Genetic variation and clonal diversity in four clonal sedges (Carex) along the Arctic coast of Eurasia

We studied the structure of genetic variation (at both ramet- and genet-level) and clonal diversity within and among populations in the four closely related arctic clonal sedges Carex bigelowii, C. ensifolia, C. lugens and C. stans by use of allozyme markers. Compared to other sedges and arctic plants, the studied taxa all had high levels of genetic variation, both within populations and taxa. These taxa contained most of the total gene diversity (H(T)) within populations and a small part of the diversity among populations (G(ST) ranged 0.05--0.43). Carex bigelowii had genetic variation (H(S) = 0.173, mean for populations) at a comparable level to other outbreeding arctic plants and to other widespread, rhizomatous and mainly outbreeding Carex species. In contrast, C. ensifolia (H(S) = 0.335), C. lugens (H(S) = 0.339) and C. stans (H(S) = 0.294) had within-population variations that were higher than in most other studied Carex species and for arctic plants in general. Genetic variation was not related to any tested environmental variable, but it was lower in areas deglaciated only 10,000 years BP compared to areas deglaciated 60,000 years BP or not glaciated at all during the Weichselian. All the populations were multiclonal, except for two populations of C. stans that were monoclonal. In contrast to genetic variation, clonal diversity decreased with latitude and did not differ between areas with different times of deglaciation. In accordance with previous studies, C. bigelowii and C. lugens were found to be outbreeding, while C. ensifolia and C. stans had mixed mating systems.     

Variation in morphological characters and habitat requirements in Carex capitata and Carex arctogena (Cyperaceae) in Norway and Sweden

Norwegian and Swedish populations of Carex capitata s. lat. were studied for variation in morphological characters. Canonical variates analysis (CVA) arranges populations in two discrete groups, corresponding to C. capitata s. str. and C. arctogena , respectively. In principal components analysis (PCA) of the total material a small overlap could be seen. However, separate PCA: s for the two regions in Scandinavia where the two taxa are sympatric show perfect separation. Student's t-test revealed significant differences in all morphological characters studied. Studies of companion plants, analysed with detrended correspondence analysis (DCA), indicate that the taxa have rather similar habitat demands in Oppland, Central Norway, although C. capitata s. str. seems to prefer wetter, and C. arctogena drier sites, respectively. Since the two taxa are distinct in morphology, they are best treated as separate species. The name Carex capitata is lectotypified.     

Defense mechanisms against grazing: a study of trypsin inhibitor responses to simulated grazing in the sedge Carex bigelowii

Trypsin inhibitors have been suggested to constitute an inducible defense in the sedge Carex bigelowii , and some former studies suggest that this might be a cause for the cyclic population dynamics in many alpine and arctic small mammals, for example lemmings ( Lemmus lemmus ). We investigated this further by using a method of simulated grazing (clipping) at different intensities, in three different habitats with varying resource availability, with different harvest times (hours after clipping), and two different stages of ramets (reproductive/vegetative) in a study from the Swedish mountain range. Our results do not indicate that C. bigelowii has an inducible defense constituted by an increase in trypsin inhibitor activity (TIA), but rather that the amount of soluble plant proteins (SPP) is lowered in wounded plants. The responses were somewhat different in the three habitats, with ramets growing in the marsh showing the highest ratio of TIA to SPP, due to low amounts of SPP. We did not find any significant effects of harvest time, or of the stage of the ramet that could support the hypothesis of an inducible defense. To conclude, we could not find any evidence for an inducible defense consisting of trypsin inhibitors in Carex bigelowii ramets, but we did find variations in the amount of SPP that may have nutritional consequences for herbivores.     

