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     

Genetic variation in Scandinavian Anthericum liliago (Anthericaceae): allopolyploidy, hybridization and immigration history

 We investigated patterns of isozyme variation and the hierarchic structure of genetic diversity in 25 Scandinavian populations of the lilioid herb, Anthericum liliago. Isozyme data suggest that tetraploid A. liliago has an allopolyploid origin and that A. ramosum may be one of its diploid progenitors. Two populations contained known or suspected hybrids between A. liliago and A. ramosum. Isozymes show that one population from S Sweden contains both triploid (hybrid) and tetraploid individuals whereas a putatively hybrid Danish population represents diploid A. liliago. There is an overall northward and eastward decline in allelic richness in the tetraploid populations, with the highest numbers of alleles in Denmark and SW Sweden. This pattern is consistent with a progressive loss of allelic variation during the species' postglacial colonization of Scandinavia. The between-population component of genetic diversity is 4% (compared with 12% in diploid A. ramosum), the between-region diversity component is 7% and 89% of the total diversity is stored within populations. Received March 13, 2002; accepted September 24, 2002 Published online: December 11, 2002     

Microsatellite polymorphism and genetic differentiation in three Norwegian populations of Elymus alaskanus (Poaceae)

 Variation at seven microsatellite loci was investigated in three local E. alaskanus populations from Norway and microsatellite variation was compared with allozyme variation. The percentage of polymorphic loci was 81%, the mean number of alleles per polymorphic locus was 5.7 and expected heterozygosity was 0.37. An F-statistic analysis revealed an overall 48% deficit of heterozygotes over Hardy-Weinberg expectations. Gene diversity is mainly explained by the within population component. The averaged between population differentiation coefficient, F _st , over 7 loci is only 0.13, which accounts for only 13% of the whole diversity and was contrary to allozyme analysis. The mean genetic distance between populations was 0.12. However, a χ² -test showed that allele frequencies were different (p < 0.05) among the populations at 5 of the 7 loci. In comparison with the genetic variation detected by allozymes, microsatellite loci showed higher levels of genetic variation. Microsatellite analysis revealed that population H10576 possesses the lowest genetic variation among the tested three populations, which concur with allozyme analysis. The dendrogram generated by microsatellites agreed very well with allozymic data. Our results suggest that natural selection may be an important factor in shaping the genetic diversity in these three local E. alaskanus populations. Possible explanations for deficit heterozygosity and incongruence between microsatellites and allozymes are discussed. Received November 6, 2001; accepted April 24, 2002 Published online: November 14, 2002 Addresses of the authors: Genlou Sun (e-mail: Genlou.sun@STMARYS.CA), Biology Department, Saint Mary's University, Halifax. Nova Scotia, B3H 3C3, Canada. B. Salomon, R. von Bothmer, Department of Crop Science, The Swedish University of Agricultural Sciences, P.O. Box 44, SE-230 53, Alnarp, Sweden.     

Allozyme variation in Carex sect. Digitatae – Evidence of introgression, genetic distinctiveness and evolution of taxa

 A subset of populations of the European taxa of Carex sect. Digitatae, including Carex digitata, C. pallens, C. ornithopoda, C. pediformis ssp. rhizodes and C. humilis have been investigated for allozyme variation. The (presumably) distantly related C. pendula was used as a reference taxon. Carex digitata, C. pallens and C. ornithopoda on the one hand, and C. humilis and C. pediformis on the other, shared the majority of the alleles. Cluster analyses based on genetic distances grouped populations of C. digitata and C. ornithopoda in a mixed subcluster whereas all populations of C. pallens formed a subcluster distinct from the digitata/ornithopoda cluster. This in spite of the fact that C. ornithopoda is morphologically clearly distinct from C. digitata whereas C. pallens is barely distinguishable from the latter. Carex pediformis and C. humilis appeared not more genetically similar to C. digitata than to the reference taxon. It is concluded that 1) C. digitata, C. pallens and C. ornithopoda are closely related and most probably forms a monophyletic group, 2) C. pallens is a genetically distinct species, 3) C. pediformis and C. humilis are not closely related to the rest of C. sect. Digitatae, 4) C. pediformis and C. humilis are relatively closely related, 5) introgression do occur between the investigated species but not to such an extent that it can explain major geographic patterns of variation in C. digitata. Received December 21, 2001; accepted November 14, 2002 Published online: March 24, 2003     

Allozyme variation in Mexican species and classification of Datura (Solanaceae)

