Variation of allozyme loci in endemic gammarids of Lake Baikal

The outstandingly rich and highly endemic fauna of Lake Baikal includes more than 200 species of gammarids (Crustacea, Amphipoda), most of which occur only in Baikal. Allelic variation at 23 enzyme loci has been studied for the first time in four species of Baikalian gammarids. The levels of polymorphism are similar to those observed in other invertebrates, including gammarids: c. 20–25% of the loci are polymorphic with 2.53 to 3.3 alleles and mean heterozygosity of 0.048 to 0.105 per polymorphic locus. The genotype frequences in local populations in most cases markedly deviate from Hardy-Weinberg equilibrium with a deficiency of heterozygotes. Populations were shown to be strongly subdivided even over a range of less than 100 m. On the larger scale genetic differences among the populations do not show any strong correlation with geographical separation. Genetic distances among species were found to be rather low: from 0.159 between closely related, but morphologically clearly distinct species of the genus Spinacanthus to 0.546 between the representatives of different genera. We discuss these patterns of population structure with respect to the problem of the origin of endemic faunas in ancient lakes. Our results suggest that speciation occurred rapidly and relatively recently. Genetic distances within and among species support models of speciation based on the founder effect or differentiation of a peripheral population rather than on divergence of large subpopulations. The rate of morphological evolution in Baikalian gammarids is very high compared with their biochemical evolution.     

Morphological variability and developmental instability in subpopulations of the Eurasian badger (Meles meles) in Denmark

Aim Local populations from different geographical regions may differ in the selection regimes to which they are exposed. Differences in environmental factors and population density may affect the relative importance of different selective forces (e.g. natural vs. sexual selection). We suggest a direction of investigation concerned with the developmental instability of morphological traits. The goal is to disclose putative small‐scale geographical differences in the evolutionary forces, which may be hard to detect. Location Craniometrical investigations were carried out on ninety‐eight skulls and teeth of the Eurasian badger (Meles meles) collected during the period 1995–97 from three different populations in Denmark. One of these thrives at low population density, whereas the two others are characterized by high local density. Methods The skulls were investigated for developmental instability (DI) using fluctuating asymmetry (FA) as its estimator. FA was measured on canines, molars, premolar teeth and other skull and mandible traits. For the statistical analyses, we applied nonparametric permutation tests. Results Evidence was found suggesting differentiation among populations in mean degree of FA, and the FA values measured on canines were higher in the high‐density populations. FA of the canines was significantly higher in males than females, in contrast to FA of the other traits. Evidence of a negative relationship between canine size and their FA was found, whereas no significant correlations were found between the molar and premolar teeth measures and their FA. Main conclusions Our results suggest that canines could be under directional selection stemming from intrasexual competition, which may be stronger in high‐density zones. The other teeth investigated seem to be under a stabilizing regime hence their FA is mainly affected by environmental stresses. The negative relationship between canine size and FA found in males suggests the capacity of badgers to respond in an evolutionary way to environmental changes, despite the low genetic variability previously found at the molecular level.     

Investigating South American biogeographic history using patterns of skull shape variation on Cerdocyon thous (Mammalia: Canidae)

Patterns of geographic variation of the canid Cerdocyon thous have historically been obscured by its remarkable intraspecific morphological variability. The observed distribution is highly associated with phytophysiognomy, a feature considered highly dynamic along geological time. In the present study, we tested whether vegetation distribution during the Holocene Glacial Maximum of South America (HGM) explains the patterns of morphological variation within Cerdocyon thous . The species was divided in groups according to paleohabitats that could support their presence during the HGM, and then tested for differences in skull morphometrics. The results obtained demonstrate that the climatic changes during the HGM influenced the population structure of this species, resulting in the establishment of geographical groups with different degrees of morphological cohesion. Higher morphological cohesion found in the Northern group might be explained by the marked discontinuity between its geographical range and the rest of the species'distribution. The Eastern and Southern morphological divergence is less striking and, although this could be related to past vegetation distribution, the disappearance of those barriers leads to a population structure that could be slowly breaking down.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 77–84.     

