Components of morphological variation in Baikalian endemial cyclopid Acanthocyclops signifer complex from different localities

Use of traditional methods for morphological studies only permits the analysis of a small part of the information embodied in morphological structures. Besides comparing populations using the mean values of characters which allows one to estimate their morphological similarity, analysis of variation among individuals within a population can be informative. Variation among individuals consists of factorial and stochastic components. The factorial component is an upper estimate of genetic heterogeneity and thus permits one to evaluate the population's adaptability. The stochastic component (estimated by fluctuating asymmetry, i.e. random deviations from perfect bilateral symmetry), being a measure of developmental stability, is an indicator of a population's fitness. Assessment of measurement error is necessary for assessment of the true value of the stochastic component and for selection of the most informative characters. Such analysis allows one to extract additional information from morphological data in comparison with methods traditionally used on copepods. This approach was applied to an analysis of morphological variation in the study of the Baikalian endemic cyclopoid Acanthocyclops signifer (Mazepova) from three different isolated localities. Characters typically used in studies of taxonomy of this group are considered here. Measurement error was rather high (more than 50% of the stochastic component), which can be explained by technical difficulties of measuring the characters. All populations differ in the mean values of the characters. This shows the taxonomic heterogeneity of this group and reveals the necessity of its taxonomic revision. Populations also differ in the level of stochastic and factorial components of the total variance. The data are interpreted from the point of view of taxonomy and the possible evolution of the group.     

Analysis of asymmetries in the African fruit bats Eidolon helvum and Rousettus egyptiacus (Mammalia: Megachiroptera) from the islands of the Gulf of Guinea. I. Variance and size components of bilateral variation

A set of cranial characters was examined in the fruit bats Rousettus egyptiacus and Eidolon helvum to compare trends and relative importance of major components of bilateral morphometric variation, and their relationship with character size. Using two‐way, sides‐by‐individuals ANOVA , four components of variation were estimated for each bilateral variable: individual variation (I), directional asymmetry (DA), non‐directional asymmetry (NDA) and measurement error (E). Both species exhibit similar major trends of variation in asymmetry across characters, as shown by principal component analysis, using variance components as variables. Degree of interspecific congruence among characters was confirmed by a two‐way ANOVA with species and variance components as fixed factors. Congruence of asymmetry patterns between species suggests that the concept of population asymmetry parameter (PAP) could be extended to higher hierarchies. PAPs above the species level may result from common mechanisms or similar developmental constraints acting on species’ buffering capacities and morphological integration processes.     

Genetic and environmental variations in quantitative characters in fishes: a comparative analysis of monoclonal triploid and bisexual tetraploid spined loaches (<Emphasis Type="Italic">Cobitis</Emphasis>, Cobitidae)

The heritability estimates of 25 external morphometric characters and 23 craniometric indices are obtained by use of variances in monoclonal all-female triploids and bisexual tetraploids of spined loaches (genus Cobitis, Cobitidae) collected from the same breeding biotope. Most of studied traits demonstrate low heritability confirming previous conclusion on the similarity between external morphometric characters and craniological indices in relative effects of genetic and environmental components in their total phenotypic variation. Low heritability estimates in most of external morphological traits correspond to their low diagnostic value in Cobitis species. As a whole, in spite of certain deviations, studies on clonal forms do not refute the concept on higher heritability estimates in diagnostically significant traits in comparison with traits without diagnostic values in the same taxonomic group. Low heritability in most morphometric traits more probably is resulted from their low additive genetic variation caused by strong selection of evolutionary developed specific body shape in spined loaches, because strong selection should reduce the genetic variance in body proportions to minimal size. Sex differences observed in heritability estimates should be interpreted as a result of linkage of several additive genes controlling these traits to sex chromosomes. A few characters demonstrating high heritability estimates up to 0.492–0.580 are of great interest for taxonomic and phylogenetic studies in genus Cobitis and related taxa.     

