Morphometrics and character state recognition for cladistic analyses in theBletia reflexa complex (Orchidaceae)

Different criteria have been suggested for decisions on character state recognition for morphological characters showing continuous variation from taxon to taxon. A method proposed byMishler andDe Luna seeks to recognize as many character states as the groups of taxon means that are revealed by an analysis of variance (ANOVA) followed by multiple range tests. We apply this method as a guide in decisions of selecting taxonomic characters in species of theBletia reflexa complex. Seven species of the complex, represented by 156 individuals belonging to 54 populations, were the terminal taxa of the study. Character state codes were assigned to 20 characters. These characters together with 12 non-quantitative characters were incorporated into cladistic analyses. Results indicate that the complex is monophyletic only under one outgroup combination.     

Orchids from the Greater Antilles I: a new species of Bletia

Bletia antillana (Orchidaceae), a species restricted to eastern Cuba, is described and illustrated. This species is similar to Bletia purpurea but differs by the following: 1) the leaves always present during anthesis; 2) flower is semi-erect; 3) flower parts larger; 4) dorsal and lateral sepals straight; 4) petal curved, enclosing the column and the lateral lobes of the lip; 5) lip whitish at base, rose-purple at the apex of the lateral lobe, and yellow at the callus; and 6) lip obovate, with a quadrangular midlobe curved downward.     

Molecular and morphological analyses of EuropeanAconitum species (Ranunculaceae)

The systematics of the yellow-floweredAconitum lycoctonum species complex (Ranunculaceae) has long been considered difficult because of high morphological variability and hypothesized hybridization. To elucidate the systematic value of these taxa, and to study their phylogeny in a broader context, we studied all species of the subgenusLycoctonum from Europe and the Caucasus Mountains and the European species of the subgen.Aconitum. We sequenced the ITS region and the chloroplast intergenic spacerpsbA-trnH and conducted parsimony analyses. We also measured morphological characters from 19A. lycoctonum populations. The two subgenera in Europe are clearly separated but hybridization between subgenera might have taken place in the past. Sequence variation withinA. lycoctonum and among all taxa of subgenusLycoctonum was very low and probably the result of recent speciation, leading to one morphologically highly variable species in which color morphs could be seen as subspecies. There was high morphological variability within and among populations and the morphological characters have no value as systematic characters.     

The evolution of novel herbicide tolerance in a noxious weed: the geographic mosaic of selection

Understanding how genetic variation is organized over geography has long been of interest to evolutionary biologists given that traits can vary within and among populations, across regions, and at continental or global scales. The pattern of regional variation can have an important impact on trait evolution at the local or population level. Using a common garden, we asked whether a geographically variable mosaic of tolerance to the widely applied herbicide RoundUp^® existed in two closely related co-occurring species of morning glory, Ipomoea purpurea and I. hederacea. We assayed RoundUp tolerance in over 1,700 plants representing 290 families from 29 populations in the southeastern United States. Our findings suggest that the two species of morning glory partition their respective levels of genetic variation for tolerance to glyphosate differently. Variation for tolerance in I. purpurea appears to exist among maternal lines and regions, whereas in I. hederacea, variation in tolerance existed only among populations. In addition, we find a significant hotspot of tolerance or positive spatial aggregation of this trait on a local scale in I. purpurea populations from the Coastal Plain. This suggests that either similar regimes of selection or gene flow between populations can produce a geographic mosaic of tolerance. These results highlight the fact that the genetic variation underlying an adaptive trait can exist at many different scales, whether it be within- or among-populations, among geographical ‘hotspots,’ or among distinct ecological regions. Given these results, the partitioning of genetic variation should be considered before making predictions about an adaptive trait’s evolutionary trajectory.     

Phenetic studies in Nuphar Sm. (Nymphaeaceae):variation in sect. Nuphar

Previous taxonomic assessments of the hydrophytic genus Nuphar Sm. (Nymphaeaceae) have inadequately considered the range of morphological variation existing in the genus. A comprehensive statistical analysis of morphological features among the recognized taxa of Nuphar is presented here, first using 10 fruit characters to delimit the separation of two infrageneric groups of taxa, and secondly using 31 characters to quantify the morphological variation found within sect. Nuphar. Results from combined phenetic analyses provided empirical support for recognition of two infrageneric groups of species in the genus (largely by fruit, stigmatic disk margin, neck, and stigmatic disk size) and for separation of taxa within sect. Nuphar (by leaf size, and fruit disk and neck size). Statistical analyses demonstrate that extensive variation in ranges of character states exists among sect. Nuphar taxa. However, a suite of qualitative and quantitative characters can be used to recognize five species and three subspecies within this predominantly Old World assemblage.     