Genetic diversity and phylogeography of Siberian roe deer,Caproulus pygargus,in central and peripheral populations

Current understanding of phylogeographical structure and genetic diversity of Siberian roe deer remains limited mainly due to small sample size and/or low geographical coverage in previous studies. Published data suggest at least two phylogroups: western (Ural Mountains and Western Siberia) and eastern (east from lake Baikal, including the Korean peninsula), but their phylogenetic relationship remains unclear. Combined sequences of cytochrome b (1140 bp) and the mtDNA control region (963 bp) were analyzed from 219 Siberian roe deer from 12 locations in Russia, Mongolia, and South Korea, which cover a large part of its range, to assess genetic diversity and phylogeographical status. Special emphasis was placed on the demographic history and genetic features of central, peripheral, and isolated populations. Results of median‐joining network and phylogenetic tree analyses indicate that Siberian roe deer from the Urals to the Pacific Ocean are genetically diverse and that geographical distribution and composition of haplogroups coincide with previously described ranges of the subspecies Capreolus pygargus pygargus and Capreolus pygargus tianschanicus. We found that peripheral populations in the northwestern parts of the species range (Urals), as well as the isolated population from Jeju Island, are genetically distinct from those in the core part of the range, both in terms of genetic diversity and quantitative composition of haplogroups. We also found that northwestern (Urals) and northern (Yakutia) peripheral populations share the same haplogroup and fall into the same phylogenetic clade with the isolated population from Jeju Island. This finding sheds light on the taxonomic status of the Jeju Island population and leads to hypotheses about the discordance of morphological and genetic evolution in isolated populations and specific genetic features of peripheral populations.     

中国淡水硅藻新记录——偏肿内丝藻兴凯变种

于2016年5月对浙江省仙居国家公园进行藻类多样性调查时,发现了中国淡水硅藻的1个新记录种——偏肿内丝藻兴凯变种[Encyonema ventricosum var.hankensis(Skvortzov) RodionovaPomazkina]。该文采用光镜和扫描电镜对偏肿内丝藻兴凯变种浙江种群的形态特征进行了观察和描述,并将其与该种的其他地理种群及内丝藻属其他相似物种的形态特征进行比较,同时讨论了该物种的生境特征。结果表明:(1)偏肿内丝藻兴凯变种壳面均具明显背腹之分,背缘明显拱起,腹缘轻微波曲;端部呈头状至喙状。(2)偏肿内丝藻兴凯变种浙江地理种群在背部中央区域具一孤点,而其兴凯种群对孤点这一特征未有描述。(3)偏肿内丝藻兴凯变种在俄罗斯的兴凯湖和贝加尔湖等地有分布,该种在中国是首次报道。     

Molecular phylogeny and phylogeography of the Cuban cave-fishes of the genus Lucifuga: evidence for cryptic allopatric diversity

Underground environments are increasingly recognized as reservoirs of faunal diversity. Extreme environmental conditions and limited dispersal ability of underground organisms have been acknowledged as important factors promoting divergence between species and conspecific populations. However, in many instances, there is no correlation between genetic divergence and morphological differentiation. Lucifuga Poey is a stygobiotic fish genus that lives in Cuban and Bahamian caves. In Cuba, it offers a unique opportunity to study the influence of habitat fragmentation on the genetic divergence of stygobiotic species and populations. The genus includes four species and one morphological variant that have contrasting geographical distributions. In this study, we first performed a molecular phylogenetic analysis of the Lucifuga Cuban species using mitochondrial and nuclear markers. The mitochondrial phylogeny revealed three deeply divergent clades that were supported by nuclear and morphological characters. Within two of these main clades, we identified five lineages that are candidate cryptic species and a taxonomical synonymy between Lucifuga subterranea and Lucifuga teresinarum. Secondly, phylogeographic analysis using a fragment of the cytochrome b gene was performed for Lucifuga dentata, the most widely distributed species. We found strong geographical organization of the haplotype clades at different geographic scales that can be explained by episodes of dispersal and population expansion followed by population fragmentation and restricted gene flow. At a larger temporal scale, these processes could also explain the diversification and the distribution of the different species.     

