Cytogenetic differentiation of natural populations of <Emphasis Type="Italic">Chironomus obtusidens</Emphasis> (Diptera,Chironomidae)

The cytogenetic structure of Siberian and European populations of Chironomus obtusidens Goetgh. was studied. The clear differentiation of these populations was revealed using quantitative analysis of sets and frequencies of inversion banding sequences. The Siberian populations were found to have six new inversion sequences, and the alternative obtG2 sequence dominated instead of the main obtG1 sequence typical of European populations. In general, chromosomal polymorphism was 3–4 times higher in the Siberian populations. Therefore, the cytogenetic distances (Dcg) between the Siberian and European populations were higher (Dcg 0.059–0.097) than Dcg between the populations in each of the geographic zones investigated (Dcg 0–0.015). The NJ phylogenetic tree constructed for the chromosomal evolution of the genus Chironomus demonstrates that Chironomus obtusidens is the most divergent member among Palaearctic species of the obtusidens group. In addition, it is closer related to the Nearctic Ch. decorus group.     

Differentiation of chromosome banding sequence pools and genomic dna in Holarctic natural populations of <Emphasis Type="Italic">Chironomus entis</Emphasis> Shobanov (Diptera,Chironomidae)

Polymorphism and differentiation of the chromosome banding sequence pools and genomic DNA were studied in three natural populations of Chironomus entis from Europe and North America. These populations showed a moderate level of chromosomal polymorphism and high RAPD polymorphism of genomic DNA. The Palearctic and Nearctic populations of this species did not differ significantly in the levels of chromosome and genomic DNA polymorphism. Estimation of the cytogenetic (GD_cg) and genetic (GD_DNA) distances between these C. entis populations showed that their chromosome banding sequence pools and cytogenetic structures are differentiated to a greater extent than genomic DNA. The values of cytogenetic and genetic distances between the Palearctic and Nearctic populations of C. entis are higher than the values of the corresponding distances between the Nearctic populations, but they do not reach the level of divergence between species.     

Intercontinental karyotypic differentiation of Chironomus entis Shobanov, a Holarctic member of the C. plumosus group (Diptera, Chironomidae).

Analysis of banding sequences of polytene chromosomes in Palearctic (Russian) and Nearctic (North American) Chironomus entis shows strong karyotype divergence between populations on the two continents. Four out of seven chromosomal arms in the North American C. entis karyotype are characterized by sequences found only in the Nearctic. In total, 44 banding sequences are now known for this species across the Holarctic, including 22 exclusively Palearctic, 6 Holarctic, and 16 exclusively Nearctic sequences. The degree of cytogenetic differentiation between Palearctic and Nearctic C. entis populations is an order of magnitude greater than differentiation among populations within either continent, but is only one third as great as the cytogenetic distance between the sibling species C. entis and C. plumosus. C. entis is the only sibling species of C. plumosus uncovered during cytological identification of Chironomus species from more than 50 North American lakes, indicating that the plumosus sibling-species group is much smaller in the Nearctic than in the Palearctic, where a dozen sibling species are known. Cytogenetic distance values calculated between Nearctic and Palearctic representatives of both C. entis and its sibling species C. plumosus are similar, but result from different patterns of karyotype divergence. New World C. entis is distinguished from Old World populations by the 16 uniquely Nearctic sequences, four of which occur in the homozygous state. In contrast, North American C. plumosus has fewer uniquely Nearctic sequences, and only one that occurs as a homozygote. However, four chromosomal arms in C. plumosus that are polymorphic in the Palearctic show fixation, or near fixation, of Holarctic sequences in the Nearctic C. plumosus karyotype. Thus, both the fixation of Holarctic sequences, and the occurrence or fixation of distinctly Nearctic sequences, contribute significantly to karyotype divergence. Patterns of karyotype divergence in Palearctic and Nearctic populations of different Holarctic chironomid species are discussed relative to intercontinental cytogenetic differentiation in other dipterans.     

