Within-island microgeographic variation in body dimensions and scalation of the skink Chalcides sexlineatus, with testing of causal hypotheses

Correlations, univariate analyses, multiple group principal component analyses, canonical analyses, contours and transects arc used to describe microgeographic variation in body dimensions and scalation of the scincid lizard Chalcides sexlineatus within the island of Gran Canaria. Patterns of geographic variation in both character systems show similar north-east/south-west clines, scalation variation is also altitude-related. Canonical variate analyses showed that the generalized patterns of geographic variation in scalation and body dimensions were largely unidimensional. Four hypotheses, each one predicting a unidimensional pattern of microgeographic variation, were tested simultaneously against each described pattern using partial correlation. This method did not reject north/south differences in climate and/or vegetation as a cause of the microevolution in body dimensions. Similarly, altitude and north/south differences in climate/vegetation were not rejected for scalation. Partial correlation rejected the hypothesis that there are two species of Chalcides with different ranges within Gran Canaria. Mantel tests rejected the altitude hypothesis alone as a cause of the generalized geographic variation in body dimensions but did not reject any of the proposed hypotheses as being causal for the generalized geographic variation in scalation. A consideration is made of how habitat variation could cause differential selection and comparisons drawn with a parallel pattern of microgeographic variation found in the colour pattern, scalation and size of a lizard on a neighbouring island.     

Clines: character number and the multivariate analysis of simple patterns of geographic variation

The stability of a clinal pattern geographic variation in relation to character number was investigated by the random selection of real characters. The characters used were selected from a set of 18 significant characters, drawn from four different character systems, and had a low within-group correlation. Principal component/coordinate analysis was used to elucidate the main pattern of geographic variation and the first principal coordinate was plotted against a geographical transect to illustrate this pattern. The existence of 'clinal' or 'monotically clinal' variation was assessed by the rank correlation between geographic position and coordinate score and the congruence between patterns of geographic variation was assessed by the correlation between the first principal coordinates of the appropriate analyses.
Two procedural models were used: A, the congruence between completely independent character sets based on 1–9 characters, and B, congruence between the 'total' analysis and those analyses based on 1–16 characters. The frequency of the types of pattern of geographic variation shown by these Model B analyses was also determined. Congruence is asymptotic in relation to character number and is described by various mathematical models, depending upon parameter and procedure. A broad pattern of clinal variation occurred in all analyses based on six or more characters and appears to be a stable feature of the racial affinities that is unlikely to be influenced by character choice. The details of the clinal pattern cannot be shown to be stable and may be influenced by the choice of character. This study had several findings in common with a previous study of categorical variation, e.g. the approximately hyperbolic relationship between character number and congruence in Model B, which may prove to be generally true in comparable situations.     

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.     

The adaptedness of the floral phenotype in a relict endemic, hawkmoth-pollinated violet. 2. Patterns of variation among disjunct populations

This study examines patterns of variation in quantitative floral traits among 18 disjunct populations of Viola cazorlensis (Violaceae), a relict endemic violet of south-eastern Spain. At all sites, the species is almost exclusively pollinated by a single species of day-flying hawkmoth. Differences between populations were significant for all traits examined, and population means exhibit a broad range of variation. When all characters are considered together, each population displays a unique combination of characters. Despite interpopulation differences in character means, local populations retain most of the variability of the entire species. Floral traits do not vary in unison among flowers, and at least four different subsets of independently varying traits are identifiable. Floral similarity between populations of V. cazorlensis was largely unrelated to geographic proximity, as revealed by analyses at both large and small geographic scales. The geographic pattern of floral variation among populations represents a random patchwork, with unique combinations of character means occurring randomly across the study region. Marked population differences in quantitative floral traits in spite of spatial constancy in the identity of pollinators, a disintegrated floral phenotype, and prevailingly random geographic variation among populations, suggest low adaptedness of the floral phenotype of V. cazorlensis to its current hawkmoth pollinators.     

