Morphologic variation in northern marginal Juniperus oxycedrus L. subsp. oxycedrus populations in Istria

Marginal tree populations are believed to be more differentiated and host less variation than central tree populations. The aim of this study was to perform a detailed morphometric study of J. oxycedrus L. subsp. oxycedrus in northern marginal populations in order to establish its phenotypic variation and geographical differentiation and to examine the eventual presence of putative species J. deltoides in Istria. Morphologic variation was studied in 16 northern marginal populations in Istria. Nine morphological traits from a minimum of 50 leaves from each of 206 individuals and two morphological traits from 30 to 50 seed cones in each of 103 females were measured. Phenotypic variation in Istrian populations was high and of a similar magnitude to that found in the centre of the distribution area. Gender dimorphism in leaf morphology was detected, but the study failed to confirm its uniform distribution pattern. Significant and surprisingly high among-population differentiation (8–30%) was revealed. Average leaf shape in the study area is clearly “delta-shaped” characterised by a relatively wide leaf base. However, no significant differences in any of the studied traits were found between the marginal Istrian populations and the more central populations analysed in our earlier studies.     

Leaflet morphometric variation of service tree (Sorbus domestica L.) in the Balkan Peninsula

Abstract

In most European countries, the service tree (Sorbus domestica L.) is a rare and threatened species and its conservation has been recognised as a priority. The aim of this study was to asses its morphologic variation in the western and central part of the Balkan Peninsula and in southern Central Europe. Three populations were analysed: one in Serbia, one in Bosnia and Herzegovina and one in Slovenia. In each population 30 trees were selected, and from each tree 30 leaves were collected for morphometric analysis based on nine leaflet morphological traits. Univariate (ANOVA) and multivariate (MANOVA) analysis of variance were used to estimate the variation within- and between populations and a discriminant analysis was performed to examine the structure of the between-population differences. The values of particular morphological traits found in our study did not differ considerably from the values reported elsewhere. The results revealed significant within- and between population variation. Variation within populations was highly significant for all the scored leaf morphological traits, while variation between populations was significant for all the studied traits except for the leaflet length. The discrimination between the three populations was significant. High percentages of correctly classified samples demonstrate good discriminating employability of the analysed leaf morphological traits and indicate differentiation of the analysed populations.     

Genetic and morphological differentiation in Populus nigra L.: isolation by colonization or isolation by adaptation?

Identifying processes underlying the genetic and morphological differences among populations is a central question of evolutionary biology. Forest trees typically contain high levels of neutral genetic variation, and genetic differences are often correlated with geographic distance between populations [isolation by distance (IBD)] or are due to historic vicariance events [isolation by colonization (IBC)]. In contrast, morphological differences are largely due to local adaptation. Here, we examined genetic (microsatellite) and morphological (from a common garden experiment) variation in Populus nigra L., European black poplar, collected from 13 sites across western Europe and grown in a common garden in Belgium. Significant genetic differentiation was observed, with populations from France displaying greater admixture than the distinct Spanish and central European gene pools, consistent with previously described glacial refugia (IBC). Many quantitative traits displayed a bimodal distribution, approximately corresponding to small‐leaf and large‐leaf ecotypes. Examination of nine climatic variables revealed the sampling locations to have diverse climates, and although the correlation between morphological and climatic differences was significant, the pattern was not consistent with strict local adaptation. Partial Mantel tests based on multivariate summary statistics identified significant residual correlation in comparisons of small‐leaf to large‐leaf ecotypes, and within the small‐leaf samples, but not within large‐leaf ecotypes, indicating that variation within the small‐leaf morphotype in particular may be adaptive. Some small‐leaf populations experience climates very similar to those in large‐leaf sites. We conclude that adaptive differentiation and persistent IBC acted in combination to produce the genetic and morphological patterns observed in P. nigra.     

