Intraspecific evolution ofMicroseris pygmaea (Asteraceae,Lactuceae) analyzed by cosegregation of phenotypic characters (QTLs) and molecular markers (RAPDs)

The Chilean annual,Microseris pygmaea, has differentiated in distinct coastal and inland series of populations after long-distance dispersal from western North America. Two plants from the most diverse biotypes were crossed, a large F₂ was raised and analysed for segregation of 30 phenotypic characters. Segregation of molecular markers (47 RAPDs, 1 RFLP, 2 isozymes) was determined in a subpopulation of 45 plants which include all extremes for the phenotypic characters. 32 marker/character cosegregations were significant at the 1% level in t-tests between dominant and homozygous recessive marker genotypes. Considering linkage among markers and pleiotropy of certain marker loci, the number of independent quantitative trait loci (QTLs) is reduced to about 18. Interactions among 2 or 3 QTLs affecting one character have been characterized. The phenotypic differentiation ofM. pygmaea during its evolution from a single founder individual begins to be understood at the level of single-gene mutants.     

Differentiation in phenotypic plasticity among populations of Arabis serrata Fr. & Sav. (Brassicaceae)

Studies on divergence of phenotypic plasticity in closely related species have suggested that character means and plasticity of these characters may evolve independently. Similar patterns of divergence between populations within a species have been reported although few plant species have been studied. Thus, in this paper, the patterns of differentiation between character means and phenotypic plasticity among eight populations of Arabis serrata are documented. Mean response and magnitude and pattern of phenotypic plasticity were measured and compared in plants growing under an environmental gradient of nutrients. Differences in means and coefficients of variation (CV as indicators of plasticity) among populations were compared using the Canberra metric and generating unrooted Wagner trees. Populations showed significant differences in character means in nine morphological traits. Magnitude and patterns of phenotypic plasticity showed a complex pattern of differentiation for each trait and population. Biomass traits were more plastic, in general, than characters associated with linear size. Comparisons between pairs of populations for nine morphological traits showed that in 28.6% of 252 possible cases, populations differed in means, magnitude and patterns of phenotypic plasticity. In almost 90% of the cases, populations differed in magnitude and/or pattern of plasticity. Considering all characters together, populations from similar habitats and with common life history features tended to respond in similar ways. The patterns of divergence, however, suggest that character means and character plasticities among populations are able to evolve independently.     

Morphological, cytogenetic and allozymic variation within Cephalofovea (Onychophora: Peripatopsidae) with descriptions of three new species

Variation within and between populations of Cephalofovea (Peripatopsidae) has been examined by allozyme, karyological and morphological analyses. Four groups are recognized on the basis of allozyme electrophoresis. One group includes specimens from the type locality of the only described species of the genus, C. tomahmontis. While karyotypic and morphological character states show considerable inter-group variation, the distributions of these states among groups are not concordant when different characters are compared. However, each group of populations is uniquely defined by the full suite of character states it possesses. The four groups are recognized here as distinct species with three species described as new.     

DISSECTING CORRELATED CHARACTERS: ADAPTIVE ASPECTS OF PHENOTYPIC COVARIATION IN MELANIZATION PATTERN OF PIERIS BUTTERFLIES

In this study we address the question of how much of the covariation among phenotypic characters observed in natural populations is adaptive. We examine covariation among a set of phenotypic characters that describe the wing-melanization pattern of Pieris butterflies. Previous functional analyses of thermoregulatory performance allow us to predict a priori whether and how different wing melanic characters should be correlated. We quantify and analyze the variation in the wing-melanization pattern within species for a series of Pieris populations from relatively cool environments in North America and compare these results with the predictions based on our adaptive hypothesis. We consider adaptive covariation both for biogeographic variation among populations and for seasonal polyphenism (phenotypic plasticity) within populations. Our hypothesis correctly predicts many of the qualitative features of covariation in melanization among major regions of the wings, at the level of biogeographic variation among populations, for both males and females of Pieris occidentalis. When within-population variation is considered, agreement with the adaptive predictions varies considerably in different populations for both P. occidentalis and P. napi males and females. Agreement for P. napi, particularly the females, is generally poorer than for P. occidentalis. In both species, there is a consistent difference in melanization pattern between alpine and arctic sites; this difference is discussed in relation to the differences in the radiative environment between these two types of “cold” habitats. Our results suggest that some important aspects of phenotypic correlation among wing melanic characters in Pieris are adaptive. We emphasize the important distinction between covariation and co-occurrence of characters, and we discuss these results in relation to the extensive biogeographic variation and phenotypic plasticity (seasonal polyphenism) in Pieris wing-melanization patterns.     

