Variability of the hard dorsal idiosomal tegument pattern and other phenotypic characters as a manifestation of microevolution in Dermacentor marginatus (Acari: Ixodidae)

The pattern of male conscutum and female scutum in Dermacentor marginatus has been examined in 6 points of the East-European and Asian parts of its range (Fig. 1). A high rate of modification variability of the pattern has been recovered in each geographical sample. It is shown that populations differ reliably from each other by the length of the dark patches on the marginal flank of the conscutum (nkappabeta, npkappabeta, 3kappabeta); some samples also differ by the number of these patches, complexes of other patches of the scutum and conscutum, and as well (visually) by the color density of light fields and contrast between them and the dark background on the conscutum and scutum (Table 1, 2). Besides, the samples differ reliably at each stage by the complexes of morphometric characters (Table 3, 4). The sample from the Peter the First Ridge is most deviate from others by the color contrast and complex of patches both in males and females. However, by the patches of marginal flank and morphometric characters of both sexes and perimaginal stages this sample differ less from others. The sample from the Stavropol plateau having most variable pattern (Fig. 3, 1-9, 4, 1-3) is characterized by the greater number of statistically reliable differences by the patches (nkappabeta, npkappabeta, 3kappabeta) and morphometric characters of all stages of the ontogenesis from all other samples. Other geographical samples have intermedial position in regard to degree of differences from each other. The samples examined are considered as morphotypes; degree of differences between morphotypes is unequal. The phenotypic differences have mosaic dispersion among ontogenetic stages and do not show a coherent direction in comparing the morphotypes. Differential characters of the morphotypes and related tick species belong to different structures or their different parameters. Possible ways of morphotype's origin and geochronological characteristics of morphotypes are discussed.     

The intraspecific differentiation of the burrow tick Ixodes crenulatus (Ixodidae)

In four extensive disjunct areas of the distribution range of Ixodes crenulatus Koch, 1844 complexes of samples in 8 locations, and separate samples in two locations have been studied (fig.). Morphological characters (sizes of organs of idiosome, gnathostoma, legs, as well as some proportions of organs), which show statistically significant differences between complexes of samples on all corresponding stages of ontogenesis were revealed (tabl. 1, 2). Statistically significant differences were determined by Student's criterion (table. 3, 4). We use the term "transit" characters to denote these characters if they have to addition a similar tendency at all stages of ontogenesis. Complexes of samples showing statistically significant differences of transit characters are considered by us as morphotypes. Apparently these differences of morphotypes were formed evolutionally. In European disjunct area morphotypes divided into two groups on the basis of the degree of differences: western (A, B), and eastern (C, D, E). Degree of differences of morphotypes within each group is low, whereas one between of these two groups is high and corresponds to differences of morphotypes A, B from all Asian morphotypes (F, G, H, I, J), and of morphotypes C, D, E from Asian ones in mountain disjunct areas I. crenulatus (G, H, I, J). At the same time Eastern-European complex of morphotypes (C, D, E) is morphologically similar to the morphotype from north Kazakhstan disjunct area (F). All Asian morphotypes have high degree of differences one from the other. Species of the genus Marmota are initial hosts of I. crenulatus, they retain the main role as hosts in Eastern-European and Asian morphotypes: C, D, E, F--M. bobac bobac (Muller, 1776), G--M. baibacina centralis (Thomas, 1909), I, J--M. sibirica sibirica (Radde, 1862). The most aberrant in all characters morphotypes are the ones inhabiting European areas (A, B), where marmots are exterminated. These morphotypes parasitize on hibernating carnivores.     

The intraspecific taxonomic structure and identification of closely related species of ixodid ticks (Ixodidae)

Two variants of intraspecific morphological differentiation of 7 palearctic species of Ixodidae and respectively two variants of their intraspecific taxonomic pattern have been revealed on the basis of the study of geographic variation of each species. Three species--Ixodes pavlovskyi Pom., 1946, Haemaphysalis erinacei Pavesi, 1884, Hyalomma asiaticum Sch. et Schl., 1929, are subdivided into subspecies, and four ones--I. ricinus (L., 1758), I. persulcatus Sch., 1930, I. crenulatus Koch, 1844, Dermacentor marginatus (Sulzer, 1776),--into morphotypes by the degree of distinction of the whole complex of active stages in ontogenesis. Acknowledging the conventional character of the categories of subspecies and morphotype their criteria have been formulated for ixodid ticks. Paleogeographic and ecological grounds are given. Some data of geographic variation of immature stages favour the identification of closely related species in concrete localities.     