Molecules and morphology in concert: tests of some hypotheses in arctic Potentilla (Rosaceae)

We developed a combined molecular and morphological approach to unravel complex variation at low taxonomic levels, exemplified by some arctic members of Potentilla. Twenty-one populations from Svalbard were analyzed for random amplified polymorphic DNAs (RAPDs) and 64 morphological characters to test the hypotheses that (1) the P. nivea complex (section Niveae) consists of three taxa (P. chamissonis, P. insularis, and P. nivea), (2) three "eco-morphotypes" in P. pulchella (section Multifidae) should be considered different taxa, and (3) P. insularis originated as an intersectional hybrid (Niveae × Multifidae). Twenty-two RAPD multilocus phenotypes were observed in the 136 plants analyzed based on 35 markers. Three fairly distinct groups of RAPD phenotypes were identified in the P. nivea complex based on multivariate analyses and an analysis of molecular variance (AMOVA; 77.6% among-group variation). The variation within the P. nivea complex was more or less continuous in multivariate analyses of the morphological data. We identified, however, several individual morphological characters that separated unambiguously among the three groups of RAPD phenotypes, revealing that these groups correspond to the previously hypothesized taxa. Many identical RAPD multilocus phenotypes were observed in the "eco-morphotypes" of P. pulchella, suggesting that its conspicuous morphological variation is caused by plasticity or by genetic variation at a small number of loci. The hypothesis of the hybrid origin of P. insularis was not supported by the RAPD data. Overall, very little RAPD variation was observed within populations of the four taxa (2.1-16.7% in AMOVA analyses; average genotypic diversity, D, was 0.10-0.30). We conclude that detailed, concerted analysis of molecules and morphology is a powerful tool in low-level taxonomy.     

黄河下游湿地芦苇形态变异研究

 以水盐为主导生态因子,分析了黄河下游湿地芦苇(Phragmites australis)种群形态变异的水平与格局。对基径、株高、叶长、叶宽、节间长、节间数和穗长7个形态特征在15个种群中的变异分析表明：种群内个体间形态差异极显著,7个形态特征在种群内的变异系数从大到小依次为节间长（0.284 6）、叶宽（0.253 6）、穗长（0.244 9）、叶长（0.208 5）、基径（0.187 5）、节间数（0.176 3）、株高（0.165 7）; 7个形态特征在种群间的变异度从大到小依次为株高、叶长、节间长、叶宽、基径、节间数、穗长,但种群间形态的显著差异主要存在于滨州种群（BZH）、淡水种群（ZHG、DPH、NYH,土壤总盐度<0.1%）、盐生种群（HS01、HS02、HS03、HS04、HS05、DWL、KD、GD、LZH、LJ、DLH,土壤总盐度>0.3%）之间,在来自相似生境的种群间形态差异普遍不显著;7个形态特征均与水盐联合影响因子显著相关;以形态特征为基础的聚类分析将研究的15个种群分为显著不同的３类。因此,根据黄河下游湿地芦苇形态变异规律和分化特点,建议将该地区芦苇分为盐生芦苇、淡水芦苇、巨型芦苇3个形态类型。     

红穗苔草对天鹅洲湿地淹水时间变化的形态学响应

以天鹅洲湿地的优势种红穗苔草（Carex argyi）作为研究对象,以淹水持续时间作为水位变化的衡量指标,选取3个淹水持续时间不同的位点,研究了淹水时间长短对红穗苔草形态指标的影响。通过野外原位观测试验发现,红穗苔草的株高及叶片的长、宽对淹水时间的长短反应敏感,而单个分株上的绿叶数对淹水时间的长短不敏感。适度的淹水可以显著增加叶片的长度和株高,长期淹水和相对干旱的环境不利于叶片的伸长和株高的增加。短时间淹水处理株高一直维持较高的水平,并保持相对的稳定。淹水会显著增加叶片的宽度,长时间淹水的处理效果更明显,且维持时间较长。长期淹水会提高红穗苔草地上部生物量的分配比例。尽管不同水位点土壤氮含量有显著差异,但是氮对红穗苔草形态指标并无显著的影响。以上实验结果表明,红穗苔草可以通过调节其叶片的形态及地下和地上部分生物量的分配比例来适应淹水胁迫的不利环境,有利于其在水位波动条件下种群的稳定和维持。     