Reaction norms of Arabidopsis. V. Flowering time controls phenotypic architecture in response to nutrient stress

The evolutionary and environmental stability of character correlations has increasingly been the focus of ecological and quantitative genetic studies. Although the genetic stability of character correlations is a central assumption of quantitative genetic models of phenotypic evolution, theoretical considerations suggest that both the genetic and the phenotypic architecture should change in response to selection and to environmental heterogeneity. We investigate genetic (population) differences and plasticity to nutrient availability of the phenotypic architecture describing the whole-plant phenotype of Arabidopsis thaliana (Brassicaceae). We found significant genetic differences among early and late flowering ecotypes in the relationships between several traits, when a path-analytical model was used to estimate character correlations. Furthermore, we found significant plasticity of several path coefficients when nutrient levels were altered. A whole-plant analysis considering all paths in the model simultaneously confirmed that populations of A. thaliana are characterized by distinct phenotypic architectures, and that these are altered in different ways by environmental changes. We discuss the implications of these findings for our understanding of selective pressure on and response by multivariate phenotypes.     

Geometric and traditional morphometrics for the assessment of character state identity: multivariate statistical analyses of character variation in the genus Arrenurus (Acari,Hydrachnidia, Arrenuridae)

Geometric and traditional morphometric approaches are tested to describe and reveal taxonomic characters and character states in variation of shape and size of idiosoma morphology in the Arrenuridae. Patterns of variation of idiosoma and glandularia features of males from 11 Mexican species of Arrenurus (Megaluracarus) and two species of subgenus Dadayella were explored with five landmark configurations and three sets of interlandmark distances. Separate principal component analyses (PCA) and canonical variate analyses (CVA) were performed for each data set. The eight multivariate analyses of variance among 13 a priori groups (species) detected significantly different morphometric variants, which were interpreted as different taxonomic character states. Patterns of character state similarity among species were examined with unweighted‐pair grouping method using averages (UPGMA) cluster analyses on Mahalanobis distances. Analyses of five landmark configurations revealed important taxonomical variation in the anterior idiosoma outline (10 character states), the outline of the posterior region or cauda (13 states), the distribution of postocularia, and the second and third pairs of dorsoglandularia (nine states), the fourth pair of dorsoglandularia (three states), and ventroglandularia on the posterior side of idiosoma (nine character states). Multivariate analyses of three sets of distance measurements also resulted in the detection of potential taxonomic characters related to idiosoma size (12 character states), postocularia and dorsoglandularia (13 states), and ventroglandularia (nine character states). Morphometric analyses of distances and shapes provide a formal basis for the interpretation of taxonomic characters, and for the discovery of character states. These characters should be investigated further in a wider sample of species for the phylogenetic systematics of these water mites. In the meantime, idiosoma regions and structures were tested for congruence in a phylogenetic analysis, and were proposed as homologous among the species sampled.     

MORPHOMETRIC VARIABILITY IN CONTINENTAL AND ATLANTIC ISLAND POPULATIONS OF CHAFFINCHES (FRINGILLA COELEBS)