Biometrical variability of foliage and cone characters inCupressus bakeri (Cupressaceae)

Biometrical variability of foliage, cones, and seeds were analyzed in 8 native populations (throughout the range) of a relictual species,Cupressus bakeri Jeps. (Cupressaceae) from California and Oregon. The partition of variation within and among populations for this rare plant was investigated and the justification for subspecies status was reviewed. Morphological characters varied significantly, with more of the variation among populations than at the population level. Multivariate analyses indicated close affinities among populations on serpentine sites suggesting the presence of a serpentine race. However, the high degree of variation among populations did not permit their clear separation into the two subspecies described byWolf (1948). We conclude thatC. bakeri should be treated as a monotypic species.     

Infra-specific genetic and morphological diversity in Linum album (Linaceae)

The genus Linum L. (Lineacea) has over 15 species, subspecies or ecotypes in Iran. These species show extensive geographical distribution and form many local populations throughout the country. Linum album is herbaceous medicinal plant containing important lignans such as podophyllotoxin (PTOX) and 6-methoxy podophyllotoxin (MPTOX), which have antiviral and anticancer properties. Studying the genetic and morphological diversity of different geographical populations produces detailed knowledge about population divergence and identification of the infra-species taxa if at all they are present. Moreover, the populations that differ in their genetic content and structure may also differ in their chemical and medicinal properties. The present study considers morphological and genetic diversity analyses of 20 L. album geographical populations by using nuclear ISSR markers, genome size, and cytogenetic characteristics. These populations differed significantly in many of their quantitative morphological characters and in some of their qualitative features. They also differed significantly in their molecular characteristics and genome size. Details of morphological and molecular variations are reported and discussed.     

小叶锦鸡儿天然居群叶形态性状变异研究

为揭示小叶锦鸡儿(Caragana microphylla)天然居群叶形态性状的变异规律及其生态适应性特征,该研究以10个小叶锦鸡儿天然居群为对象,通过多重比较、巢式方差分析、相关性分析、聚类分析和主成分分析等方法,对7个叶形态性状进行分析。结果表明：(1)小叶锦鸡儿叶形态性状在居群内和居群间均存在极显著差异(P < 0.01),平均变异系数为10.13%,不同性状的变异幅度为6.23%~12.78%;平均叶形态性状的表型分化系数为43.62%,居群内变异(30.09%)大于居群间变异(24.91%),说明居群内是其叶形态性状变异的主要来源。(2)相关性分析表明,环境因子对小叶锦鸡儿的叶形态性状变异有很大的影响,在地理空间上主要呈现出沿海拔梯度的变异模式;主成分分析的结果显示,小叶宽、叶柄宽和叶柄长对小叶锦鸡儿叶形态变异起主导作用;利用欧式距离对小叶锦鸡儿居群进行UPGMA聚类分析结果显示,基于叶形态性状和环境因子可分别将小叶锦鸡儿10个居群分为3类和2类,Mantel检验结果表明,小叶锦鸡儿的叶形态性状变异不存在地理连续性。研究结果为小叶锦鸡儿的适应性进化和开发利用提供了理论依据。     

The morphological variation and geographical distribution ofBiscutella glacialis (Boiss. etReuter)Jordan in Spain

A study was made of the morphological variation in seven population samples ofB. glacialis. Data on quantitative and qualitative characters reveal that one of the populations differs considerably from the rest. Observations on geographical distribution show that this species is not endemic to the peaks of Sierra Nevada but far more widely distributed than previously realized. Two varieties ofB. glacialis, one of which is new, are described.     

紫距淫羊藿的地理分布新记录及形态描述

淫羊藿属依据少量栽培个体命名的类群普遍存在,往往缺乏足够的野外调查和形态性状统计,对其性状变异幅度、变异式样和分类价值研究不够。紫距淫羊藿(Epimedium epsteinii)由Stearn于1997年依据引种个体命名,为掌握其资源分布及形态性状变异,该研究基于居群概念,对淫羊藿属分布区展开调查,在各分布区选择一个居群(30个体/居群)于花期进行形态观察、统计和描述。结果表明:紫距淫羊藿以往被认为狭域分布于湖南天平山,该研究调查发现广东省和湖北省均有分布,为新分布记录。此两处新分布分别位于天平山的南部和北部,将其自然分布区显著扩大。基于居群观测的形态变异较原描述更广泛。6个主要数量性状(株高、序轴长、小花数、叶长、叶宽和根茎)和花部颜色在居群间和个体间均呈现丰富的变异。生物种的分布范围并非完全连续,而是由若干个多少间断的居群组成,任何一个居群都存在不同程度的连续性变异,要更好地认识各物种,就应当足够重视这些居群系统。紫距淫羊藿的新分布记录对淫羊藿属的资源利用、扩散机制和系统地理学研究具有重要意义。     