Geographic differentiation of genomic DNA of <Emphasis Type="Italic">Chironomus plumosus</Emphasis> (Diptera,Chironomidae) in natural holarctic populations

Using RAPD markers, polymorphism and differentiation of genomic DNA was examined in seven natural populations of Chironomus plumosus from Europe, Siberia, and North America. All these populations showed high polymorphism of genomic DNA. The Palearctic and Nearctic populations of this species were not statistically significantly different in the genomic DNA polymorphism level. The genetic distance (GD), which characterizes the extent of intraspecific differentiation of population genetic structure, was determined among the natural populations of C. plumosus. The genetic distance was on average 0.245. It was demonstrated that genetic structures of the Palearctic and Nearctic populations of C. plumosus was differentiated to a higher extent than in Palearctic. However, the genetic distances between the populations from different zoogeographical zones (0.313) did not exceed the level characteristic of the among-population differences, which do not disturb the species genetic integrity.     

Geographic variability of the polytene chromosome banding sequence of non-biting midge <Emphasis Type="Italic">Chironomus pseudothummi</Emphasis> Str. (Diptera,Chironomidae)

The karyotypes and chromosomal polymorphism of Chironomus pseudothummi were investigated in different parts of its areal. It was established that chromosomal variability in the natural populations of this species was represented mainly by the inversion polymorphism of arm G. Only rare and unique inversions were found to be heterozygous in arms C, D, and E. In total, 14 banding sequences of polytene chromosomes form the banding sequence pool of C. pseudothummi. Geographic differences in the distribution of chromosomal banding sequences throughout the areal were established. The presence of banding sequences pstG1 and pstG2 is characteristic of European populations. The banding sequence pstG1 disappeared completely with a simultaneous increase in the frequency of pstG2 and with the appearance of a new inversion banding sequence pstG3 in Siberian populations. Differences in the set of rare and unique inversions in arms C, D, and E between the West-European and West-Siberian populations have been revealed.     

The geographic varyability of the polytene chromosome banding sequence of non-biting midje Chironomus pseudothummi str. (Diptera, Chironomidae)

The karyotypes and chromosomal polymorphism of Chironomus pseudothummi were investigated in different parts of its range. It was established that chromosomal variability in the natural populations of this species was represented mainly by the inversion polymorphism of arm G. Only rare and unique inversions were found as heterozygous in arms C, D, and E. In total, the 14 banding sequences of polytene chromosomes form the banding sequences pool of Ch. pseudothummi. Geographic differences in distribution of chromosomal banding sequences throughout the range were established. The presence of banding sequences pstG1 and pstG2 is characteristic of European populations. Banding sequence pstG1 was completely vanished with simultaneous increase in frequency of pstG2 and appearance of new inversion banding sequence pstG3 in Siberian populations. The differences in the set of the rare and unique inversions in arms C, D, and E between west-European and west-Siberian populations were revealed.     

The chromosomal polymorphism and divergence of populations in Chironomus nuditarsis Str. (Diptera, Chironomidae)

The karyotype structure and chromosomal polymorphisms were investigated in 6 natural and 2 laboratory populations of Chironomus nuditarsis from Europe and Asia. The pool of rearranged polytene chromosome banding sequences of this species was determined that includes 16 inversion banding sequences and sequences with giant DNA-knobs (ndtG1k, ndtG2k). Obvious differences were demonstrated in the level of chromosomal polymorphism between European and Asian (Siberian) populations: the former were highly polymorphic, while the latter were practically monomorphic. It was suggested to consider the Siberian populations as marginal one. Cytogenetic distances between populations of C. nuditarsis as well between C. nuditarsis and the related species C. plumosus were estimated. The data obtained show that chromosomal rearrangements play a very important role in cytogenetic divergence of populations.     