 Fifty natural Datura populations, belonging to eleven species (D. ceratocaula, D. discolor, D. inoxia, D. kymatocarpa, D. lanosa, D. metel, D. pruinosa, D. quercifolia, D. reburra, D. stramonium, D. wrightii) from Mexico and adjacent USA, were investigated using starch gel electrophoresis. A total of 64 alleles were scored at 17 loci (DIA1, DIA2, GOT1, GOT2, G6PDH, IDH, MDH1, MDH2, MDH3, ME, PGD1, PGD2, PGM1, PGM2, PHI, SAD, SOD). The heterozygosity among the species ranged from 0.166 (D. ceratocaula) to 0.276 (D. wrightii). Most genetic diversity was found within populations (average H_s=0.242), while values between populations are relatively low (average Dst=0.066, Gst=0.171). The analysis of the genetic distance suggested new taxonomic relationships among the species. Rather than supporting the conventional infrageneric classification with three sections, the results revealed that the herbaceous members of the genus Datura form four groups. One group included four of the eight species of the section Dutra and was more similar to the section Ceratocaulis than it was to the other group that contained the remaining taxa of Dutra. Received February 13, 2001 Accepted December 25, 2001     

Geographic pattern of genetic variation in photosynthetic capacity and growth in two hardwood species from British Columbia

Geographic patterns of intraspecific variations in traits related to photosynthesis and biomass were examined in two separate common garden experiments using seed collected from 26 Sitka alder (Alnus sinuata Rydb.) and 18 paper birch (Betula papyrifera Marsh.) populations from climatically diverse locations in British Columbia, Canada. Exchange rates of carbon dioxide and water vapour were measured on 2-year-old seedlings to determine the maximum net instantaneous photosynthetic rate, mesophyll conductance, stomatal conductance, and photosynthetic water use efficiency. Height, stem diameter, root and shoot dry mass and fall frost hardiness data were also obtained. Mean population maximum photosynthetic rate ranged from 10.35 to 14.57 μmol CO₂ m^–2 s^–1 in Sitka alder and from 14.76 to 17.55 μmol CO₂ m^–2 s^–1 in paper birch. Based on canonical correlation analyses, populations from locations with colder winters and shorter (but not necessarily cooler) summers had higher maximum photosynthetic rates implying the existence of an inverse relationship between leaf longevity and photosynthetic capacity. Significant canonical variates based on climatic variables derived for the seed collection sites explained 58% and 41% of variation in the rate of photosynthesis in Sitka alder and paper birch, respectively. Since growing season length is reflected in date of frost hardiness development, an intrinsic relationship was found between photosynthetic capacity and the level of fall frost hardiness. The correlation was particularly strong for paper birch (r=–0.77) and less strong for Sitka alder (r=–0.60). Mean population biomass accumulation decreased with increased climate coldness. These patterns may be consequential for evaluation of the impact of climate change and extension of the growing season on plant communities. Received: 12 July 1999 / Accepted: 24 November 1999     

Geographic isolation and evolution of Mediterranean endemic Cyclamen: insights from chloroplast trnL (UAA) intron sequence variation

 In this study we construct a phylogenetic hypothesis for the relatedness among disjunct subspecies of Cyclamen repandum and its two allopatric congeners, C. creticum and C. balearicum in order to examine the evolutionary divergence of currently isolated populations across the western Mediterranean. The most parsimonious phylogenetic tree obtained from sequencing the cpDNA trnL (UAA) intron suggests a major phylogeographic divide in southern Greece between two clades. The first clade comprises samples of C. repandum subsp. peloponnesiacum (from the Peloponnese) and C. creticum (from Crete). The second comprises samples of C. repandum subsp. repandum (from Croatia, Italy, southern France, Corsica, Sardinia and Sicily), C. repandum subsp. rhodense (from Rhodes and Kos) and C. balearicum (from the Balearic Islands and southern France). These data suggest that C. creticum has evolved in allopatry from C. repandum subsp. peloponnesiacum and that C. balearicum and C. repandum ssp. rhodense have diverged from C. repandum subsp. repandum at its western and eastern distribution limits. At one small site on Corsica, a population of C. repandum may have introgressed with relictual populations of C. balearicum. These divergence patterns illustrate how a phylogenetic perspective can be used to better understand the evolution of endemism in the Mediterranean flora. Received February 19, 2001 Accepted August 22, 2001     

Infraspecific genetic variation in Biscutella laevigata (Brassicaceae): new focus on Irene Manton's hypothesis