Quantitative morphological variation inTetramolopium (Asteraceae) in Hawaii and the Cook Islands

Multivariate analyses of quantitative characters were undertaken to examine the pattern of variation resulting from adaptive radiation ofTetramolopium in Hawaii and to investigate the systematics of an undescribed species in the Cook Islands and a group of populations of uncertain affinity on Kalaupapa Peninsula in Hawaii. Taxa differentiated on quantitative characters in the absence of qualitative characters. Differentiation was observed in multiple characters which could be organized into a few principal components. The analyses supported the species rank of the Cook Islands plants. The relationship of the Kalaupapa Peninsula populations to the rest of the taxa needs further study.     

Morphology and taxonomy of the Puccinia species on corn and sorghum

Morphological characters of the sori and spore forms ofPuccinia polysora andP. purpurea are described and illustrated indicating the range of variation as determined from herbarium and field materials. Morphological variation inP. sorghi clones and inbred lines from these clones were studied earlier. The genes conditioning morphology apparently function independently of those conditioning pathogenicity. Race 13 ofP. purpurea was morphologically similar toP. sorghi and probably was not derived from the recombination of gametes ofP. purpurea as indicated in an earlier paper. Morphological distinctions between the 3 pathogens viz.P. sorghi, P. polysora andP. purpurea are wide. Taxonomically they are retained as valid species. A study of the complete life cycle of the latter 2 species and the genetic control of morphological characters, however, is essential to the final solution of the problem.     

Phylogenetic relationships of three genera in Anacardiaceae: Bonetiella, Pseudosmodingium, and Smodingium

Phylogenetic analyses were conducted to determine the relationships between and characterize the morphological variation among three genera of Anacardiaceae:Bonetiella, Pseudosmodingium, andSmodingium. The first two genera are endemic to Mexico and the third to South Africa.Bonetiella andSmodingium are monotypic.Pseudosmodingium has eight species, six extant and two fossil, and all were included in the analyses. Fifty-six anatomical characters for leaves, wood, and fruit, as well as morphological characters, were considered. Our results indicate thatSmodingium is not closely related toPseudosmodingium andBonetiella, and that these last two genera are sister taxa.Pseudosmodingium is a monophyletic group and the two fossil species are related to species from the same geographical area of central Mexico where they were found. Fruit anatomical characters were significant at the generic level, while leaf anatomical characters were important at the species level inPseudosmodingium.

---

Resumen  En este trabajo se llevaron a cabo análisis filogenéticos para determinar las relaciones filogenéticas así como caracterizar la variación morfológica de tres géneros de Anacardiaceae:Bonetiella, Pseudosmodingium ySmodingium. Los dos primeros son endémicos a México y el terceroo a Sudáfrica.Bonetiella ySmodingium son monotípicos.Pseudosmodingium incluye ocho especies, dos fósiles y el resto vivientes, y todas fueron consideradas en los análisis. Se consideraron 56 caracteres tanto anatómicos como morfológicos de hojas, madera y frutos. Nuestros resultados indican queSmodingium no está cercanamente relacionado aPseudosmodingium yBonetiella, y que estos dos últimos géneros son taxa hermanos.Pseudosmodingium es un grupo monofilético y las dos especies fósiles están relacionadas a especies que crecen en las mismas áreas geográficas del centro de México, donde fueron descubiertas. Los caracteres anatómicos del fruto fueron significativos a nivel genérico, mientras que los de hoja fueron importantes a nivel de especies enPseudosmodingium.