同质园栽培下的湖南鱼腥草居群叶表型性状多样性分析

将采自湖南的20个鱼腥草(Houttuynia cordata Thunb.)居群栽培于同质园内,通过测定9个叶片表型性状和6个叶片相对性状,采用方差分析、变异系数、相关性分析、主成分分析以及聚类分析等方法,对鱼腥草居群叶片形态的变异特征进行研究。结果表明:(1)同质园栽培的20个鱼腥草居群叶片15个表型性状均存在显著差异,居群间差异大于居群内差异;(2)居群内叶性状的平均变异系数变化幅度为5.63%~9.08%,说明居群内多样性程度较低,其中株洲攸县居群(P16)变异系数最高(9.08%),怀化溆浦居群(P3)变异系数最小(5.63%);(3)15个叶性状的平均变异系数变化幅度为3.71%~10.28%,说明各性状的多样性程度也较低,其中叶面积变异系数最大(10.28%),叶宽与叶中部宽之比(W_2/W_4)的变异系数最小(3.71%);(4)相关性分析显示,叶面积与叶宽、叶中部宽、叶长、叶基至叶宽处的距离、叶柄长都呈现出显著正相关,与叶尖夹角呈显著负相关,而与叶基夹角关系不大;(5)叶片15个性状中前3个主成分的贡献率为83.66%,聚类分析可将20个居群划分为心形、小心形、狭心形和大心形4类,呈现出间断分布的地理格局;(6)同质园实验说明叶表型性状多样性主要是由遗传物质决定的,对其多样性分析能为优良种质资源的选育提供理论依据。     

高山红景天形态分化与生存环境关系的研究

对采自黑龙江省平顶山和吉林省长白山的14 个天然居群高山红景天的分株数、株高、基径、叶长和叶宽5 个形态性状进行了比较分析, 探讨了高山红景天适应环境的形态变异机制。单因素方差分析F 检验表明, 除分株数外的4 个形态性状在14 个居群中均表现为差异极显著。对这4 个性状的多重比较分析表明, 各形态性状在不同生态环境下表现出一定程度的分化。对14个天然居群形态性状的聚类分析表明, 高山红景天的形态变异与其生态环境间存在着明显的相关关系。     

Biosystematic Studies on Adenophora potaninii Korsh. Complex (Campanulaceae). Ⅰ. Phenotypic Plasticity

Phenotypic plasticity is the environmental modification of genotypic expression and an important means by which individual plants respond to environmental heterogeneity. The study of phenotypic plasticity in the genus Adenophora, which is very complicated taxo nomically because of great morphological variation, proves to be helpful in both investigating the phenotypic variation so as to evaluate potential taxonomic value of their characters and providing important sources of information on the variation, adaptation and evolution of the genus. Twenty-three populations representing all the six species in Adenophora potaninii complex were transplanted into the garden. Of them six populations were selected for study ing their performance in the field and in the garden, in addition to cultivation experiment under different treatments. The results show that there exists considerable developmental plasticity in some leaf, floral and capsule characters. In particular, the leaf shape and length of calyx lobe display significant developmental variation with the maximum being three times as great as the minimum, which is noteworthy because they were previously considered as diagnostic. The characters of root, caudex, stem and inflorescence are found to be very plastic, especially the root diameter, the number of stems, stem height and inflorescence length with great environmental plasticity. In addition, the populations from different habi tats show distinct amounts of plasticity. On the contrary, the characters of leaf, floral, cap sule and seed are less influenced by environments. It seems that the considerable variation in the characters of leaf is attributed mainly to genetic differences. Finally, the phenotypic plasticity of morphological characters of A. potaninii complex and its taxonomic significanceis discussed.     

泡沙参复合体（桔梗科）的物种生物学研究：Ⅰ表型的可塑性

表型可塑性是生物变异中由环境引起的一种变异,是植物适应的一种重要方式。对沙参属这样一个形态上复杂多变、分类上很难处理的类群,研究其表型可塑性不仅为探讨性状变异、判断其系统学意义及选择可靠的分类性状提供了有益的资料,而且有助于揭示沙参属植物变异、适应和进化的机制。本文从泡沙参复合体中选择了6个居群,利用播种和移栽试验,通过对不同个体和居群在一致条件下的表现及野外和移栽后的对比,对根、茎、叶、花和果等形态性状的表型可塑性进行了初步的观测分析。结果表明,一些叶片、花部和果实性状具有较大的发育可塑性,尤其是叶形、花萼裂片不仅发育变化大,而且随发育过程定向变化。环境可塑性较大的性状主要是根、茎、花序分枝等性状,而叶片、花部、果实和种子性状的环境饰变能力都较小。最后,对泡沙参复合体形态性状的变异从发育可塑性和环境可塑性的角度进行了讨论。     