 Genetic variation in 42 populations throughout the range of Biscutella laevigata L. (Brassicaceae), a morphologically variable central European species, has been investigated by enzyme electrophoresis with three loci (Amy1, Amy2, and Gpi2). Genetic identities and the Fitch-Margoliash tree suggest differentiation into four regional groups: 1) a northwestern diploid group (northern France and northern Germany), 2) a northeastern diploid group (southern Germany, Upper Austria, northern Lower Austria, Poland, and Romania), 3) a central diploid group in southern Lower Austria corresponding to subspecies austriaca, and 4) a southern tetraploid group in Alpine areas of France, Germany, Switzerland, Austria, Italy, and Slovenia corresponding to subspecies laevigata. Geographically isolated diploid relic populations that are genetically depauperate are found in the NW and NE diploid groups. On the other hand, the diploid relic subspecies austriaca from the NE Prealps and Alps is highly variable. Subspecies laevigata appears to be a genetical autotetraploid with multiple origins involving several diploid progenitors (the NW diploids, subspecies austriaca and B. prealpina). Received April 6, 2001; accepted March 6, 2002 Published online: October 14, 2002 Addresses of the authors: Karin Tremetsberger (e-mail: k.tremetsberger@gmx.net), Christiane K?nig, Rosabelle Samuel, Tod F. Stuessy, Department of Higher Plant Systematics and Evolution, Institute of Botany, University of Vienna, Rennweg 14, A-1030 Vienna, Austria. Wilhelm Pinsker, Institute of Medical Biology, University of Vienna, Waehringerstra?e 10, A-1090 Vienna, Austria.     

多叶重楼遗传多样性的RAPD分析

应用RAPD技术检测了多叶重楼(Paris polyphyfzo)2个变种4个居群的遗传多样性,并与1个凌云重楼(P．cronquistii)居群进行了比较。选择的16个随机引物在5个居群中共检测到246个多态位点。在居群水平上,滇重楼2个居群的多态位点百分比(PP鳓分别为57．43％和54．67%,Shannon指数分别为0．3080和0．2830;七叶一枝花2个居群的PPB分别为56．33％和57．75%,Shannon指数分别为0、3080和0．3293。在变种水平上,滇重楼的PPB为75．14％,Shannon指数为0．3922,遗传分化系数(Gst)为0．3085;七叶一枝花的PPB为80．31％,Shannon指数为0．3992,遗传分化系数(Gst)为0．3726;在种的水平PPB达92．05%,遗传分化系数Gst达0．5151。聚类分析显示滇重楼和七叶一枝花有较近的亲缘关系,而与凌云重楼遗传距离较远。此结果从分子水平上支持了过去将滇重楼和七叶一枝花划分为1个种下2个变种的形态分类观点。     

Genetic structure and differentiation of wild and domesticated populations of Capsicum annuum (Solanaceae) from Mexico

 Genetic variation and structure of ten wild, three domesticated and one wild-cultivated populations of pepper (Capsicum annuum) from northwestern Mexico were studied in order to find out if the domestication process has reduced the genetic variation of the modern cultivars of this species. The analysis was based on 12 polymorphic loci from nine isozymes. Wild populations were sampled in different habitats along a latitudinal gradient of ca. 500 km. All populations had high genetic variation (i.e. wild: A = 2.72, P = 90.8%, He = 0.445; wild-cultivated: A = 2.50, P = 92.3%, He = 0.461; domesticated: A = 2.60, P = 84.6%, He = 0.408), indicating little genetic erosion in modern cultivars of pepper. Genetic diversity estimated by Nei's method showed that most genetic variation is found within, rather than among populations. However, genetic differentiation is greater among cultivated (G _ST=0.167) than among wild (G _ST=0.056) populations. Wild populations had an average genetic identity (I) of 0.952, indicating little differentiation and high gene flow (Nm=4.21) among these populations. Average genetic identity between wild and domesticated populations was of I=0.818, revealing that the domestication process has modified the genetic composition of commercial varieties of pepper. Changes in genetic composition among commercial varieties seem to have occurred in different directions, as indicated by the average value of I = 0.817 among these populations. The high level of diversity found in wild populations of C. annuum suggests that the wild relatives of cultivated peppers are a valuable genetic resource which must be conserved. Received May 5, 1999 Accepted October 30, 2000     

Allozyme variation within and among populations of onion (Allium cepa L.) from West Africa