Chromosomal polymorphism in two species of Hypancistrus (Siluriformes: Loricariidae): an integrative approach for understanding their biodiversity

Structural chromosome changes are widely described in different vertebrate groups and generate genetic, phenotypic and behavioral diversity. During the evolution of loricariids, several rearrangements (fissions, fusions, inversions) seem to have occurred. Hypancistrus, tribe Ancistrini, are highly demanded for fishkeeping around the world. In this tribe, the diploid chromosome number 2n = 52 is considered a synapomorphy, and paracentric-type inversions appear to be involved in the chromosomal evolution of the tribe. The present study investigated the karyotypes of H. zebra and H. cf. debilittera using cytogenetic, classical and molecular tools, as well as DNA barcoding. Data reveal that, although diploid number in both species corroborates the proposed synapomorphy for the tribe, there is a complex karyotype dynamics, reflected in the intense chromosomal polymorphism, resulting from rearrangements involving ribosomal regions (5S and 18S rDNA), which are suggested to be paracentric inversions. Besides, DNA barcode confirms reciprocal monophyletism between the species, validating the existence of two species, only. This scenario, coupled with genomic instability caused by exogenous sequences such as Rex-3 retrotransposons and the species’ sedentary lifestyle, which helps the fast polymorphism fixation, may reflect different phenotypic color patterns in natural populations, as observed in H. cf. debilittera.     

A genealogical view of chromosomal evolution and species delimitation in the Drosophila virilis species subgroup

Chromosomal arrangement was a historically important character used for defining taxonomic boundaries. The Drosophila virilis species group exhibits a series of chromosomal rearrangements, and the resulting differences among karyotypes were primary characters originally used to define taxa within the group. However, some chromosomally divergent forms have not been sufficiently resolved in phylogenetic reconstructions of DNA sequences from several nuclear genes. Sequences of mitochondrial regions have the potential for finer-scale resolution of closely related taxa; therefore, sequences of two mitochondrial genes were used to examine phylogenetic relationships within the chromosomally variable virilis subgroup. Sequences were obtained from multiple strains of the Palearctic species, D. virilis and D. lummei, and the Nearctic species, D. novamexicana and two chromosomal forms of D. americana. Analyses support the recent emergence of the different chromosomal forms in North America. However, none of these chromosomally divergent forms exhibit reciprocal monophyly of their mtDNA sequences, which is the requirement for attaining genealogical species status.     

Genomic Organization of Repetitive DNA Elements and Its Implications for the Chromosomal Evolution of Channid Fishes (Actinopterygii,Perciformes)

Channid fishes, commonly referred to as “snakeheads”, are currently very important in Asian fishery and aquaculture due to the substantial decline in natural populations because of overexploitation. A large degree of chromosomal variation has been found in this family, mainly through the use of conventional cytogenetic investigations. In this study, we analyzed the karyotype structure and the distribution of 7 repetitive DNA sequences in several Channa species from different Thailand river basins. The aim of this study was to investigate the chromosomal differentiation among species and populations to improve upon the knowledge of its biodiversity and evolutionary history. Rearrangements, such as pericentric inversions, fusions and polyploidization, appear to be important events during the karyotypic evolution of this genus, resulting in the chromosomal diversity observed among the distinct species and even among populations of the same species. In addition, such variability is also increased by the genomic dynamism of repetitive elements, particularly by the differential distribution and accumulation of rDNA sequences on chromosomes. This marked diversity is likely linked to the lifestyle of the snakehead fishes and their population fragmentation, as already identified for other fish species. The karyotypic features highlight the biodiversity of the channid fishes and justify a taxonomic revision of the genus Channa, as well as of the Channidae family as a whole, as some nominal species may actually constitute species complexes.     

DNA barcoding as a heuristic tool for classifying undescribed Nearctic Myrmica ants (Hymenoptera: Formicidae)

The Palearctic species of the ant genus Myrmica are well studied. In contrast, the taxonomy of the Nearctic species is outdated, making identification impossible. We collected Myrmica samples in the Holarctic and investigated their diversity using mtDNA data. We analysed a barcode sequence of the Cytochrome Oxidase I gene for 57 Palearctic and 293 Nearctic Myrmica samples. We used sequences of known Palearctic species to search for Myrmica barcode patterns. All but one Palearctic species groups were recovered. The Nearctic diversity was much higher than known. We retrieved the punctiventris, crassirugis and incompleta groups, and established nine additional tentative species groups. Genetic distance analysis revealed a large overlap of intra- and inter-specific distances in Palearctic species and species groups. We could not find a variation gap to separate Nearctic sequences into species with COI data only. Variation in scape morphology divided two genetic groups further. Scape morphology correlated with most molecular groups, except three specimens. Our results illustrate that barcoding, using only a limited amount of genetic information, cannot serve as a universal proxy for taxonomy and species demarcation. It should be considered a first step in understanding the taxonomic diversity of an unknown group of organisms.     