Evolution of termite functional diversity: analysis and synthesis of local ecological and regional influences on local species richness

Aim To (1) describe termite functional diversity patterns across five tropical regions using local species richness sampling of standardized areas of habitat; (2) assess the relative importance of environmental factors operating at different spatial and temporal scales in influencing variation in species representation within feeding groups and functional taxonomic groups across the tropics; (3) achieve a synthesis to explain the observed patterns of convergence and divergence in termite functional diversity that draws on termite ecological and biogeographical evidence to‐date, as well as the latest evidence for the evolutionary and distributional history of tropical rain forests. Location Pantropical. Methods A pantropical termite species richness data set was obtained through sampling of eighty‐seven standardized local termite diversity transects from twenty‐nine locations across five tropical regions. Local‐scale, intermediate‐scale and large‐scale environmental data were collected for each transect. Standardized termite assemblage and environmental data were analysed at the levels of whole assemblages and feeding groups (using components of variance analysis) and at the level of functional taxonomic groups (using correspondence analysis and canonical correspondence analysis). Results Overall species richness of local assemblages showed a greater component of variation attributable to local habitat disturbance level than to region. However, an analysis accounting for species richness across termite feeding groups indicated a much larger component of variation attributable to region. Mean local assemblage body size also showed the greater overall significance of region compared with habitat type in influencing variation. Ordination of functional taxonomic group data revealed a primary gradient of variation corresponding to rank order of species richness within sites and to mean local species richness within regions. The latter was in the order: Africa > south America > south‐east Asia > Madagascar > Australia. This primary gradient of species richness decrease can be explained by a decrease in species richness of less dispersive functional taxonomic groups feeding on more humified food substrates such as soil. Hence, the transects from more depauperate sites/regions were dominated by more dispersive functional taxonomic groups feeding on less humified food substrates such as dead wood. Direct gradient analysis indicated that ‘region’ and other large‐scale factors were the most important in explaining patterns of local termite functional diversity followed by intermediate‐scale geographical and site variables and, finally, local‐scale ecological variables. Synthesis and main conclusions Within regions, centres of termite functional diversity lie in lowland equatorial closed canopy tropical forests. Soil feeding termite evolution further down food substrate humification gradients is therefore more likely to have depended on the long‐term presence of this habitat. Known ecological and energetic constraints upon contemporary soil feeders lend support for this hypothesis. We propose further that the anomalous distribution of termite soil feeder species richness is partly explained by their generally very poor dispersal abilities across oceans. Evolution, radiation and dispersal of soil feeder diversity appears to have been largely restricted to what are now the African and south American regions. The inter‐regional differences in contemporary local patterns of termite species richness revealed by the global data set point to the possibility of large differences in consequent ecosystem processes in apparently similar habitats on different continents.     

Tracing geographical patterns of population differentiation in a widespread mangrove gastropod: genetic and geometric morphometrics surveys along the eastern African coast

In the present study, we assessed the inter‐ and intrapopulation genetic and morphological variation of Cerithidea decollata along the eastern coast of Africa. The population structure of C. decollata along the latitudinal gradient was examined by sequencing 420 bp of the mitochondrial cytochrome c oxidase I (COI) gene in 172 snails from 29 sites, in a combined analysis with geometric morphometrics in 1799 snails from 32 sites. Analysis of molecular variance and spatial analysis of molecular variance showed a moderate spatial population differentiation from Kenya to the Republic of South Africa, suggesting genetic divergence between the northern, central, and southern regions. This structure appears to be the result of life‐history traits combined with oceanographic features. Haplotype network and mismatch analysis suggest a recent population expansion during the Holsteinian interglacial period in the northern region and several colonization events in the central and southern regions. The morphometric approach suggests that morphological variation in shell shape is somewhat independent of the genetic divergence, revealing an overlap of shape across the latitudinal gradient but significant differences among‐population at a local level. This may indicate that similar ecological pressures are acting along the coast, leading to the occurrence of similar morphological characters. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 647–663.     

Molecular phylogeny and ecogeography of Onobrychis viciifolia Scop. (Fabaceae) based on nrDNA ITS sequences and genomic ISSR fingerprinting

In this study 40 specimens from 8 representative Iranian populations of Onobrychis viciifolia SCOP. were collected from their natural habitats. The specimens were biometrically assessed using 43 quantitative and 15 qualitative morphological characters. At each sample site we recorded ecogeographical variables. In order to evaluate the difference between the variation due to phenotypic responses and genetic adaptation, we generated nucleotide sequence data from the internal transcribed spacer of the nuclear rDNA genes (ITS) and carried out genomic fingerprints using inter‐simple sequence repeat PCR (ISSR). Cluster analysis of morphological characters divided the eight populations into three major groups. Leaf length, leaflet length and width, calyx length, plant height, and stem length were introduced as diagnostic characters for three different morphological groups. Furthermore, canonical correspondence analysis (CCA) of ecogeographical data showed correlations between morphological variations and ecogeographical factors. Clay%, saturation percentage (SP), total neutralizing value (TNV%), texture, minimum temperature, and geographic separation were the main environmental variables associated with morphological groups of O. viciifolia. ISSR analysis revealed three main groups which are in correspondence with morphological groups, indicating that the three morphological groups have been shaped by genetic and phenotypic responses to environmental conditions. (© 2012 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