Local germplasm of onion (Allium cepa L.) in West Africa is threatened by extinction. Sixteen populations of onion collected in five countries in West Africa were investigated for isozyme polymorphism using four polymorphic enzyme systems (ADH, MDH, 6-PGDH and PGI) among nine enzyme systems assayed. This is the first report on the genetic diversity of local landraces of onion. The inheritance of two dimeric enzyme systems PGI and MDH was demonstrated using F₂ progeny arrays. The PGI system revealed a single locus with three alleles, and the MDH system revealed three loci with four alleles. Four polymorphic systems revealed nine alleles (adh-a1 and a2, mdh-c1 and c2, 6-pgdh-a1 and a2, and pgi-a1, a2 and a3) in the 16 local populations observed. The mean number of alleles per polymorphic locus was 2.25, and 67% of the alleles were present in all populations. Allele 6-pgdh-a2 was present in only two landraces (from Niger and Nigeria); it is considered to be a rare allele (frequency approximately 2%). Among the 16 populations, within-population diversity was greater (90%) than between-population diversity (10%). Genetic distance analyses showed an aggregate of all populations except for two, which originated from Nigeria, an English-speaking country. Received: 24 August 1995 / Accepted: 26 February 2001     

Molecular differentiation of two vicariant neotropical tree species, Plathymenia foliolosa and P. reticulata (Mimosoideae), inferred using RAPD markers

 Plathymenia foliolosa and P. reticulata are two tropical tree species economically important for their high quality wood. P. foliolosa occurs mainly in areas of Atlantic Forest (a tropical forest) whereas P. reticulata occurs mainly in the Cerrado region (a savanna vegetation), both Brazilian biomes classified as hotspots for conservation priorities. In the present study RAPD markers were used to differentiate P. foliolosa from the vicariant species P. reticulata. Using 10 random primers it was possible to obtain 84 markers, of which 22 (26.2%) were present in just one of the two species. Nine of the ten primers were successful in discriminating between the species through the detection of diagnostic markers or band frequency differences. One of the populations of P. foliolosa, located in a region where the two species co-occur, presented several markers that were characteristic of P. reticulata, suggesting the occurrence of gene flow between the two species. Received September 3, 2001; accepted August 19, 2002 Published online: November 22, 2002 Addresses of the authors: Daniela Rodrigues Lacerda (dlacerda@dedalus.lcc.ufmg.br), Maria Bernadete Lovato (lovatomb@icb.ufmg.br), Maria Dolores Porto Acedo (acedomd@icb.ufmg.br), Dept. Biologia Geral, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, CP: 486, Belo Horizonte, MG, 31270-901, Brasil. José Pires de Lemos Filho (lemos@icb.ufmg.br), Dept. Botanica, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, CP: 486, Belo Horizonte, MG, 31270-901, Brasil.     

Genetic variation and population structure ofCarex breviculmis (Cyperaceae) in Korea

We determined the genetic diversity and population structures ofCarex breviculmis (Cyperaceae) populations in Korea, using genetic variations at 23 allozyme loci.C. breviculmis is a long-lived herbaceous species that is widely distributed in eastern Asia. A high level of genetic variation was found in 15 populations. Twelve enzymes revealed 23 loci, of which 11 were polymorphic (47.8%). Genetic diversity at the speciesand population levels were 0.174 and 0.146, respectively. Total genetic diversity (H_T = 0.363) and within-population genetic diversity (H_s = 0.346) were high, whereas the extent of the population divergence was relatively low (G_ST = 0.063). Deviation from random mating (Fis) within the 15 populations was 0.206. An indirect estimate of the number of migrants per generation(Nm = 3.69) indicated that gene flow was extensive among Korean populations of this species. Analysis of fixation indices revealed a substantial heterozygote deficiency in some populations and at some loci. Genetic identity between popu-lations was high, exceeding 0.956.     

The effects of two parasitoids on the life history and metapopulation structure of the intertidal snail Littoraria filosa in different-sized patches of mangrove forest

We describe differences in life history of the intertidal arboreal snail, Littoraria filosa, among patches of mangroves ranging in size from isolated trees to large stands several square kilometres in area. Recruitment of L. filosa occurred from mid spring (October) to early winter (June), recruits grew rapidly and copulating adults were found during the following September–April. Populations within large patches of forest were annuals; all or most individuals died between October–January (spring–midsummer). In contrast, those in smaller peripheral patches were more likely to survive over the summer but survival differed among patches and years. These differences in life history were caused by a parasitoid fly (genus Sarcophaga) that attacked L. filosa 10 mm and longer and was present in all large patches, but absent from, or rare, in smaller peripheral patches. Experimental introductions to isolated trees confirmed that the fly could kill L. filosa. Another sarcophagid parasitoid that attacked L. filosa from 4 to less than 10 mm long was also found in every patch. The combined effects of these parasitoids appear to determine the metapopulation structure of L. filosa. Most adults in large patches were killed by the larger fly during early summer. Summer recruits were often killed by the smaller fly within a month of settlement and when this happened effective recruitment of L. filosa was reduced to autumn. The planktotrophic larval stage of L. filosa lasts less than 1 month, so the source of autumn recruits to all patches must have been adults that survived the early summer, most of which were in small patches or on isolated trees. Consequently these ”peripheral sources” are likely to be important for persistence of the metapopulation of L. filosa. The results of this study demonstrate that metapopulation structure may be determined by complex interactions and that common models cannot be assumed to apply in all habitats. Received: 15 September 1999 / Accepted: 31 January 2000     

