extensive evolution of number of trunk vertebrae in Batrachoseps. Analysis of intrapopulational variation revealed sexual dimorphism in both body shape and number of trunk vertebrae, but no correlation between these variables in either sex. Females are more elongate than males, a pattern that has been attributed to fecundity selection in other taxa. Patterns of covariation among different classes of vertebrae suggest that some intrapopulational variation in number results from changes in vertebral identity rather than changes in segmentation.