华南沿海地区西部入海水系中间黄颡鱼的形态变异及地理分化

为探讨不同水系中间黄颡鱼(Pelteobagrus intermedius)的形态差异及地理分化问题,采用形态学方法, 测量12个水系中间黄颡鱼150尾样本的18个形态性状,进行主成分分析.其结果显示,前3个主成分累积贡献率为58.918%,并可将这12个水系的群体分为3簇.华南沿海地区西部诸独立入海小水系中的中间黄颡鱼群体与海南岛群体之间的相似性大于与西江干流群体的相似性.防城港地区3条直接入海的小水系的群体在形态特征上已发生了一定程度的分化,其分化主要表现在吻、眼和尾的变异.     

Stromatoporoid palaeoecology in the Frasnian (Upper Devonian) Belgian platform,and its applications in interpretation of carbonate platform environments

Abstract: Stromatoporoid faunas in the Frasnian of southern Belgium are abundant in the carbonate platform environments present in this area. Stromatoporoids dominate the large skeletal organisms and occur principally in biostromes. The stromatoporoid assemblage is represented by a small number of taxa. Stromatoporoid genera include Actinostroma, Amphipora, Atelodictyon, Clathrocoilona, Salairella, Stachyodes, Stictostroma, Stromatopora and Trupetostroma which are present in environments ranging from the outer, outer intermediate, inner intermediate and inner zones and associated biostromes. Most large skeletal stromatoporoids are low profile, which reinforces the conclusions of previous studies that low‐profile growth forms were the most successful stromatoporoid forms. These low‐profile forms are likely to have been important sediment stabilisers that may have led to expansion of the carbonate factory. Growth forms vary between facies, indicating some degree of environmental control on form; for example, laminar in the intermediate zone, bulbous and domical in the inner and outer zones. Stromatoporoid taxa vary in occurrence across the environmental gradient from shallow to deep. There is some taxonomic control on growth forms, with some taxa showing more variability than others in different environments.     

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.     

The oldest labechiid stromatoporoids from intraskeletal crypts in lithistid sponge–Calathium reefs

Early Ordovician (early Floian) reefs of South China include lithistid sponge–Calathium reefs with a three‐dimensional skeletal framework. These structures are among the first post‐Cambrian skeletal‐dominated reef structures and provides an opportunity to test how the novel metazoan builders changed the environments and increased topographic complexity within benthic communities. We document the oldest labechiid stromatoporoid (Cystostroma) in a lithistid sponge–Calathium reef of the Hunghuayuan Formation in southeastern Guizhou, South China. These earliest stromatoporoids may have originated in reefs, and we argue that the complex topography created by the hypercalcified sponge Calathium facilitated the emergence of stromatoporoids. Beyond Cystostroma, keratose sponges, Pulchrilamina (hypercalcified sponge) and bryozoans have also inhabited in the micro‐habitats (cavities and hard substrates) provided by Calathium. These findings suggest that ecosystem engineering by Calathium played an important role in the further diversification of reefs during the Ordovician.     

Natural selection on the genetical component of variance in body condition in a wild bird population