Complex clines: the predictivity of complicated patterns of geographic variation portrayed by multivariate analysis

The stability of complex patterns of geographic variation was investigated by assessing the congruence between multivariate ordinations derived from randomly chosen real characters. Two series of populations were analysed representing two situations with complex patterns of geographic variation. The first, a ‘Eurasian’ series of populations, showed a strongly structured hierarchical pattern, the second, an ‘eastern’ series of populations, showed a more subtle complex pattern of smooth clines and steps. The characters were selected from a total of 81 (Eurasian) or 61 (eastern) within-population independent characters from six different systems. The congruence between ordinations of the geographical populations was measured by the rotational fit statistic, R². Three procedures were used to compare ordinations based on from two to up to 80 characters randomly chosen to give: A, completely independent character sets; B, subsets compared to the total set; and C, potentially overlapping sets. All three procedures showed that congruence between the ordinations was asymptotic in relation to character number. This relationship was described by one of two mathematical models (procedure B did not result in a hyperbolic model as found with simple patterns of geographic variation). Generally speaking, once a sufficient number of characters are used, the complex patterns of geographic variation are stable, reliable and predictive and not substantially influenced by character choice. The strongly structured hierarchical pattern required 15 or so characters to achieve reliability whilst the more subtle patterns required 20 or so characters. However, the addition of further characters does improve reliability in both cases. The greater percentage of variance portrayed by three-dimensional ordinations compared to two-dimensional ordinations is achieved at the cost of lower congruence when a sufficient number of characters are used. If case studies of geographic variation were to adopt these procedures (preferably using completely independent character sets; procedure A) the reliability of their results would be indicated).     

MICROGEOGRAPHIC PATTERNS OF GENETIC AND MORPHOLOGICAL VARIATION IN SAVANNAH SPARROWS (PASSERCULUS SANDWICHENSIS)

Surveys of genetic population structure are often limited to large geographic scales because geographically close populations are indistinguishable. Genetic uniformity across adjacent demes can be interpreted as evidence for cohesion (panmixia) or recent divergence. However, poor genetic resolution at microgeographic scales can also arise from the use of overly conservative (slowly evolving) markers. This study examines the ability of hypervariable, minisatellite loci to discriminate among geographically close populations of Savannah sparrows (Passerculus sandwichensis) and to track morphological differentiation at a microgeographic scale (interregional distance < 55 km). Savannah sparrows breeding at five island and two mainland sites in the Bay of Fundy, New Brunswick, Canada, show concordant patterns of variation in external morphology (seven characters) and multilocus DNA fingerprinting profiles (S_xy): island sparrows are phenotypically larger and genetically more similar to each other than they are to mainland sparrows. This pattern of variation is consistent with both adaptive (natural selection) and nonadaptive (genetic drift) mechanisms of population divergence. Based on minisatellite diversity, the effective size of both island and mainland populations is 37, an estimate substantially lower than census population sizes. These data are discordant with observations of sparrow vagility and abundance and suggest a closer examination of microgeographic patterns in avian systems.     

长白落叶松种群地理变异研究

根据17年生长白落叶松种源试验资料,从生态遗传学的观点出发,采用典型相关分析等方法,揭示了长白落叶松地理变异规律可以认为,以海拔垂直梯度为主、纬向渐变为辅的连续型变异;低海拔、低等效纬度的小北湖种源是长白落叶松优良基因资源中心,生长较快、生长期较长;水热因子的综合选择作用是长白落叶松地理变异的重要因素,其中温度是主导因子;在地理种群的遗传分化中,生长性状的分化最为明显,可以作为种源区区划的主要性状;从长白落叶松分布的低等效纬度地区向高等效纬度地区调种(一般不超过2个等效纬度),可望获得较大的遗传增益.     

The effect of insignificant characters on the multivariate analysis of simple patterns of geographic variation

The multivariate analysis of a set of significant characters portrays a simple geographic cline which is evident even when very few characters are used. The effect of adding insignificant characters to the set of significant characters is studied, as is the effect of replacing significant characters by insignificant characters. The former (addition) causes only a slight decline in congruence between patterns of geographic variation but the latter (replacement) causes a substantial decline in congruence. The congruence between patterns of geographic variation obtained by multivariate analysis of independent character sets is plotted against character number and gives an S-shaped relationship when insignificant or significant plus insignificant characters are used. This is distinct from the convex asymptotic curve obtained when only significant characters are used. In spite of the reduction in congruence caused by the use of insignificant characters, multivariate analysis of insignificant characters consistently revealed the 'same' geographic pattern (i.e. 'cline') as the set of significant characters. However, it required 10 times as many insignificant characters to achieve this.     