Evidence of gynodioecy and sex ratio variation in Wurmbea biglandulosa (Colchicaceae)

 A valuable approach to understanding the evolution of gender dimorphism involves studies of single species that exhibit intraspecific variation in sexual systems. Here we survey sex ratios in 35 populations of Wurmbea biglandulosa, previously described as hermaphroditic. We found pronounced intraspecific variation in sexual systems; populations in the northeastern part of the species' range were hermaphroditic, whereas other populations were gynodioecious and contained 2–44% females. Populations with lower annual rainfall were more likely to be gynodioecious, supporting the view that gender dimorphism evolves more frequently in harsher environments. In gynodioecious populations, however, female frequency was not related to either annual rainfall or habitat, indicating that other factors are important in determining sex ratio variation. Females had smaller flowers and shorter stems than did hermaphrodites, potentially providing a basis for resource compensation. A female fecundity advantage may contribute to the maintenance of females in populations because females produced more ovuliferous flowers and had more ovules per flower than did hermaphrodites. Received March 2, 2001 Accepted February 25, 2002     

微生境下羊草两种生态型种群的遗传多样性及遗传分化——等位酶分析

采用水平淀粉凝胶电泳技术,应用等位酶分析方法测定了松嫩平原南部微生境条件下羊草灰绿色和黄绿色两种生态型９个种群的遗传多样性和遗传分化程度。羊草种、种群和生态型水平都维持较高的遗传多样性。两种生态型之间有明显的遗传多样性差异及遗传分化。灰绿型和黄绿型的多态位点百分率分别为６７．３％和５７．１％,等位基因平均数分别是２．２和１．９,期望杂合度分别为０．３４８和０．２９８。两种生态型种群之间平均遗传距离     

影响披碱草属植物遗传分化和亲缘关系的地理因素分析

用微卫星标记法对产自中国青藏高原和蒙古高原地区9种披碱草属（Elymus L．）植物的40个居群进行遗传变异和亲缘关系分析,居群问遗传相似系数为0．020～0．957,居群间遗传距离与地理因素关系密切。以垂穗披碱草（Elymus nutctns Griseb．）为例,采用二维和三维Mantel检验,对遗传距离和地理因素间的关系（纬度、经度和海拔）作进一步分析,结果表明,地理位置（纬度和经度）是影响垂穗披碱草居群微卫星遗传差异的最重要因素（R=0．478;P＜0．05）,其次是海拔（R=0．258;P＞0．05）。单一地理位置因素（纬度或经度）与遗传距离的相关性并不明显,但从单一地理位置因素与遗传距离的相关系数、可信度区间来看,纬度对遗传分化的影响程度大于经度。     

Studies in the Melica ciliata – complex: 1. Distribution of allozyme variation within and among individuals, populations and geographic regions

Allozyme variation in 83 European populations of the Melica ciliata/ transsilvanica species complex has been investigated. In spite of a diploid chromosome number, most loci were duplicated and patterns of within-population variation were not explainable unless apomixis or uncommon patterns of chromosomal segregation were assumed. Allelic richness and genetic diversity were very high but most of the variation (55-98%) was due to population differentiation. Geographic patterns were analysed both using phenetic, based on allele frequencies within populations, and cladistic, based on multi-locus genotypes, methods. Both methods congruently recognised three large groups of populations with discrete geographic distributions, in addition to a number of smaller groups. Of the three main groups, one was distributed in eastern and central Europe, one was restricted to western Europe north of the Alps and one was distributed in Iberia, Brittany and along the western shores of the Mediterranean. These groups are likely to have had separate histories but there appears to have been significant gene-flow among them. Among the less distinct groups, it was suggestive that some, e.g. the group containing all populations from the disjunct distribution in the Baltic basin, have been formed by introgression between the three major groups, whereas other deviating populations may represent additional distinct entities.