Variation in morphological traits and trait asymmetry in field Drosophila serrata from marginal populations

Drosophila serrata occurs along the eastern coast of Australia with a southern range boundary near Sydney. To compare levels of phenotypic variation in marginal and central populations, we examined morphological variation in populations of this species from the southern range boundary and two more northerly populations. The populations differed for wing traits and there was an increase in wing size in the marginal locations which persisted under laboratory culture. The means of wing and bristle traits increased under laboratory culture, whereas wing trait coefficients of variation and variances decreased. Heritability estimates for wing size traits tended to be lower in the field compared with the laboratory, whereas bristle and crossvein length heritabilities were similar across environments. There was evidence for heritable variation in wing and bristle traits in both the marginal and more northern populations, suggesting that genetic variation was not limiting in marginal populations. Fluctuating asymmetry (FA) was also assessed as a measure of genomic and environmental stress. There were no consistent differences among populations for the FA of individual traits, or for a total FA score summed across traits. FA levels in field parents and laboratory‐reared progeny were similar. Overall, the results do not support the conjecture that levels of phenotypic and genetic variability differ between central and marginal D. serrata populations.     

Effects of copper on the fitness of the Chilean scallop Argopecten purpuratus (Mollusca: Bivalvia)

Exposure to high concentrations of metals can affect populations and individuals at morphological, physiological, biochemical or genetic levels. Metal pollution is a source of environmental stress that can have deleterious effects on organisms and generate selective pressure upon populations. This work attempts to establish whether concentrations of copper, above physiological requirements, can affect the genetic structure of a cohort of Chilean scallop Argopecten purpuratus. Following the determination of LC₅₀ for 96 h experiments, other acute toxicity tests were carried out, exposing the juveniles to solutions of 150 ppb copper for 120 h. Dead, surviving and control individuals from these bioassays were genetically characterized for five polymorphic loci, Isocitrate dehydrogenase (Idh), Octopine dehydrogenase (Ocdh), Phosphogluconate dehydrogenase (Pgd), Leucine aminopeptidase (Lap) and Phosphoglucose isomerase (Pgi). Results showed no significant differences in allele and genotypic frequencies between surviving and dead individuals, although for the same groups significant differences were found in estimated mean heterozygosity. Pgi and Ocdh showed significantly different heterozygosity values for dead and surviving individuals. A positive relationship between multilocus heterozygosity and survival was found for young A. purpuratus exposed to high copper concentrations; thus, we found evidence of a differential response to exposure to high copper concentrations, related to degree of heterozygosity.     

不同种源樟树叶片形态特征及生长差异分析

为了解不同种源樟树叶片形态特征和生长差异,该文以30个种源樟树为研究对象,对其叶长、叶宽、叶柄长、周长、叶面积、长宽比、形态因子、株高、地径等指标进行测定和差异性分析.结果表明:(1)30个种源间叶片性状的变异系数为3.88％～16.14％,显示不同种源樟树叶片形态特征存在显著差异;叶长、叶宽、叶柄长、周长、面积、叶厚...     

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。     

Variation and selection in western montane species II. Variation within and between populations of white firon an elevational transect

Summary Four populations of Abies concolor, white fir, were sampled along an elevational transect in the central Sierra Nevada mountains of California. This paper is based on data taken during the first two years' growth in a nearby nursery at Placerville, California. Ten of the thirteen growth, size, and needle morphological characteristics had significant differences between population samples, while eleven characteristics had significant differences between open-pollinated families within the population samples. High-elevation population samples were smaller in size and needle measurements, had fewer adaxial stomatal rows, blunter needle tips, and a shorter growing season. Most characteristic values were distributed ecotypically between the two lower- and the two higher-elevation population samples. Analyses of the patterns of variation demonstrated that characteristics of size and growth showed greater differentiation between population samples than did characteristics of needle morphology. It was concluded that the differences between characteristic patterns were caused by the more intense selection pressures acting upon the characteristics of size and growth. No consistent differences were discovered between these population samples in terms of total amounts of within-population genetic variation. The failure to find differences in intra-population variation may be due to the high rates of migration that probably exist within such a transect, and the fact that none of the populations included in this study are truly marginal. There were large differences between characteristics for total within-population variation; a positive but non-significant correlation was found between high coefficients of variation and those characteristics with much differentiation between populations. It is postulated that the maintenance of this intra-population variation was caused by microhabitat adaptations and gene flow. Evidence for increased genetic variation within open-pollinated families due to pollen immigration is presented, and its implications discussed.     