     

Species delimitation in the Luziola peruviana (Poaceae) complex

Luziola is a small genus from aquatic environments of the New World. The most widespread species in the genus isL. peruviana. Morphological variation has been documented for this species throughout its entire range of distribution, however this variation has been difficult to characterize. A population aggregation analysis was performed in order to determine how many species can be identified when analyzing characters from the individuals of this taxon as well as with individuals from closely related taxa such asL. divergens, L. doelliana, andL. pittieri. Multivariate analysis, ANOVA and ANCOVA analyses were also performed to detect if quantitative morphological variation is related to altitude and latitude. Results indicate that within this complex only one species should be recognized, referable toL. peruviana, and that plants near the equator are larger, and have larger lemmas and paleas in both female and male flowers. An identification key for the nine species now recognized inLuziola is included.     

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.     

Patterns of morphometric variation and a new supraspecific classification of apomictic taxa of Hieracium (Asteraceae) from Denmark and southern Sweden

Patterns of variation among all micro-species of Hieracium sect. Hieracium and H. sect. Vulgata known from Denmark and southern Sweden are investigated. Forty-three, possibly independent morphological characters are used. Significant correlations were revealed among the majority of the characters. Multivariate analyses did not reveal any discontinuities in the variation or any discrete groups of species. However, there were obvious correlations between previous taxonomic classifications and the result of the multivariate analyses. Based on the statistical analyses, three sections and 38 informal species aggregates are recognized. H. sect. Bifidum T. Tyler is described anew. The characters that turned out to be most influential in the multivariate analyses, and thus most important for the classification, were the distribution of stellate tomentum on the phyllaries, the amount of glandular hairs and the amount and pigmentation of simple hairs on the phyllaries, the shape of the leaf-base and the regularity of the leaf dentation. The evolution of the group and the taxonomic treatment is thoroughly discussed in the light of the results.     

贵州省野生皂荚种实表型性状多样性分析

为深入了解贵州省野生皂荚(Gleditsia sinensis)荚果表型性状的遗传多样性及其变异类型,为皂荚的遗传改良、种质鉴定、亲本选择以及品种培育奠定理论基础。该研究以贵州省7个野生皂荚群体70个个体为研究对象,采用方差分析、主成分分析、相关性分析及多性状综合评价等方法对皂荚群体的10个种实表型性状进行系统分析和综合评价。结果显示：(1)所测皂荚的表型性状差异在群体内均达到极显著水平(P＜0.01);在群体间,除每荚粒数、种子宽、种子长宽积以及种子长宽比之外,其余表型性状的差异均达极显著水平(P＜0.01)。(2)7个居群野生皂荚各性状平均变异系数为21.16%,其中凯里市(P4)居群的变异系数最高(24.44%);居群间荚果的变异(29.22%)高于种子的变异(11.04%),且变异主要来自于群体内。(3)相关分析显示,皂荚种实各性状之间存在不同程度的关联性;主成分分析显示,前4个主成分(皂荚种子大小、单个荚果出籽数量、种子形态指数因子、与荚果长和种子厚相关的因子)的累积贡献率达69.783%,可基本反映皂荚表型性状的大部分信息;以10个种实性状对皂荚野生群体进行综合评价发现,来自于惠水县(P7)群体的皂荚种实性状综合评价最高。研究表明,贵州省野生皂荚在群体间及群体内具有丰富的表型变异,且群体内的变异大于群体间的变异,变异主要来自于群体内。     

Estimates of hierarchical variation in flower morphology in natural populations ofScleranthus annuus (Caryophyllaceae), an inbreeding annual

The variation in two sets of morphological characters of the flowers of the highly inbreedingScleranthus annuus (Caryophyllaceae) was assessed using 15–20 plants from each of 20 natural populations from the southernmost region of Sweden. The stamen fertility data set consisted of 10 characters describing the degree of fertility of the ten stamens/staminoids, while the sepal/gynoecium data set comprised 10 sepal characters, together with style and stigma length. Substantial variation was found in both the degree of development and the fertility of the stamens, the degree of variability in stamen fertility being related to stamen position within the flower. Considerable variation was found in the characters of the sepal and the gynoecium. Hierarchical analyses of variance indicated that 29% of the variation in total male reproductive effort was distributed among populations, 28% among plants within populations and 43% represented within-plant variation. The corresponding averages for the characters from the sepal/gynoecium data set are 26, 38, and 35%: a greater proportion of the total variance in female reproductive characters is accounted for by among individual variation than is the case with the male reproductive characters. Significance tests of Mahalanobis distances derived by canonical variate analyses indicated that all populations were significantly separated using the sepal/gynoecium data set, while only 50% of the pairwise comparisons on the basis of the stamen fertility data set were significant. Cluster analysis did not reveal any aggregation of the populations. The incongruence of the two data sets and their ability to discriminate between the populations is discussed.