Evolutionary and biogeographical implications of variation in skull morphology of raccoon dogs (Nyctereutes procyonoides,Mammalia: Carnivora)

Morphological characteristics reflect geographical variation resulting from adaptation to varying environmental conditions. Carnivore species distributed over a wide geographical range generally have highly polymorphic morphological variation. The raccoon dog (Nyctereutes procyonoides) has a longitudinal distribution restricted to East Asia and the northern Indochina Peninsula. Its unique geographical range makes it an appropriate model to examine how morphological differences are influenced by geography. To demonstrate morphological evolution of Russian, Chinese, Korean and Japanese raccoon dogs predicted by geographical differences, we tested the island rule and Bergmann's rule. We compared craniodental variation among populations and examined morphological implications for intraspecific taxonomic status. Insular raccoon dogs possessed substantially smaller body size than those from the mainland. Moreover, different island effects among Japanese islands were demonstrated by markedly larger occipital condyle breath in the Hokkaido population. Larger skull size in Russian and Hokkaido raccoon dogs could be explained by Bergmann's rule. Based on previous chromosomal and molecular studies and results of our morphological analyses, we suggest Japanese raccoon dogs are a distinct species from the mainland N. procyonoides.     

The role of geography and ecology in shaping repeated patterns of morphological and genetic differentiation between European minnows (Phoxinus phoxinus) from the Pyrenees and the Alps

Neutral and selective processes can drive repeated patterns of evolution in different groups of populations experiencing similar ecological gradients. In this paper, we used a combination of nuclear and mitochondrial DNA markers, as well as geometric morphometrics, to investigate repeated patterns of morphological and genetic divergence of European minnows in two mountain ranges: the Pyrenees and the Alps. European minnows (Phoxinus phoxinus) are cyprinid fish inhabiting most freshwater bodies in Europe, including those in different mountain ranges that could act as major geographical barriers to gene flow. We explored patterns of P. phoxinus phenotypic and genetic diversification along a gradient of altitude common to the two mountain ranges, and tested for isolation by distance (IBD), isolation by environment (IBE) and isolation by adaptation (IBA). The results indicated that populations from the Pyrenees and the Alps belong to two well differentiated, reciprocally monophyletic mtDNA lineages. Substantial genetic differentiation due to geographical isolation within and between populations from the Pyrenees and the Alps was also found using rapidly evolving AFLPs markers (isolation by distance or IBD), as well as morphological differences between mountain ranges. Also, morphology varied strongly with elevation and so did genetic differentiation to a lower extent. Despite moderate evidence for IBE and IBA, and therefore of repeated evolution, substantial population heterogeneity was found at the genetic level, suggesting that selection and population specific genetic drift act in concert to affect genetic divergence.     

Speciation and chromosomal evolution of the Baikalian endemic chironomids of the genus Sergentia Kief. (Diptera, Chironomidae): Karyotype divergence and chromosomal polymorphism in the populations of eurybathic species Sergentia flavodentata Tshern. and littoral species Sergentia baicalensis Tshern

Specific karyotype structure and chromosomal polymorphism was investigated in the populations of the Baikalian endemic chironomid species, eurybathic Sergentia flavodentata Tshernovskij, 1949 and littoral Sergentia baicalensis Tshernovskij, 1949. In addition to two fluctuating homozygous inversions in arms A and E, both species were characterized by species-specific karyotype features, namely, nucleolar organizer in the region 1 of chromosome IV in S. flavodentata, and inversion in arm F in S. baicalensis. Moreover, S. baicalensis demonstrated the presence of intraspecific population polymorphism. The populations differing in the presence of secondary overlapping inversion in arm A were found. The highest number of chromosomal rearrangements (7) was detected in S. flavodentata. Most of these rearrangements (six) were found in the population from the underwater thermal spring at a depth of 420 to 430 m (Northern Baikal, Frolikha Bay). In the populations from Middle and Southern Baikal, a rare pericentric inversion in chromosome I was described. In S. baicalensis, in addition to two common heterozygous inversions in arms C and F, disturbance of homologous pairing in different regions of the remaining arms were often detected. Stable chromosomal polymorphism preserved during 13 years in the population of S. flavodentata from the region of hydrothermal venting, serves as an evidence of high genetic plasticity of the species, which favors successful colonization of different Baikal depths and biotopes. Original Russian Text ? V.I. Proviz, 2008, published in Genetika, 2008, Vol. 44, No. 9, pp. 1191–1202.     