Phenotypic plasticity of Vaccinium meridionale (Ericaceae) in wild populations of mountain forests in Colombia

Vaccinium meridionale is a promising crop for the Andean region of South America and is currently available only in the wild. Spontaneous populations of this plant are found across the Colombian mountains, but very few published records on this plant morphology are available. A zonification study of V. meridionale was conducted in four principal areas of a low mountain forest of Colombia (Provinces of Boyacá, Cundinamarca, Santander and Nari?o) in 2007. A total of 20 populations and 100 plants of V. meridionale were individually characterized and surveyed, using a list of 26 characters of morphological variables (9 quantitative and 17 qualitative characters). Our results indicated that natural populations of V. meridionale might be found in the tropical forest under a highly heterogeneous climate and microclimate conditions, at different mountain regions between 2 357 and 3 168masl. The shrubs of V. meridionale exhibited a high level of intra-population variation in several quantitative (plant height, stem diameter) and qualitative (growth habit, ramification density, presence of anthocyanins in stems) morphological characters, suggesting an environmentally induced phenotypic plasticity. Plant height, stem diameter and foliar density were the most variable morphological traits, with coefficients of variation higher than 50%. However, several quantitative characters of its reproductive potential, such as berry dimensions, rachis length and number of flowers per inflorescence, resulted with low plasticity with coefficients of variation lower than 30.2%, indicating that these characters were genetically determined. The highest correlation coefficients (p < 0.05) resulted to be between fruit length and fruit width (0.90), leaf length and leaf width (0.78), plant height and stem diameter (0.60), and inflorescence length and flowers number per inflorescence (0.57). The results suggest that an important genetic resource exists for this species in the wild. Low variation in fruit size, which constitutes a target trait for plant breeders, could be useful for selection of cultivars of V. meridionale. The results of this study could also be applied in conservation programs aimed to protect these diverse populations in the mountain forests of Colombia.     

Leaf anatomy of Carex humilis does not correlate with orographic,geological and bioclimatic habitat conditions in C&SE Europe

This paper presents the results of a multivariate morphometric study of leaf anatomical characters in different, geographically very distant populations of taxon Carex humilis from Hungary, Romania, Serbia, Bosnia & Herzegovina, Montenegro and Albania with the aim to reveal the trends of population differentiation. Analyses were performed on the cross-section of 173 leaves collected from 12 populations. In order to establish the overall morphological variation and relationships between individuals from all populations, principal component analyses (PCA) and canonical discriminant analysis (CDA) have been done. The UPGMA clustering analyses based on leaf anatomical characters and habitat climatic characteristics were carried out to explore whether the observed anatomical differences are a result of adaptive responses. Regression analysis (linear regression) was performed to identify the level of correlation between leaf anatomical characters and basic orographic, geological, and bioclimatic habitat characteristics. Quite unexpectedly, most of observed groups are formed of geographically very distant populations which are living in extremely different climatic and geological conditions, indicating that general anatomical differentiation in Carex humilis in C&SE Europe cannot be explained by the environmental impacts, and basically do not represent an adaptive response to different climatic or geological condition.     

Integrating environmental,molecular, and morphological data to unravel an ice‐age radiation of arctic‐alpine Campanula in western North America

Many arctic‐alpine plant genera have undergone speciation during the Quaternary. The bases for these radiations have been ascribed to geographic isolation, abiotic and biotic differences between populations, and/or hybridization and polyploidization. The Cordilleran Campanula L. (Campanulaceae Juss.), a monophyletic clade of mostly endemic arctic‐alpine taxa from western North America, experienced a recent and rapid radiation. We set out to unravel the factors that likely influenced speciation in this group. To do so, we integrated environmental, genetic, and morphological datasets, tested biogeographic hypotheses, and analyzed the potential consequences of the various factors on the evolutionary history of the clade. We created paleodistribution models to identify potential Pleistocene refugia for the clade and estimated niche space for individual taxa using geographic and climatic data. Using 11 nuclear loci, we reconstructed a species tree and tested biogeographic hypotheses derived from the paleodistribution models. Finally, we tested 28 morphological characters, including floral, vegetative, and seed characteristics, for their capacity to differentiate taxa. Our results show that the combined effect of Quaternary climatic variation, isolation among differing environments in the mountains in western North America, and biotic factors influencing floral morphology contributed to speciation in this group during the mid‐Pleistocene. Furthermore, our biogeographic analyses uncovered asynchronous consequences of interglacial and glacial periods for the timing of refugial isolation within the southern and northwestern mountains, respectively. These findings have broad implications for understanding the processes promoting speciation in arctic‐alpine plants and the rise of numerous endemic taxa across the region.     