The magnitudes of phenotypic variances in peripherally isolated populations of common chaffinches (Fringilla coelebs) in the Azores, Madeira, and Canaries relative to their continental source populations in Iberia and Morocco have played a pivotal role in the formulation of competing hypotheses of phenotypic evolution in these isolates. Because Van Valen's niche variation hypothesis and Grant's model of island evolution were based on museum skins prone to measurement error and temporal variation, we re-examined the patterns of phenotypic variability using more precise skeletal measurements from freshly collected specimens. Levene's tests showed that univariate character variances were homogeneous in all island and continental populations, although there was a consistent trend for the magnitude of the variances to be lower for all characters in all Canary island populations. Multivariate Levene's tests, however, revealed significantly reduced total variances in the Hierro and Madeira populations compared to some Azores and continental populations. The Azores and continental populations did not differ in variability, and lower variances in the Canaries were not related to the presence or absence of the congeneric blue chaffinch (F. teydea), contrary to the predictions of Van Valen's niche variation hypothesis. Population variability was not inversely related to differentiation or isolation within the Azores or Canaries archipelagoes, opposite to the association reported by Grant. Our results also differ from both previous studies which reported much larger differences in population variabilities, and this likely reflects the use of heterogeneous samples of museum specimens, less precise external characters, and the use of tests sensitive to sample size. Differentiation among populations has been markedly greater in the Canary islands, implicating founder events and possibly historically stronger directional selection as determinants of this enhanced divergence relative to the Azores. These variance-reducing processes are unlikely to explain current lower levels of phenotypic variability because there has been sufficient time since colonization for replenishment of variability in polygenic characters. Average heterozygosities at putatively neutral allozyme loci are 1.6 times higher in the Azores compared to the Canaries, and support the view that effective population sizes are smaller in the latter archipelago. We argue that reduced variances in the Canary island populations represent lower equilibrium levels maintained by drift and mutation in populations with smaller long-term effective sizes, consistent with Lynch and Hill's neutral model of phenotypic evolution. Although episodes of selection in the past may have been interspersed with long periods of effective neutralism and drift, adaptationist hypotheses invoking a primary role for variance-reducing selection appear to be unwarranted.     

Modular genetic architecture of floral morphology in Nicotiana: quantitative genetic and comparative phenotypic approaches to floral integration

Animal‐pollinated flowers are complex structures that may require a precise configuration of floral organs for proper function. As such, they represent an excellent system with which we can examine the role of phenotypic integration and modularity in morphological evolution. We use complementary quantitative genetic and comparative phenotypic approaches to examine correlations among floral characters in Nicotiana alata, N. forgetiana and their artificial fourth‐generation hybrids. Flowers of both species share basic patterns of genetic and phenotypic correlations characterized by at least two integrated character suites that are relatively independent of each other and are not disrupted by four generations of recombination in hybrids. We conclude that these integrated character suites represent phenotypic modules that are the product of a modular genetic architecture. Intrafloral modularity may have been critical for rapid specialization of these species to different pollinators.     

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。     

TESTS OF INBREEDING EFFECTS ON HOST-SHIFT POTENTIAL IN THE PHYTOPHAGOUS BEETLE OPHRAELLA COMMUNA

Although inbreeding, on average, decreases additive genetic variance, some inbred populations may show an increase in phenotypic variance for some characters. In those populations with increased phenotypic variance, character changes by peak shifts may occur because of the effects of the higher variance on the adaptive landscape. A population's increased phenotypic variance may place it in the domain of attraction of a new adaptive peak or increase the likelihood of a selection-driven peak shift as the landscape of mean fitness flattens. The focus of this study was to test for increased variance, in inbred populations, in a behavioral character involved in adaptive diversification and probably speciation. We examined the effect of inbreeding on feeding responses of the leaf beetle Ophraella communa in a series of inbred lineages across a range of levels of inbreeding (f = 0.25, 0.375, 0.5). We measured the feeding response of inbred lineages of O. communa on its normal host, Ambrosia artemisiifolia, and on two novel plants, Chrysopsis villosa and Iva frutescens, that are the hosts of other Ophraella species. The results show that feeding responses on the different plants are not correlated, indicating that the feeding responses to the different plants are to some degree genetically independent. Despite apparent genetic variation in lineage feeding responses, we could not statistically demonstrate increases in phenotypic variance within the lineages. Thus, the experimental results do not support the idea that host shifts in this beetle evolved by peak shifts in bottlenecked populations.     