Morphological vs. molecular delineation of taxa across montane regions in Europe: the case study of Gammarus balcanicus Schäferna, (Crustacea: Amphipoda)

Mountainous areas are characterized by substantial biodiversity and endemicity due to their complex geological history and habitat fragmentation. Hence, it can be assumed that particularly high species richness can be found in organisms with limited dispersal capabilities that inhabit mountain streams. A number of scientific papers focus on molecular phylogeography or traditional taxonomy of species or species groups inhabiting such habitats. However, there is a lack of studies that integrate morphological and molecular data to identify and delineate cryptic species. For practical reasons, uncovering cryptic diversity is crucial in taxa used in biomonitoring. Distinct species, hard to separate based on morphology only, may have different tolerance ranges towards a variety of factors. Thus, our goal is to combine the two approaches to reveal potential patterns of diversification within a species widely distributed across European mountains: the amphipod crustacean Gammarus balcanicus. The data were obtained from 13 populations spread across the range of the species. Individuals were initially ascribed to G. balcanicus based on conventional fauna key morpho‐anatomical diagnostic features and were further analysed for 23 additional features to explore any putative diversification. Morphometric data were analysed with use of the multiple correspondence analysis and anova . Molecular distances were calculated for 551‐bp‐long COI sequences. Test for isolation by distance was performed for both morphological and molecular data. The morphometric studies showed that some of the analysed features differed significantly between populations, although there was only a weak correlation between the morphological divergence and the between‐population geographical distances. Moreover, high morphological diversity was present within sites. A set of 42 COI haplotypes was identified among the 135 individuals sequenced. No haplotype was shared among populations. The molecular p‐distances within the nine localities presenting more than one haplotype were either almost null (ca. <0.003 for 7 localities) or relatively low (ca. 0.01–0.02 for 2 localities). In opposite, the molecular p‐distances between localities were mostly at a high level (94% of pairwise comparisons being >0.14), similar as between other well‐defined species of the genus Gammarus. Surprisingly, G. balcanicus appears to be polyphyletic based on topology of the neighbour‐joining tree. The level of genetic distance between localities was not correlated with their geographical proximity. Globally, combining spatial patterns of morphological versus molecular divergence indicates a high level of cryptic diversity within a species conventionally defined based upon fauna key morphological features. In this context, the name G. balcanicus should be applied only to the population from locus typicus, while the other populations represent a number of putative distinct species. We may expect that such phenomenon would apply also to other animal taxa with conserved morphology, which are widespread over different mountain ranges in Europe.     

The species composition and distribution of free-living nematodes (Nematoda) in an area of natural oil and gas seeps in Lake Baikal

The species composition and quantitative parameters of free-living nematode fauna have been studied in an area of natural oil seeps off Cape Gorevoi Utes (Middle Baikal). A total of 34 species of 17 genera, 10 families, and 8 orders have been found, 18 of them endemic to Lake Baikal. The nematode fauna of bitumen volcanoes was found to be represented by three specialized species (Monhystera naphthera Gagarin et Naumova, 2010; Eumonhystera abyssalis Gagarin et Naumova, 2010; and Eutobrilus mirandus Gagarin et Naumova, 2011). Areas of the bottom with scattered oil seeps are dominated by eurybathic species endemic to Lake Baikal. The maximal values of nematode population density and maximum proportion of nematodes in the meiozoobenthos were recorded in areas directly adjacent to oil seeps.     