Genet age in marginal populations of two clonal Carex species in the Siberian Arctic

During a Swedish-Russian expedition to northern Siberia 1994, we sampled two marginal populations of two Carex species at two high arctic sites ( C, stans Drej. on Faddeyevsky Island and C. ensifolia V. Krecz ssp. arctisibirica Jurtz. at north-eastern Taymyr Peninsula), both north of previously documented localities in that areas for the two species. These populations were composed of a few distinct patches of ramet colonies, some of them shaped like fairy rings with dead centres. We measured the size of all colonies and collected samples for detailed morphometric analyses of rhizome growth. By using RAPD (random amplified polymorphic DNA) analysis we established that the largest colony at each site consisted of a single genet, based on 41 polymorphic bands amplified with three primers. Pooled samples from each of two additional colonies of C. stans on Faddeyevsky Island were analysed and showed that clones of the same species at the same site were relatively dissimilar (Dice's similarity index 0.26 0,43), We then assumed that each ramet colony represented a single genet. Based on the morphometric data, we developed a deterministic growth model that simulates the clonal growth of these species and enabled estimates of the time since establishment of the genets. The estimated age of the five C. Stans clones varied from 17 to 154 yr and the age of the two C. ensifolia ssp, arctisibirica clones was well over 3000 yr.     

Trans-Atlantic dispersal and large-scale lack of genetic structure in the circumpolar, arctic-alpine sedge Carex bigelowii s. l. (Cyperaceae)

Paradoxically, several of the ecologically most important plant groups in the Arctic are little understood in terms of taxonomy and biogeographic history. The circumpolar Carex bigelowii s. l. (Cyperaceae) is abundant in the Arctic and is one of the most complicated arctic plant groups. While its ecology and population genetics have been extensively studied, its taxonomy is largely unexplored. We analyzed the large-scale geographical structuring of amplified fragment length polymorphisms (AFLPs) covering most of the distribution range. We detected high levels of genetic variation, most (66%) within populations, and a fairly weak genetic structure. Only the Central Asian populations, referred to as C. orbicularis, were strongly divergent. For the remaining populations, Bayesian clustering separated three distinct clusters (one European, one amphi-Atlantic, and one broadly amphi-Beringian), probably reflecting different major glacial refugia and recent transoceanic dispersal. The isolated central European populations were most closely related to those from a larger distribution area in northern Europe. Differences in genetic diversity suggest that the Alpine and Tatra populations have experienced strong bottlenecks, whereas the Krkono?e population may have been part of a continuous distribution area during the cold stages of the Pleistocene. Finally, we discuss the relevance of our results for a uniform, range-wide taxonomic concept.     

沙生植物沙鞭不同居群形态变异研究

以沙鞭(Psammochloa villosa Bor.)20个自然居群为研究对象,对其株高、花序长度、旗叶长度、旗叶宽度和小穗长度等12个表型性状进行形态变异研究。结果显示,12个性状群体间F值为1.832~8.958,达到显著或极显著水平,表明沙鞭不同居群表型性状存在广泛变异,且变异程度各不相同;主成分分析结果显示,前4个主成分代表了沙鞭形态多样性的82.277%,其中旗叶长度、旗叶宽度、颖片长度、小穗长度等是造成不同居群表型性状差异的主要因素;依据欧式距离对参试居群进行的UPGMA聚类分析结果表明,当遗传距离为20.5时,可以将20个沙鞭野生居群划分为两类,且各表型性状并没有依居群地理分布而聚类。     

Architecture of coastal and desert Encelia farinosa (Asteraceae): consequences of plastic and heritable variation in leaf characters