Phenotypic plasticity inCrepis tectorum (Asteraceae)

Two experiments were carried out using two different approaches to compare populations ofCrepis tectorum (Asteraceae). One was based on a comparison of means of various vegetative and reproductive characters and another was based on a comparison of response patterns of the same characters in a series of environments. Population divergence within two earlier recognized form series, one from weed habitats and one from alvar habitats on Baltic islands, resulted in a partially overlapping pattern in cluster analyses based on character means. However, the pattern revealed by a comparison of the direction and amount of plastic response suggested that populations within the two form series had more similar response patterns than other combinations of populations. It was concluded that patterns of plasticity may provide useful additional information on the overall similarity among taxa. An hypothesis that plants in weed populations should exhibit a greater phenotypic response to the environments than plants in alvar populations was rejected.     

Spontaneous variability in a population of Candida albicans strains]

The spontaneous variability of the populations of C. albicans strains of different genesis in the morphological properties of their colonies and in the potential of the activity of their extracellular proteolytic and phospholipid enzymes has been studied. The isolated types of colonies, differing in their morphology, have the phenotypic character of variability. Different populations of strains exhibited variability in the activity of enzymes, depending on morphological variants isolated from these populations. Selected morphological variants with high potential of their proteolytic enzymes retained stability in this property for 5 generations and can be used in medical practice for the isolation of C. albicans antigens.     

油松天然群体的种实性状表型多样性研究

为了揭示油松天然种群在不同地理环境条件下表型变异的程度和规律,在油松整个天然分布范围内选择了12个具有代表性的居群作为研究对象,对其球果、种子、种翅等12个种实性状的变异程度及其与环境因子间关系进行了比较分析。结果显示:(1)各个性状在居群内和居群间均存在较大的变异(CV>12%)。其中千山(QS),曾家镇(ZJ)和互助(HZ)3个居群表现出了较高的变异(CV>20%),而球果干重(CDW)和种子长(CL)是所有表型性状中变异幅度最大的(CV分别为31%和21%),但种翅性状与其他性状相比具有较高的稳定性。(2)巢式设计方差分析表明,在居群内表型分化系数(Vst)变化在3.18%~89.86%之间,而群体间的Vst为38.97%;与其他针叶树种相比,油松拥有较高的表型分化系数,且居群内的变异程度远高于居群间的变异,尤其是千山(QS)、曾家镇(ZJ)和互助(HZ)3个居群,这说明油松具有较高的环境异质性适应能力或恶劣环境耐受能力。(3)相关性分析表明,该研究的各形态特征与潜在蒸发量均为负相关,且大部分形态指标间及它们与潜在蒸发量间存在显著相关性,表明潜在蒸发量是油松形态特征变化的最重要环境影响因子,预示油松最适宜生长于温暖潮湿的环境中;并表明因各形态特征间相互紧密关联,所以它们受环境条件影响而共变。     

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.     

浙江省野生蜡梅花部形态变异及其与环境因子的相关性

以浙江省蜡梅自然分布区的5个居群为研究对象,采用巢式方差分析、主成分分析、相关分析、聚类分析等多种分析方法,探究蜡梅花部的形态变异及其与环境因子相关性以及居群间和居群内的表型多样性。结果表明:蜡梅花部性状中除雄蕊长度、雌蕊数、雌蕊长度、内被片数以外,其他12个性状在居群间均有极显著差异。胚珠数的平均变异系数最大,花冠内径的平均变异系数最小,各性状的平均变异系数为15.03%(10.81%—23.38%)。五尖山的表型多样性最丰富,碧东山居群最小,种群间平均表型分化系数为44.38%(1.57%—89.62%),种群内变异大于种群间变异,种群内变异是花部变异的主要来源。主成分分析显示花冠直径、花冠内径、花筒深度和中被片长对种群变异起主要贡献作用。花部多数性状间存在显著或极显著的相关关系,生态因子中花部性状与土壤酸度相关性最大,土壤中花部性状与大量元素相关性最大,微量元素次之,中量元素最小。通过UPGMA聚类分析可以将5个种群分成两组。浙江省野生蜡梅花部形态存在丰富的变异和多样性,花部部分性状与土壤中钙、铁、铜含量和海拔有显著或极显著相关关系。     