Finding a new form of the grayling Thymallus arcticus (thymallidae) in the basin of lake baikal

A dwarf form of the Arctic grayling Thymallus arcticus, inhabiting a group of small lakes at the origin of the Yakchii stream, which flows into the Verkhnyaya Angara River (basin of Lake Baikal), has been found in the northeastern watershed of the Baikal and Lena basins. The form is similar to upper Lena populations in body coloration and the pattern of dorsal fin. Our comparison by meristic characters indicated that the grayling of the Yakchii Lakes is more similar to the fish from Kutima River (basin of the upper courses of the Lena River) than to the black Baikal grayling T. arcticus baicalensis. The presence in Lake Baikal of a population similar to upper Lena graylings may have three causes: (1) possible drainage from Baikal to the pra-Lena via ancient valleys of the Barguzin and the Upper Angara; (2) appearance of transit zones as a result of glacial or tectonic events during the periods of watershed development between the Baikal and Lena basins; (3) the upper Lena grayling could be an endemic of the Baikal basin which was replaced by graylings penetrating from the Yenisei basin and remained in the form of relic populations in the upper courses of certain tributaries of the northern and northeastern parts of Lake Baikal.     

Morphological divergence among Spanish Common Crossbill populations and adaptations to different pine species

Crossbills (Loxia spp.) provide a classical avian model of ecological specialization on food resources. Previous studies have suggested that morphometric, genetic and vocal diversification among Common Crossbill Loxia curvirostra populations is better explained by ecological distance (use of different conifers) than by geographical distance, indicating that populations have diverged adaptatively. We tested for adaptive divergence in Iberian crossbills using bill and body size measurements of 6082 crossbills from 27 sites, each consisting of a dominant or single pine (Pinus) of four possible species. Crossbills using different pines differed significantly in body size and bill size and shape. There was no correlation between geographical and morphological distance among sampling sites, consistent with the hypothesis that the morphological divergence of Iberian crossbills is shaped by their ecological differences (foraging on alternative conifers) rather than geographical distance. However, for unknown reasons, Common Crossbills foraging on Pinus sylvestris in Iberia have on average much smaller bills than Parrot Crossbills Loxia pytyopsittacus feeding on the same pine species in northern Europe. The extent to which crossbills specialize on Iberian P. sylvestris remains to be established. Specialization on conifers with overlapping geographical distributions may be facilitated by matching habitat choice of crossbills as a function of their local intake rates.     

Regional genetic variability among South African vervet monkey<Emphasis Type="Italic">Chlorocebus aethiops</Emphasis> populations

A South African rehabilitation centre for illegally kept vervet monkeysChlorocebus aethiops Linnaeus, 1758 required an evaluation of the genetic status of vervet monkeys, to determine whether animals from different geographical areas may be mixed. We studied animals originating from three geographical regions and used biochemical genetic and morphological approaches to address this question. The most prominent trend from allozyme data was derived from the locusPrt-2 (an unspecified serum protein), where each of the three populations could be characterised by the absence or presence of unique alleles. Nevertheless, statistical coefficients indicated little genetic divergence, with genetic distance values of 0.001–0.003 and an overall fixation index value of 0.046. Average heterozygosity did not differ appreciably among populations (2.5–3.3%). The morphological study identified suitable traits, free from the influence of growth allometry, that can be used for inter-population comparisons. No significant morphological differences between con-specific populations were however found. It is concluded that vervet monkeys from the species’ wider distribution range is relatively monotypic, but that monkeys from different geographical areas should not be unduly mixed, pending the results of finer grained molecular studies.     

Special traits of growth and feeding of the stone sculpin <Emphasis Type="Italic">Paracottus knerii</Emphasis> (Cottidae) in water bodies of different types

Growth and feeding of the stone sculpin Paracottus knerii (Cottidae) is investigated in Water Bodies of Different Types: Baikal (the littoral of its southern part), the Angara River, and the mountainous Sobolinoe Lake (the southern Baikal basin). The highest growth rate is characteristic of the South-Baikalian population, followed by the Angara population. The stone sculpin from Sobolinoe Lake is slow-growing. The growth rate of specimens of the latter is by 1.5–2 times lower than in the two aforementioned populations due to inferior feeding conditions of this water body. The size-age changes in the food spectrum of all investigated populations are consecutive change of all principal components and increase in size of the consumed prey     

A molecular phylogenetic and phylogeographic study of two forms of Calliptamus barbarus (Costa 1836) (Orthoptera: Acrididae,Calliptaminae) from two regions of Algeria