Although there is substantial evidence that skeletal measures of body size are heritable in wild animal populations, it is frequently assumed that the nonskeletal component of body weight (or ‘condition’) is determined primarily by environmental factors, in particular nutritional state. We tested this assumption by quantifying the genetic and environmental components of variance in fledgling body condition index (=relative body weight) in a natural population of collared flycatchers (Ficedula albicollis), and compared the strength of natural selection on individual breeding values with that on phenotypic values. A mixed model analysis of the components of variance, based on an ‘animal model’ and using 18 years of data on 17 717 nestlings, revealed a significant additive genetic component of variance in body condition, which corresponded to a narrow sense heritability (h²) of 0.30 (SE=0.03). Nongenetic contributions to variation in body condition were large, but there was no evidence of dominance variance nor of contributions from early maternal or common environment effects (pre‐manipulation environment) in condition at fledging. Comparison of pre‐ and post‐selection samples revealed virtually identical h² of body condition index, despite the fact that there was a significant decrease (35%) in the levels of additive genetic variance from fledging to breeding. The similar h² in the two samples occurred because the environmental component of variance was also reduced by selection, suggesting that natural selection was acting on both genotypic and environmental variation. The effects of selection on genetic variance were confirmed by calculation of the selection differentials for both phenotypic values and best linear unbiased predictor (BLUP) estimates of breeding values: there was positive directional selection on condition index both at the phenotypic and the genotypic level. The significant h² of body condition index is consistent with data from human and rodent populations showing significant additive genetic variance in relative body mass and adiposity, but contrasts with the common assumption in ecology that body condition reflects an individual’s nongenetic nutritional state. Furthermore, the substantial reduction in the additive genetic component of variance in body condition index suggests that selection on environmental deviations cannot alone explain the maintenance of additive genetic variation in heritable traits, but that other mechanisms are needed to explain the moderate to high heritabilities of traits under consistent and strong directional selection.     

Principal component analysis as an alternative treatment for morphometric characters: phylogeny of caseids as a case study

In a recent study, the phylogeny of Caseidae (a herbivorous family of Palaeozoic synapsids belonging to the paraphyletic grade known as pelycosaurs) was analysed with a dataset employing more than three hundred continuous morphological characters in an effort to follow the principles of total evidence. Continuous characters are a source of great debate, with disagreements surrounding their suitability for and treatment in phylogenetic analysis. A number of shortcomings were identified in the handling of continuous characters in this study of caseids, including the use of gap weighting to discretize the characters and potential issues with redundancy and character non‐independence. Therefore, an alternative treatment for these characters is suggested here. First, rather than using gap weighting, the continuous characters were analysed in the program TNT, in which the raw values can be treated as continuous rather than discrete. Second, prior to the phylogenetic analysis, the continuous characters were subjected to a log‐ratio principal component analysis, and then the principal components were included in the character matrix rather than the raw ratios. Analysing the original data in TNT produced little difference in the results, but using the principal components as continuous characters resulted in alternative positions for Caseopsis agilis, Ennatosaurus tecton and Caseoides sanangeloensis. The differences are judged to be due to the reduced redundancy of the characters, the smaller number of principal components not overwhelming the discrete characters and the use of a scaling method which allows principal components with a higher variance to have a greater influence on the analysis. The positions of highly fragmentary fossils depended heavily on the method used to treat the missing characters in the principal component analysis, and so the method proposed here is not recommended for analysing very incomplete taxa.     

Climatic variability and body size variation in the muskrats (Ondatra zibethicus) of North America

Summary Patterns of geographic variation in nine morphological characters of adult muskrats (Ondatra zibethicus) are investigated by multiple regression and canonical correlation analysis. Three variables describing the annual precipitation regimen account for 16 to 33% of the variance in each of the skeletal measurements. Highly variable precipitation patterns result in low rates of germination and survivorship for preferred aquatic food plants which in turn reduces average total food availability for muskrats. In large individuals nutritive demands may exceed supply, thus selection favors smaller body size in areas of low food availability.After removing the variation attributable to three precipitation variables, the residual variation in the morphological variables is exposed to canonical correlation analysis with a set of 10 environmental and geographic variables. A canonical variate loaded for climatic seasonality accounts for 60% of the variance in a canonical variate of the morphological residuals which load as a general body size variate. It is argued that seasonality is a major factor selecting for large body size in muskrats and other organisms. During seasonal periods of resource abundance natural selection favors individuals with rapid growth to a large size, while concurrently enhancing survivorship through oncoming periods of resource shortage.     

Variability and correlations in muskmelon in relation to the cultivation method

Summary Six fruit characters have been measured in 23 cultivars of Cucumis melo, representing a wide geographical range. Plants were grown both in the greenhouse and in the field. When the 23 cultivars were analyzed together, the largest component of variance was found between cultivars under both growth conditions, suggesting the existence of large genetic diversity for all the characters studied. Generally, variance between plants within cultivars was less than or equal to variance between fruits within plant. This indicates that environmental variation is the most important part of the variation within cultivars. Correlations between pairs of characters at cultivar, plant and fruit levels were calculated from the variance-covariance components. In the majority of paired traits, the correlation values indicated that genetic and environmental factors may act in the same direction.     