SELECTION FOR NATIVE CHARACTERS IN HYBRIDS BETWEEN TWO LOCALLY ADAPTED PLANT SUBSPECIES

Gene exchange between locally adapted plant populations can have significant evolutionary consequences, including changes in genetic diversity, introduction of adaptive or maladaptive traits, disruptive of coadaptive gene complexes, and the creation of new ecotypes or even species. The potential for introgression between divergent populations will depend on the strength of selection against nonnative characters. Morphologically variable F₂ hybrids of two Gilia capitata subspecies were used to evaluate the strength of phenotypic selection and the response to selection in the home habitats of each subspecies. At both sites, traits diagnostic of the subspecies were subject to significant phenotypic selection, probably mediated by direct selection on unmeasured correlated characters. Phenotypic selection favored native morphologies in all but a single case; leaf shape of one subspecies was favored in both habitats. The strength of selection varied between sites, with one site selecting more strongly against nonnative characters. Offspring of the F₂ hybrids showed a significant evolutionary response to selection when grown in a common environment. Evolution was in the direction of similarity with the subspecies native to the site where selection was imposed. This result reveals that native character states are adaptive and suggests that selection will maintain native morphologies even after a substantial influx of genes from an ecologically and morphologically distinct, and locally adapted subspecies.     

疏花水柏枝结构性状多样性研究

以疏花水柏枝(Myricaria laxiflora)茎的11个解剖结构参数作为表型性状,对覆盖整个分布区域的居群进行部分抽样取样,研究了该物种表型性状的多样性及其格局、表型性状的分化及其与环境的关系。结果表明,各性状在居群间以及不同居群总体性状之间都存在显著或极显著的差异,63％的表型分化属居群内分化,37％为居群间地理差异引起的分化,居群内性状的变异程度与立地环境条件的分化程度一致,奉节以上居群相似程度较大,而以下居群相似程度则较小,并有独立分化的居群出现,这表明表型性状的多样性中心是在三峡河谷区域内,而不是在三峡以上的河谷区段。     

The extent of variation in male song, wing and genital characters among allopatric Drosophila montana populations

Drosophila montana, a species of the Drosophila virilis group, has distributed around the northern hemisphere. Phylogeographic analyses of two North American and one Eurasian population of this species offer a good background for the studies on the extent of variation in phenotypic traits between populations as well as for tracing the selection pressures likely to play a role in character divergence. In the present paper, we studied variation in the male courtship song, wing and genital characters among flies from Colorado (USA), Vancouver (Canada) and Oulanka (Finland) populations. The phenotypic divergence among populations did not coincide with the extent of their genetic divergence, suggesting that the characters are not evolving neutrally. Divergence in phenotypic traits was especially high between the Colorado and Vancouver populations, which are closer to each other in terms of their mtDNA genotypes than they are to the Oulanka population. The males of the Colorado population showed high divergence especially in song traits and the males of the Vancouver population in wing characters. Among the male song traits, two characters known to be under sexual selection and a trait important in species recognition differed clearly between populations, implying a history of directional and/or diversifying rather than balancing selection. The population divergence in wing characters is likely to have been enhanced by natural selection associated with environmental factors, whereas the male genitalia traits may have been influenced by sexual selection and/or sexual conflict.     

Independent contrasts succeed where ancestor reconstruction fails in a known bacteriophage phylogeny

Abstract.— Methods of ancestor reconstruction are important tools for evolutionary inference that are difficult to test empirically because ancestral states are rarely known with certainty. We evaluated reconstruction methods for continuous phenotypic characters using taxa from an experimentally generated bacteriophage phylogeny. Except for one slowly evolving character, the estimated ancestral states of continuous phenotypic characters were highly inaccurate and biased, even when including a known ancestor at the root. This error was caused by a directional trend in character evolution and by rapid rates of character evolution. Computer simulations confirmed that such factors affect reconstruction of continuous characters in general. We also used phenotypic viral characters to evaluate two methods that attempt to estimate the correlation between characters during evolution. Whereas a nonphylogenetic regression was relatively inaccurate and biased, independent contrasts accurately estimated the correlation between characters with little bias.