Variation in morphology of <Emphasis Type="Italic">Teucrium polium</Emphasis> aggr. populations in Bulgaria

The aim of this study was to investigate natural serpentine and non-serpentine populations of Teucrium polium aggr. and to document the differences in their morphological traits, as well as estimate which characters are most likely contributing to differentiation of the populations. Nine populations distributed both on and off serpentine soils were investigated, and 12 different morphological features of 270 individuals were studied by univariate and multivariate statistical analyses. The results did not demonstrate clear delimitation of the serpentine from the non-serpentine populations. Using these ecotypes as classification factors in the discriminant analysis, it can be stated that stem height, leaf length in the fifth leaf pair, internode length between the fourth and the fifth leaf pairs, and corolla tube length are features with the greatest discriminant ability. The character variation was generally higher for the vegetative features, while reproductive features failed to demonstrate clear differences. The similarity groupings indicated by the cluster analysis were synchronous with groups distinguished by both the discriminant and PCA analyses. Significant differences were observed between sets of vegetative characters in all populations investigated.      

Food preference and performance of the larvae of a specialist herbivore: variation among and within host-plant populations

Specialist herbivores are suggested to be unaffected by or attracted to the defense compounds of their host-plants, and can even prefer higher levels of certain chemicals. Abrostola asclepiadis is a specialist herbivore whose larvae feed on the leaves of Vincetoxicum hirundinaria, which contains toxic alkaloids and is unpalatable to most generalist herbivores. The food choice, leaf consumption and growth of A. asclepiadis larvae were studied to determine whether there is variation among and within host-plant populations in their suitability for this specialist herbivore. There was significant variation in food preference and leaf consumption among host-plant populations, but no differences were found in larval growth and feeding on different host-plant populations. A. asclepiadis larvae preferred host-plant populations with higher alkaloid concentrations, but did not consume more leaf material from plants originating from such populations in a no-choice experiment. There was also some variation in food preference of larvae among host-plant individuals belonging to the same population, suggesting that there was variability in leaf chemistry also within populations. Such variation in larval preference among host-plant genotypes and populations may create potential for coevolutionary dynamics in a spatial mosaic.     

Geographical variation in Fucus vesiculosus morphology in the Baltic and North Seas

Effects of stress and disturbance on morphology, reproductive effort, size and sex ratio were studied for Fucus vesiculosus populations from the Baltic Sea at Askö and the North Sea on the west coast of Sweden at Tjäm[otilde]. High morphological variation was found between Fucus populations, with significant differences in length and weight of individuals, thallus breadth, number of branches and receptacles and receptacle weight, not only between Baltic and North Sea populations but also between populations within the same area, differing in wave exposure. With increasing disturbance, individuals in both studied populations were smaller and less branched. Differences were observed in plant size, with longer, broader and more branched plants being found in Askö compared with Tjärnö. Fucus populations at Tjämö allocated more biomass to reproduction and had longer, heavier receptacles than at Askö. Although the observed morphological changes may be partly explained by differences in wave exposure and salinity between the two sites, it is not possible to rule out genetic differences between the Baltic and North Sea populations. However, it is unlikely that the variations observed within the populations and between populations from the same area are genetically determined.     

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。     

Large‐scale patterns in genetic variation,gene flow and differentiation in five species of European Coenagrionid damselfly provide mixed support for the central‐marginal hypothesis

Recently, an increased effort has been directed towards understanding the distribution of genetic variation within and between populations, particularly at central and marginal areas of a species’ distribution. Much of this research is centred on the central‐marginal hypothesis, which posits that populations at range margins are sparse, small and genetically diminished compared to those at the centre of a species’ distribution range. We tested predictions derived from the central‐marginal hypothesis for the distribution of genetic variation and population differentiation in five European Coenagrionid damselfly species. We screened genetic variation (microsatellites) in populations sampled in the centre and margins of the species’ latitudinal ranges, assessed genetic diversity (H_S) in the populations and the distribution of this genetic diversity between populations (F_ST). We further assessed genetic substructure and migration with Bayesian assignment methods, and tested for significant associations between genetic substructure and bioclimatic and spatial (altitude and latitude) variables, using general linearized models. We found no general adherence to the central‐marginal hypothesis; instead we found that other factors such as historical or current ecological factors often better explain the patterns uncovered. This was illustrated in Coenagrion mercuriale whose colonisation history and behaviour most likely led to the observation of a high genetic diversity in the south and lower genetic diversity with increasing latitude, and in C. armatum and C. pulchellum whose patterns of low genetic diversity coupled with the weakest genetic differentiation at one of their range margins suggested, respectively, possible range shifts and recent, strong selection pressure.     