Multivariate analysis of the phenotypic relationships of the species of Paragalaxias and Galaxias (Pisces: Galaxiidae) in Tasmania

The 10 species of Galaxias in Tasmania, G. olidus from mainland Australia and the four species of Paragalaxias were studied using principal co-ordinates analysis (PCOA) and cluster analysis of a standardized Euclidean distance matrix based upon variate means, and by canonical variate analysis (CVA) conducted as a stepwise multiple discriminant analysis. Thirty-five variables comprising 30 morphometric and five meristic characters were analysed. The meristic characters were not included in the CVA. Excellent separation of the two genera was achieved in all analyses. The multivariate analyses were repeated on each genus separately to see if relationships suggested by the overall analysis remain stable. When the resultant groupings of species are compared for the different analyses, no consistent, distinct groupings of species within each genus are apparent. Despite the absence of distinct groupings, some trends in the affinities of some species are evident. In particular, species affinities as indicated by the CVA are more consistent with established opinions of species relationships. From the results of the study it is suggested that caution be exercised in the application of multivariate statistical analyses of morphological data to ichthyological systematics and phylogeny.     

Population differentiation in scalation of the Iberian rock lizard (Lacerta monticola)

The rock lizard Lacerta monticola is a climatic relict confined to three mountain ranges and one coastal region of the Iberian peninsula. Scalation was studied in specimens from localities encompassing all major parts of its range. Population differentiation was analysed using analyses of variance, z-transformations of within-locality character-correlations, canonical variate analyses, principal components analysis, and matrix association tests. Substantial differentiation was found between Pyrenees specimens and those from other localities, supporting a recent study which indicated that the Pyrenees populations could represent a different species. Considerable geographic variation is also found among populations from the remaining parts of its range, the pattern of which is not compatible with existing racial categories designated for this species.     

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.     

Adaptive variation in head size in Vipera berus L. populations

To prove that predators are morphologically adapted to the size of their prey one has to demonstrate that the morphological variation in the trophic apparatus is related to the prey size distribution and that the variation in the trait has some effect on individual fitness. I have studied geographic variation in relative head length (RHL) of adders, Vipera berus , on the Swedish mainland and on groups of islands in the Baltic Sea, and the relationship between RHL and physical condition, a character related to fitness. I also examined the relationship between RHL and sex and colour morph. Relative head length of adders was smallest on the mainland and increased on the islands with increasing body size of the main prey, Microtus agrestis , suggesting stabilizing selection for head size within each population. There was no difference in RHL between sexes or colour morphs. However, physical condition was positively correlated with RHL, indicating directional selection for larger heads. The observed pattern is interpreted as an evolutionary response to the geographic variation in body size of the main prey species and the smaller number of alternative prey species available on islands.     

Geographic patterns of song variation in four species of Malurus fairy‐wrens

Geographic variation in song is widespread among birds, particularly in species that learn vocalizations. The relationship between geographic distance and song variation is likely related to the degree of isolation between populations. To assess this effect of geographic isolation on song divergence, we examined patterns of geographic song variation in four species of Australian fairy‐wrens (Malurus), two with suspected histories of geographic isolation and two without. Song variation in all four species was consistent with patterns of isolation by distance, and allopatric subspecies in two species were more divergent in song than predicted by distance alone. Each species’ pattern was unique, and some interspecific variation could not be explained by geographic distance. These results indicate that patterns of geographic variation can be influenced by more than geographic distance and historical isolation alone. We suggest that morphological constraints, environmental influences, and sexual selection may all contribute to the variation observed for each species.     

Microgeographic Variation, Habitat Effects and Individual Signature Cues in Calls of Chiffchaffs Phylloscopus collybita canarensis

Variation of signals is a widespread feature in animal communication. Signals usually vary among signallers and in some cases vary across geographic areas. Individual variation provides the basis for individual recognition and thus has important implications for social interactions. Microgeographic variation can reveal insights into patterns of dispersal and into timing and mechanism of acquisition or development of behavioral traits. In addition, it may reflect adaptations of signals to the transmission characteristics of local habitat types. In song birds these kinds of variation have been documented in particular for male song. Here, we examined individual signature cues, site-specific variation, and variation among habitat types in tonal contact calls of 26 chiffchaffs, Phylloscopus collybita canarensis , recorded on two Canary islands. Multiparametric analyses of calls and subsequent discriminant function analyses revealed clear individual differences as well as microgeographic variation in call structure. Call structures differed not only between islands but also among the different locations on Tenerife, indicating that individuals on the same island do not share the same call, as suggested earlier. Calls recorded in different types of habitat, however, did not differ in structure as predicted by the 'acoustic adaptation hypothesis'. The findings indicate that individual recognition may be possible on the basis of calls and they contribute to the understanding of development of behavioural trails in relation to life history patterns such as the timing and pattern of dispersal.     