Inferring evolutionary scenarios with geostatistics and geographical information systems for the viperid snakes Vipera latastei and Vipera monticola

The present study aimed to infer evolutionary scenarios for Vipera latastei and Vipera monticola in the Iberian Peninsula and the Maghreb through the identification of spatial patterns in morphological character variation and biogeographic patterns in morphological variability distribution. Ten morphological traits from 630 vipers were analysed with geostatistic and ecological niche modelling in a geographical information system. Interpolation by Kriging was used to generate surfaces of morphological variation, which were combined with spatial principal components analysis (SPCA). Putative morphological differentiated groups generated by SPCA maps were tested with discriminant function analysis (DFA). Maximum entropy modelling and nine environmental variables were used to identify factors limiting the distribution of groups and areas for their potential occurrence. Groups supported by DFA were: Western Iberia, Eastern Iberia, Rif plus Middle Atlas, Algeria, and High Atlas. Their distribution is influenced by common environmental factors such as precipitation. Areas of probable sympatry between Iberian groups matched the morphological clines observed by geostatistics tools. Geographic variation patterns in V. latastei‐monticola are probably due to vicariant separation of Iberian and African populations during the opening of the Strait of Gibraltar, and population refugia during the Quaternary glaciations with secondary contact. The taxonomic status of northern Morocco and Algerian groups should be further investigated. We conclude that geostatistics and niche‐modelling tools are adequate to infer morphological variability across wide geographic ranges of species. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 790–806.     

高原鼢鼠(Eospalax baileyi)头骨形态的地理分化

运用单因素方差分析、主成分分析、判别分析和聚类分析,对高原鼢鼠(Eospalax baileyi)4个地区(临潭、天祝、玛曲和碌曲)8个地理群体的头骨形态特征进行综合分析,比较13个特征变量的地理分异,挖掘形态特征的主要变量,并探讨其形态变异与环境的关系。结果发现,群体间13个特征变量均存在显著差异,主成分分析表明13个头骨特征变量可用3个主成分来表述,其累计贡献率达96.981%。颅全长、上齿列长及齿隙长是高原鼢鼠头骨发生分化的主要变量。天祝种群与碌曲种群,及玛曲种群与临潭种群都有重叠,其中碌曲的两个种群相似性最高,玛曲种群和临潭种群与其他种群几乎无重叠,显示完全分化,聚类分析结果与判别分析结果一致。头骨形态与地理因子的相关性分析发现头骨形态大小与海拔成显著的正相关关系。综合分析认为地理隔离、栖息地海拔因子影响下的生态条件等是高原鼢鼠头骨分化的主要因素。     

CLASSIFICATION AND CONTROLS OF INTERNAL BANDING IN PALAEOZOIC STROMATOPOROIDS AND COLONIAL CORALS

Abstract:  Palaeozoic corals and stromatoporoids exhibit a variety of internal banding phenomena, many of which have been commonly interpreted as annual growth bands. We evaluate bands through analysis of colonial corals and stromatoporoids from three stratigraphic intervals: Upper Ordovician of Manitoba Canada, and Llandovery–Wenlock and Ludlow of Gotland, Sweden. Banding features are divided into four categories: (1) absence of banding; (2) density banding formed by variation in density or form of elements; (3) growth-interruption banding indicating growth cessation and regeneration; and (4) post-mortem banding caused by compaction or diagenesis. For discrimination of band types, it is essential to examine internal structures and skeletal margins in thin sections or acetate peels. Species vary considerably in degree and type of banding; each has a distinct pattern of variation. We propose criteria to determine if banding is consistent with seasonally induced growth variation: (1) consistency in band character and thickness; (2) continuity of skeletal growth; (3) marginal features; and (4) evidence of diagenetic alteration. Density bands in tabulate and rugose corals probably represent annual growth variations, but results for stromatoporoids are more ambiguous; although stromatoporoids commonly show banding, unequivocal density banding is poorly developed and growth interruption generated most stromatoporoid banding. Cerioid rugose and tabulate corals possess the thickest density bands; the thinnest bands are in stromatoporoids and heliolitid tabulates.