Calliptamus barbarus (Orthoptera: Acrididae) is the most polymorphic species within the genus Calliptamus. It shows a morphological polymorphism (three hind femoral spots, or only one hind femoral spot). Several studies have been made in order to distinguish the two forms: morphometry, number of ovarioles, sound production, protein and enzyme system. The aim of our work is to assess whether the two forms can be considered as different taxa and to perform a molecular phylogenetic study of two populations of C. barbarus collected from two different Algerian localities. No clear genetic differentiation was found between the samples with different morphologies. Additionally, the samples from Algeria do not form a monophyletic sister clade compared to the one formed by the sequences from GenBank from other geographical regions. Despite the morphological differences shown between the two populations, our molecular study indicates that there are no differences at a molecular level using the two mitochondrial genes COI and 16S.     

Geographical variation in the cranial and external characters of the little tube-nosed bat,Murina silvatica in the Japanese archipelago

The distribution ofMurina silvatica (Yoshiyuki, 1983) in the Japanese archipelago extends over about 2000 km from north to south. Specimens were obtained from populations in Hokkaido and Yakushima, which are at the northern and southern ends of the range, and from two intermediate locations in Honshu. Measurements of cranial and external morphology were examined for evidence of geographical variation. The results of both multivariate analysis of variance and cluster analysis showed that there was no distinct cline in skull morphology among the Hokkaido, Tohoku and Chubu populations. However, the results of multivariate analysis of variance showed that all measures were significantly greater for the Yakushima population than for the others. Similarly, in a dendrogram of cluster analysis, the Yakushima population formed a cluster that was distinct from the other populations. However, as the difference between the Yakushima population and the other populations was less than the variation found within the Hokkaido, Tohoku and Chubu populations, morphological divergence of the Yakushima population was attributable to intraspecific variation. The island of Yakushima is the most isolated of the four locations and the morphological divergence of this population may be associated with its relative geographic isolation.     

Phenetic Analysis of Aboriginal and Introduced Populations of Sable (Martes zibellina) in Russia

Using standard and multivariate statistic methods, an epigenetic character—foramina in fossa condyloidei inferior(FFCI)—was studied in sable populations. This character was shown to be most frequent in southeastern populations of the species (Primorye and the Baikal region) while its distribution in the remaining part of the range was polyclinal. The expression of FFCI was directly associated with coat color and longitude, and inversely associated with skull size. This trend was broken by some western populations formed in the 1950s by introduction, which exhibited stable morphological differences with adjacent aboriginal sable populations. Most populations of the species exhibit differences in the manifestation of the studied characters. Frequency of the FFCI manifestation can be used as an additional population characteristic, an associative diagnostic character that shows high discriminating capability in detecting phenogenetic relationships of intraspecific groups.     

Song complexity is associated with habitat quality in an upland passerine

Understanding the influence of intrinsic (genetic and morphological) and extrinsic (geographical, environmental and social) factors on the performance and spatial differentiation of sexual signals, such as bird song, can help identify behavioural indicators of individual quality, habitat degradation and social environment. We used the Iberian Bluethroat Luscinia svecica azuricollis, a migratory bird that breeds in fragmented landscapes dominated by shrublands, as a case study to: (1) assess how a set of acoustic indicators of song performance are driven by intrinsic and extrinsic factors; and (2) contrast deterministic (adaptations to the environmental context and morphological constraints) vs. stochastic (differentiation by geographical isolation) explanations for song differentiation patterns. We explored acoustic indicators of song performance (spectral, temporal and song complexity) in relation to parameters related to genetic structure, body size, habitat type, habitat quality (assessed through a spatially explicit modelling approach) and social context (population abundance and songbird community composition). Then, we explored the contribution of genetic, geographical and environmental dissimilarity to song diversification across space. Our results highlight an association of song spectral variables with genetic structure and a significant connection between song complexity and duration with habitat quality. We found no relationship between social features and acoustic variables, or between song differentiation and genetic or geographical distances. There was, however, a correlation between song differentiation and environmental dissimilarity. We recommend the consideration of song complexity as an indicator of habitat quality.