FLUCTUATING ASYMMETRY IN CENTRAL AND MARGINAL POPULATIONS OF LYCHNIS VISCARIA IN RELATION TO GENETIC AND ENVIRONMENTAL FACTORS

Developmental instability in the form of increased fluctuating asymmetry can be caused by either genetic or environmental stress. Because extinctions can be attributed broadly to these factors, fluctuating asymmetry may provide a sensitive tool for detecting such stresses. We studied the level of fluctuating asymmetry of flowers of a perennial outcrossing plant species, Lychnis viscaria, both in natural and common-garden populations. The degree of flower asymmetry was higher in small, isolated, and marginal populations of the species range. These marginal populations also were the most homozygous. In the core area of the species' range, flowers were more symmetrical The level of asymmetry was correlated with both population size and heterozygosity. However, a partial correlation analysis revealed that when the impact of population size was controlled for, there was a negative relationship between fluctuating asymmetry and heterozygosity, whereas when controlling for heterozygosity, no relationship between population size and fluctuating asymmetry was found. This indicates that genetic consequences of small population size probably underlie the relationship between the level of asymmetry and population size. Results from a transplantation experiment showed that individuals subjected to a higher environmental stress had an increased level of asymmetry compared to control plants. In the common-garden conditions the level of fluctuating asymmetry did not differ between the central and marginal populations. This suggests that presumably both genetic and environmental factors affected to the higher level of asymmetry among marginal populations compared to central ones. In all we conclude that even though fluctuating asymmetry seems to be a sensitive tool for detecting stresses, results from studies focusing on only one factor should be interpreted with caution.     

Isozyme Variability in Plants Regenerated from Calli of Cereus peruvianus (Cactaceae)

Morphological and isozyme variation was observed among plants regenerated from callus cultures of Cereus peruvianus. Different morphological types of shoots (68%) were observed in 4-year-old regenerated plants, while no distinct morphological variants were observed in plants grown from germinated seeds. Isozyme patterns of 633 plants regenerated from calli and of 261 plants grown from germinated seeds showed no variation in isocitrate dehydrogenase isozyme, and the differential sorbitol dehydrogenase, alcohol dehydrogenase, malate dehydrogenase, acid phosphatase, and peroxidase isozyme patterns observed in regenerated plants were attributed to nonallelic variation. Allelic variation was detected at three isoesterase loci. The proportion of polymorphic loci for both populations was 13.6% and the deviation from Hardy–Weinberg equilibrium for the Est-1 and Est-7 loci observed in somaclones was attributed to the manner in which the regenerant population was established. The high values for genetic identity among regenerant and seed-grown plant populations are in accordance with the low levels of interpopulation genetic divergence. In somaclones of C. peruvianus, morphological divergence was achieved within a short time but was not associated with any isozyme changes and also was not accompanied by biochemical genetic divergence.     

Morphological and molecular variation in Mitchella undulata,with special reference to the systematic treatment of the dwarf form from Yakushima

Morphological and molecular variation in Mitchella undulata Siebold et Zucc. was examined to evaluate the genetic basis for recognizing the dwarf variety, M. undulata var. minor Masamune. Considerable variation in leaf size in M. undulata, but no obvious morphological discontinuities, were found between the normal and dwarf varieties. Instead, a weak cline running from the Pacific Ocean to the Sea of Japan was found. Anatomical observations of leaf blades revealed that the large variation in leaf size can be attributed to variation in the number of leaf cells and not to differences in cell size. A molecular analysis based on sequences of rDNA internal transcribed spacer regions indicated that there were two major genotypes in M. undulata with minor variation in haplotypes resulting from additional substitutions or putative recombination. The dwarf form from Yakushima was neither genetically uniform nor apparently differentiated from other populations. From these results, we conclude that the dwarf form of M. undulata should be treated at the rank of forma.     