Variations in <Emphasis Type="Italic">Microtus arvalis</Emphasis> and <Emphasis Type="Italic">Microtus agrestis</Emphasis> (Arvicolinae,Rodentia) Dental Morphologies in an Archaeological Context: the Case of Teixoneres Cave (Late Pleistocene,North-Eastern Iberia)

Morphological and morphometric variations in the first lower molars of Microtus arvalis and Microtus agrestis from the late Pleistocene site of Teixoneres Cave (Barcelona, Spain) have been investigated in order to understand the modifications in dental patterns occurring in these two species in a peripheral region of their distribution area. It was possible to identify along the sequence differences in size and frequencies of morphotypes within the two populations, corresponding to environmental and climatic oscillations. Hypotheses to explain these intraspecific changes are discussed, and the variations are ascribed primarily to small-scale intraregional movements of these two rodent populations.     

The comparative morphology of the larynx in snakes

Young, B.A. 2000. The comparative morphology of the larynx in snakes. —Acta Zoologica (Stockholm) 81 : 177–193 The larynx and glottal tube were examined in 10 specimens each of 22 snake species. Qualitative analysis through dissection and clearing and staining revealed distinct morphological variation in the cartilage and intrinsic musculature of the larynx. Quantitative analysis of nine morphometric characters revealed laryngeal sexual dimorphism in three species, significant interspecific differences in the coefficients for every morphometric feature regressed on body size, no significant intraspecific differences in the log transformed means, and poor discrimination of species using principal component analysis. The observed variations in laryngeal morphology did not correlate with the phylogeny of the species, or with habitat preferences.     

Morphological variability between wild populations and inbred stocks of a Chinese minnow, Gobiocypris rarus

Gobiocypris rarus, a small, native cyprinid fish, is currently widely used in research on fish pathology, genetics, toxicology, embryology, and physiology in China. To develop this species as a model laboratory animal, inbred strains have been successfully created. In this study, to explore a method to discriminate inbred strains and evaluate inbreeding effects, morphological variation among three wild populations and three inbred stocks of G. rarus was investigated by the multivariate analysis of eight meristic and 30 morphometric characters. Tiny intraspecific variations in meristic characters were found, but these were not effective for population distinction. Stepwise discriminant analysis and cluster analysis of conventional measures and truss network data showed considerable divergence among populations, especially between wild populations and inbred stocks. The average discriminant accuracy for all populations was 82.1% based on conventional measures and 86.4% based on truss data, whereas the discriminant accuracy for inbred strains was much higher. These results suggested that multivariate analyses of morphometric characters are an effective method for discriminating inbred strains of G. rarus. Morphological differences between wild populations and inbred strains appear to result from both genetic differences and environmental factors. Thirteen characters, extracted from stepwise discriminant analysis, played important roles in morphological differentiation. These characters were mainly measures related to body depth and head size.     

Microevolutionary patterns in the common caiman predict macroevolutionary trends across extant crocodilians

Both extinct and extant crocodilians have repeatedly diversified in skull shape along a continuum, from narrow‐snouted to broad‐snouted phenotypes. These patterns occur with striking regularity, although it is currently unknown whether these trends also apply to microevolutionary divergence during population differentiation or the early stages of speciation. Assessing patterns of intraspecific variation within a single taxon can potentially provide insight into the processes of macroevolutionary differentiation. For example, high levels of intraspecific variation along a narrow‐broad axis would be consistent with the view that cranial shapes can show predictable patterns of differentiation on relatively short timescales, and potentially scale up to explain broader macroevolutionary patterns. In the present study, we use geometric morphometric methods to characterize intraspecific cranial shape variation among groups within a single, widely distributed clade, Caiman crocodilus. We show that C. crocodilus skulls vary along a narrow/broad‐snouted continuum, with different subspecies strongly clustered at distinct ends of the continuum. We quantitatively compare these microevolutionary trends with patterns of diversity at macroevolutionary scales (among all extant crocodilians). We find that morphological differences among the subspecies of C. crocodilus parallel the patterns of morphological differentiation across extant crocodilians, with the primary axes of morphological diversity being highly correlated across the two scales. We find intraspecific cranial shape variation within C. crocodilus to span variation characterized by more than half of living species. We show the main axis of intraspecific phenotypic variation to align with the principal direction of macroevolutionary diversification in crocodilian cranial shape, suggesting that mechanisms of microevolutionary divergence within species may also explain broader patterns of diversification at higher taxonomic levels.     