A multivariate analysis of geographic variation in morphology inPersoonia mollis (Proteaceae)

This study highlights the utility and limitations of multivariate procedures for the analysis and taxonomic interpretation of geographic variation within a complex species. Morphological data from herbarium specimens ofPersoonia mollis (Proteceae) were analyzed by two ordination procedures, multidimensional scaling (MDS) and canonical variate analysis (CVA). Support for the earlier recognition of nine infraspecific taxa was found. However, plots of canonical variate scores against latitute or longitude across parapatric subspecies boundaries revealed unexpected phenetic homogeneity within subspecies with sharp transitions between subspecies. The geographic variation in morphology withinP. mollis can be best described as a mosaic of nine recognisably distinct and internally homogeneous allopatric or parapatric taxa, where neighbouring taxa share surprisingly narrow zones of morphological transition at ecotones. The utility of these procedures was highlighted by the recovery of unexpected patterns of variation. However, questions of rank remain unresolved by phenetic analysis alone.     

Interaction between area effects and variation with habitat in Cepaea

Populations of Cepaea nemoralis in Warwickshire occupy habitats of considerable temporal stability, most being at least 250 years old, and some much older. As expected from earlier work, shell pattern polymorphisms in these populations show variation with habitat of a kind suggesting the operation of visual selection for crypsis. They also show patterns of microgeographical variation unrelated to habitat. Although of a less extreme character, this variation resembles the 'area effects' seen in downland populations of Cepaea , in the lack of coincidence of variation at different loci, and in the existence of stronger and larger scale patterns in banding than in colour morphs. A similar explanation is advanced for their occurrence: previous bottlenecks and colonization from small relicts with founder effects. The less marked character of the variation is expected from the greater habitat stability and continuity in Warwickshire compared with downland. A clear colonization effect is seen in the inverse relationship between age of habitat and frequency of yellow in woodland populations. The results suggest that population histories affect variation in Cepaea even in areas of relative habitat stability.     

Geographic variation in the garter snakes (Thamnophis sirtalis) of the north-central United States, a multivariate study

A study of geographic variation in three subspecies of garter snake (Thamnophis sirtalis sirtalis, T. s. parietalis, T. s. semifasciata) in the north-central United States indicates patterns of irregular non-clinal microgeographic variation for 40 characters of coloration, scalation, and internal anatomy. Multivariate analyses show that geographically close populations are not always phenetically close and that overall differentiation of the phenotype is greater in males than in females. Univariate and multivariate studies indicate that T. s. semifasciata Cope 1892, restricted to the Chicago area, is not uniform within its small range and that it has no constant distinguishing features: it is shown to be taxonomically invalid.     

Morphometric variation in a hybrid zone of two subspecies of Gerris costae (Heteroptera: Gerridae) in the Maritime Alps

Two morphologically distinct subspecies of Gerris costae form a contact zone extending from the Maritime to the Western Alps. Within this area canonical trend surface analysis revealed a geographic pattern of morphometric variation consistent with topography. From North to South, and from high to low elevation there was a transition from G. c. costae-like phenotypes to phenotypes resembling pure G. c. fieberi. The same pattern was confirmed with canonical variate analysis not taking geographic location into account; it is therefore not an artifact of trend surface analysis. Comparisons of the pattern of morphometric variation of laboratory-reared offspring with the pattern of their parents sampled from natural populations show that geographic variation is mostly determined genetically. Intermediate individuals from field populations probably are natural hybrids between the two subspecies, because laboratory-reared hybrids were intermediate between the offspring of pure strains. We did not find increased morphometric variation within the contact zone. This suggests unimpeded introgression and is in contrast with an increase in size variability that is predicted to be associated with a transition between uni- and bivoltine forms.