Comparative morphology of populations of Monstera Adans. (Araceae) from natural forest fragments in Northeast Brazil using elliptic Fourier Analysis of leaf outlines

Summary  A comparative study of the leaf outline morphometrics of Monstera adansonii var. klotzschiana, M. adansonii var. laniata and M. praetermissa was carried out. The study focused on populations in isolated montane humid (brejo) forests of Ceará state in Northeast Brazil and compared them with populations from Amazonia and the Brazilian Atlantic Forest. Digitised outlines were prepared from a total of 1,695 field-collected leaf images from 20 populations, and elliptic Fourier analysis was used to generate matrices of coefficients, from which six shape variables (principal components) were extracted using Principal Components Analysis. Intra-population variability and inter-population differences were analysed with multivariate distance methods. Separate analyses were carried out for each of three leaf size classes (juvenile, submature, mature) because of the strong heteroblasty typical of this genus. Juvenile leaves were the least variable size class within populations of M. adansonii var. klotzschiana. The shape variables expressed very similar types of variation in all three size classes. The Ceará brejo populations of M. adansonii var. klotzschiana showed significant differences between mature leaf outlines in all pairwise comparisons; the Pacatuba population was the most distinct. The Ceará populations did not cluster together exclusively. In all three size classes, populations clustered together into their taxonomic groups, most clearly so in mature leaves. No correlation between morphological and geographic distance matrices was found, nor between morphological and molecular distance. The study showed that leaf outline shape is a practicable and useful quantitative trait for studying morphological variability at species, varietal and population levels. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Sex-specific physiological and growth responses to water stress in <Emphasis Type="Italic">Hippophae rhamnoides</Emphasis> L. populations

Dioecious plant species and those occupying diverse habitats may present special analytical problems to determine effects of environmental stress. Here, sex-specific physiological and growth responses of two contrasting sea buckthorn (Hippophae rhamnoides L.) populations were recorded after exposure to different watering regimes. The populations used were from wet and dry climate regions in China, respectively. In the semi-controlled environmental study, the well-watered and water-deficiency plants which were watered to 100 % and 50 % field capacity were used, respectively. Sexual differences in height growth (HT), dry matter accumulation (DMA), root/shoot ratio (RS), specific leaf area (SLA), net photosynthesis (A), transpiration (E), instantaneous water use efficiency (WUE_i) and carbon isotope composition (δ¹³C) between the male and female individuals were detected under water-deficiency treatment in both populations tested. However, these sexual differences were not detected under well-watered treatment. On the other hand, compared with the wet climate population, the dry climate population showed lower HT, DMA, SLA, A and E, and higher RS under both watering regimes. The dry climate population also showed higher WUE_i and δ¹³C as affected by water deficit than the wet climate population. These morphological and physiological responses to drought showed that the different populations and the different sexual individuals may employ different survival strategies under environmental stress. The male individuals and the dry climate population would have a conservative water-use strategy in response to drought stress.     

A genetic analysis of the photosynthetic properties of populations of Danthonia spicata that have different growth responses to light level

Summary Three populations of the grass Danthonia spicata were observed to have different rates of biomass accumulation when grown in common environment treatments. The populations were native to adjacent sites of different successional age and different levels of shading. Twelve individuals from each population were clonally replicated and two replicates were grown in each of two light treatments, 100% and 22% of unshaded sunlight. Following growth in the treatments the populations all exhibited the same mean light-saturated photosynthetic rate of 11.7 mol m^-2s^-1. This rate is intermediate for published values of sun and shade species and for species from along a successional gradient. There was no difference in photosynthetic rate among treatments. There was significant genetic variation for lightsaturated photosynthetic rate within populations but no significant differences among populations. The populations had similar leaf water potential values of-1.12 MPa in all treatments. There were significant differences among treatments and genotypes for specific leaf weight which resulted in significant differences among treatments and no significant differences among genotypes in light-saturated photosynthetic rate expressed on a leaf weight basis. Lightsaturated photosynthetic rate had a high heritability and low plasticity. We postulate that photosynthetic rate is under strong selection and that the observed rates permit populations of D. spicata to grow in a wide range of habitat light levels.     

Variation in Cnidoscolus texanus in relation to herbivory

Summary Twelve Oklahoma populations of Cnidoscolus texanus were investigated for morphological variation between and within populations, and between site classes with contrasting histories of use by domesticated livestock. Characters scored were numbers of stinging hairs on petioles and midribs, and depth of leaf lobing. Leaf lobing was not significantly variable within or among populations or grazing categories. Stinging hair numbers showed significant differences within and among populations. Petiole stinging hairs were more numberous in heavily grazed sites, suggesting population differentiation in response to the selective pressures of herbivory.