Geographic variation in Austrolebias luteoflamulatus Vaz‐Ferreira, Sierra & Scaglia (Cyprinodontiformes, Rivulidae)

The morphological variation in the annual killifish Austrolebias luteoflamulatus was studied using meristic and pigmentation characters analysed with univariate and multivariate analysis, and morphometric characters analysed with a geometric morphometry approach (thin plate spline). The results showed that morphological variation in this species was associated with a geographical pattern. In the case of meristic and pigmentation characters, the strongest distinction occured in males, while females were more similar among different drainages. Males from Rocha (eastern Uruguay) exhibited the greatest differences from the others. The number of dorsal fin rays and bars on the flanks were the characters that contributed most to this segregation. When morphometrics were considered, the three populations analysed could be distinguished well from each other in both sexes. Some of the characters that contributed to this pattern were dorsal fin origin, position of eyes and length of snout. Since pigmentation characteristics and some characters of the dorsal fin in males were involved in the differences observed, sexual selection could have played an important role in the differentiation of populations of this species. Since Austrolebias luteoflamulatus has a restricted geographical distribution, conservation of this species should include consideration of these populations individually since they might represent different evolutionary units.     

Morphological differences among three color morphotypes of Sebastes inermis (Scorpaenidae)

 Morphological differences among three color morphotypes occurring almost sympatrically (A, reddish type; B, blackish type; C, brownish type) of Sebastes inermis are clarified. Twenty-six morphometric characters and seven meristic characters were examined on about 20 specimens of each morphotype, 15 of the morphometric characters differing significantly. Principal component analysis (PCA) of the morphometric characters showed morphotype C to differ from the other two morphotypes. Canonical discriminant analysis (CDA) of the morphometric characters showed clear separation of each morphotype. Four meristic characters also differed significantly. Modes of pectoral fin ray counts of morphotypes A, B, and C were 15, 16, and 17, respectively, and the modes of anal fin ray counts of morphotypes B and C were 7 and 8, respectively. Many individuals of morphotype A had seven or eight anal fin rays. These differences suggested that the three morphotypes may be reproductively isolated from each other. Received: October 15, 2001 / Revised: March 14, 2002 / Accepted: April 9, 2002     

Discrimination of snipefish Macroramphosus species and boarfish Capros aper morphotypes through multivariate analysis of body shape

The existence of two species of the genus Macroramphosus Lacepède 1803, has been discussed based on morphometric characters, diet composition and depth distribution. Another species, the boarfish Capros aper (Linnaeus 1758), caugth along the Portuguese coast, shows two different morphotypes, one type with smaller eyes and a deeper body than the other, occurring with intermediate forms. In both snipefish and boarfish no sexual dimorphism was found with respect to shape and length relationships. However, females in both genera were on average bigger than males. A multidimensional scaling analysis was performed using Procrustes distances, in order to check if shape geometry was effective in distinguishing the species of snipefish as well as the morphotypes of boarfish. A multivariate discriminant analysis using morphometric characters of snipefish and boarfish was carried out to validate the visual criteria for a distinction of species and morphotypes, respectively. Morphometric characters revealed a great discriminatory power to distinguish morphotypes. Both snipefish and boarfish are very abundant in Portuguese waters, showing two well-defined morphologies and intermediate forms. This study suggests that there may be two different species in each genus and that further studies on these fish should be carried out to investigate if there is reproductive isolation between the morphotypes of boarfish and to validate the species of snipefish.     

Taxonomy of the Solidago virgaurea Group (Asteraceae) in Poland,with Special Reference to Variability along an Altitudinal Gradient

The morphological differentiation and taxonomic treatment of lowland and high-mountain morphotypes within the Solidago virgaurea group are controversial. To clarify the taxonomic status of these taxa, we conducted a morphometric analysis of 1,746 individuals from 80 localities along an altitudinal gradient from the lowlands of northern Poland to the Carpathians and Sudetes of southern Poland. Multivariate morphometric analyses, cluster analyses and principal component analyses, were used to examine the morphological differentiation within the S. virgaurea group in Poland. Canonical discriminant analysis was applied to determine the morphological characters that best discriminate among the taxa. The stability of the high-mountain Solidago minuta morphotype was tested in an experimental field established in lowland Poland; individuals transplanted from various mountain sites were cultivated at this site, and the morphotypes remained stable in terms of their floral and vegetative characters. Multivariate analyses revealed two morphologically distinct taxa in the S. virgaurea group, which correspond to lowland S. virgaurea s. str. and high-mountain S. minuta as recognised in some European floras. The most important morphological characters for distinguishing the taxa are the number of tubular florets per capitulum, inner involucral bract width and involucre height. Vegetative and inflorescence characters appear to have less taxonomic value because they changed continuously with altitude. A key for identifying S. virgaurea and S. minuta in Poland is presented.     