The shrub Encelia farinosa (Asteraceae) exhibits geographic variation in aboveground architecture and leaf traits in parallel with environmental variation in temperature and moisture. Measurements of plants occurring across a natural gradient demonstrated that plants in desert populations produce smaller, more pubescent leaves and are more compact and branched than plants in more mesic coastal environments. This phenotypic variation is interpreted in part as adaptive genetic differentiation; small size and pubescence reduce leaf temperature and thus increase water-use efficiency but at the cost of lower photosynthetic rate, which results in slower growth and more compact growth form. We explored the basis of phenotypic variation by planting seed offspring from coastal and desert populations in common gardens in both environments. Phenotypic differences among populations persisted in both common gardens, suggesting a genetic basis for trait variation. Desert offspring outperformed coastal offspring in the desert garden, suggesting superior adaptation to hot, dry conditions. Herbivore damage was greater for all offspring in the coastal garden. Phenotypic characters also showed plastic responses; all offspring had smaller, more pubescent leaves and more compact growth form in the desert garden. Our results confirm that leaf size and pubescence are heritable characters associated with pronounced variation in plant architecture.     

贵州梵净山亮叶水青冈解剖特征的生态格局及主导因子分析

使用具有不同生物学意义的生态因子为环境变量,应用多元统计方法,分析了贵州梵净山的亮叶水青冈（Ｆａｇｕｓ ｌｕｃｉｄａ Ｌ．）叶片及木材形态解剖特征的生态格局和影响其变化的主导生态因子。结果表明,随着温度要素的增加,叶片的所有解剖特征、叶表面的气孔面各以及木材的导管直径、纤维长度均显示下降趋势,而叶表面的气孔器密度、木材的导管频率和单列射线高度呈上升趋势;随着寒冷指数（负积温）和水分因子（年降水和水     

Quantitative variation within and among populations of Arabis serrata Fr. & Sav. (Brassicaceae)

Phenotypic and genetic variation within and among eight populations of Arabis serrata are documented in this study. This species shows great morphological variation throughout its geographical distribution in Japan. Plants are located in habitats with different types of soils and degree of disturbance. Half-sibs progenies from eight populations were collected and cultivated in a garden experiment. Nine morphological traits representing size and shape of rosette leaves were recorded. Univariate analyses of measured traits showed that phenotypic means differed among populations for all characters. Leaves of plants from disturbed habitats had the longest petioles (lanceolate) and plants from limestone habitats showed the most roundness in leaf shape (ovate). The northernmost populations always revealed the smallest leaves. Multivariate principal component analyses also showed that leaf shape and size varied among populations. The first three principal components explained 98.5% of the variation. Coefficients of variation had a very wide range and differed from one population to another. Some traits (e.g. leaf width/leaf length ratio) were consistently less variable while others (e.g. leaf area and petiole length) were more plastic. All traits had significant genetic variance in all populations. Intra-class correlation coefficients differed for most of the traits and each population presented a different range of values. Most of the leaf traits were intercorrelated in all the populations studied, although some populations were integrated more tightly for some traits. Populations of A. serrata are differentiated in phenotypic means but they display a mosaic of traits with slight morphological differences in each locality (i.e. a quantitative genetic variation). Some traits can be correlated to the habitats that they occupy but for some of them it is difficult to assign an actual adaptive value.     

Taxonomy of the Carex flava complex (Cyperaceae) in Finland

The morphological variation in the Carex flava complex (sect. Ceratocystis ) in Finland, particularly taxa described by Alvar Palmgren as endemic to northern Europe, was studied using numerical analyses. Discriminant analyses were used to maximize the separation of the taxa. However, the separation of the taxa described by Palmgren was weak. The taxonomic treatment of the C. flava complex in Finland based on morphology, ecology, distribution and hybridization of the taxa is presented. Four species are recognized: Carex flava (the lectotype is selected here), C. lepidocarpa, C. demissa and C. viridula. C. lepidocarpa includes ssp. lepidocarpa and ssp. jemtlandica . In the polymorphic C. viridula , three taxa are recognized: var. viridula , var. bergrothii and var. pulchella . A key to the taxa is presented. The distribution of the taxa in Finland is presented, and their ecology is briefly discussed.