Phylogenomic and morphological data reveal hidden patterns of diversity in the national tree of Brazil,Paubrasilia echinata

Premise

Paubrasilia echinata (common names, pau brasil, brazilwood) is the national tree of Brazil and an endangered species endemic to the Brazilian Atlantic Forest. Over its wide distribution of 2000 km, its leaflets morphology exhibits extensive plasticity. Three morphotypes are commonly identified based on leaf size, but it is unclear if they represent distinct taxa or a single polymorphic species. This study aims to clarify the taxonomic position of the three morphotypes to inform conservation decisions.

Methods

A morphometric study of leaf characters of herbarium specimens was coupled with genetic analyses using genotype-by-sequencing data. We used maximum-likelihood and coalescent methods to evaluate the phylogenetic and population structure of the species. We compared these with a morphological dendrogram built from hierarchical clustering.

Results

Two of the three morphotypes formed separately evolving lineages, the third morphotype formed two geographically separate lineages, and northern trees with intermediate leaf morphology formed a separate fifth lineage. Leaflet size varied by over 35-fold, and although morphological clustering generally matched the genetic patterns, there were some overlaps, highlighting the cryptic diversity within this group.

Conclusions

Our genetic and morphological results provide some evidence that cultivated trees from different states in Brazil seem to have a limited genetic origin and do not reflect the broader genetic and geographical diversity of the species. As a result, more care is likely needed to preserve the overall genomic diversity of this endangered and iconic species.     

Variation in Body Shape across Species and Populations in a Radiation of Diaptomid Copepods

Inter and intra-population variation in morphological traits, such as body size and shape, provides important insights into the ecological importance of individual natural populations. The radiation of Diaptomid species (~400 species) has apparently produced little morphological differentiation other than those in secondary sexual characteristics, suggesting sexual, rather than ecological, selection has driven speciation. This evolutionary history suggests that species, and conspecific populations, would be ecologically redundant but recent work found contrasting ecosystem effects among both species and populations. This study provides the first quantification of shape variation among species, populations, and/or sexes (beyond taxonomic illustrations and body size measurements) to gain insight into the ecological differentiation of Diaptomids. Here we quantify the shape of five Diaptomid species (family Diaptomidae) from four populations each, using morphometric landmarks on the prosome, urosome, and antennae. We partition morphological variation among species, populations, and sexes, and test for phenotype-by-environment correlations to reveal possible functional consequences of shape variation. We found that intraspecific variation was 18-35% as large as interspecific variation across all measured traits. Interspecific variation in body size and relative antennae length, the two traits showing significant sexual dimorphism, were correlated with lake size and geographic location suggesting some niche differentiation between species. Observed relationships between intraspecific morphological variation and the environment suggest that divergent selection in contrasting lakes might contribute to shape differences among local populations, but confirming this requires further analyses. Our results show that although Diaptomid species differ in their reproductive traits, they also differ in other morphological traits that might indicate ecological differences among species and populations.     

Coexisting Cyclic Parthenogens Comprise a Holocene Species Flock in Eubosmina

Background

Mixed breeding systems with extended clonal phases and weak sexual recruitment are widespread in nature but often thought to impede the formation of discrete evolutionary clusters. Thus, cyclic parthenogens, such as cladocerans and rotifers, could be predisposed to “species problems” and a lack of discrete species. However, species flocks have been proposed for one cladoceran group, Eubosmina, where putative species are sympatric, and there is a detailed paleolimnological record indicating a Holocene age. These factors make the Eubosmina system suitable for testing the hypotheses that extended clonal phases and weak sexual recruitment inhibit speciation. Although common garden experiments have revealed a genetic component to the morphotypic variation, the evolutionary significance of the morphotypes remains controversial.

Methodology/Principal Findings

In the present study, we tested the hypothesis of a single polymorphic species (i.e., mixing occurs but selection maintains genes for morphology) in four northern European lakes where the morphotypes coexist. Our evidence is based on nuclear DNA sequence, mitochondrial DNA sequence, and morphometric analysis of coexisting morphotypes. We found significant genetic differentiation, genealogical exclusivity, and morphometric differentiation for coexisting morphotypes.

Conclusions

We conclude that the studied morphotypes represent a group of young species undergoing speciation with apparent reproductive barriers despite coexistence in the freshwater pelagic zone.