Parallel evolution in ecological and reproductive traits to produce cryptic damselfly species across the holarctic

The damselfly genus Enallagma originated in the Nearctic, and two Nearctic lineages recently underwent radiations partly associated with multiple independent habitat shifts from lakes dominated by fish predators into lakes dominated by dragonfly predators. A previous molecular study of four Palearctic morphospecies and all representative Nearctic species identified the presence of two cryptic species sets, with each set having Palearctic and Nearctic representatives. However, the cryptic species within each set are not sibling species. Here, we present quantitative data on ecologically important larval morphologies and behaviors involved in predator avoidance and on adult male morphological structures involved in mate recognition to quantify the phenotypic relationships among these cryptic species sets. For the adult stage, our data indicate strong parallel evolution of the structures involved in specific mate recognition-the male cerci. For the larval stage, morphometric analyses show that the Palearctic species evolved a nearly identical morphology to the sibling-clade members in the Nearctic that live in waters where dragonflies are the top predators. This implicates the importance of dragonfly predation in the history of the Palearctic clade. Behavioral analyses suggest population differentiation in response to the actual predator environment in the Palearctic clade, consistent with the species differentiation seen in the Nearctic. Our results suggest parallel evolution of adult traits that influence specific mate choice and larval traits that influence ecological performance underlie the striking similarity of Enallagma species across continents. This concurrent parallel evolution in both stages of a complex life cycle, especially when both stages do not share the same selective environment, may be a very unusual mechanism generating cryptic species.     

Karyofund of Chironomus plumosus (Diptera, Chronomidae) in the Palearctic region

Quantitative and qualitative analyses of chromosomal polymorphism in 38 Palearctic populations of Chironomus plumosus were made. It was shown that most of the populations studied were highly polymorphic: in average 63.2 +/- 4.0% of larvae were heterozygous for inversions with 0.95 + 0.08 heterozygous inversion per larvae. Polymorphism on the size of centromeric heterochromatin and the presence of B-chromosomes were observed in many populations studied. The karyofund of Ch. plumosus in Palearctic was estimated. In total 35 banding sequences were found in Palearctic Ch. plumosus. Fifteen banding seguences have been described for the first time. On mapping the used banding sequences, we employed the conventions of Keyl (Keyl, 1962; Devai et al., 1989) and Maximova (Maximova, 1976; Shobanov, 1994a) for arms A, C, D, E and F, and the conventions of Maximova for arms B and G.     

Karyotypic features of Chironomus entis and Chironomus borokensis (Diptera, Chironomidae) from Lake Kotokel (Lake Baikal Basin)

Karyotypes and chromosomal polymorphism of sympatric sibling species Chironomus entis and Chironomus borokensis from Lake Kotokel were studied. Five inversion banding sequences were found in C. entis, including p’entA16, p’entB9, and p’entF6 that are new for the species; in C. borokensis, four inversion banding sequences were revealed, borA4 being new for the species. The level of chromosome polymorphism in C. entis and C. borokensis can be considered similar: 54.2 and 53.8% heterozygosity, 0.67 and 0.73 inversions per individual, respectively. The new data are compared with those on C. entis population from Lake Kotokel obtained in 1981 and those of C. borokensis from Lake Dukhovoe (Lake Baikal basin) obtained in 1983.     

Inversion polymorphism of <Emphasis Type="Italic">Glyptotendipes glaucus</Emphasis> Mg. (Diptera: Chironomidae) from the reservoirs of Kaliningrad

The karyotypes of larvae Glyptotendipes glaucus Mg. 1804 from three reservoirs of Kaliningrad city (Pen’kovoe and Karasevka lakes and Chistyi pond) were studied. The levels of the natural inversion polymorphism for the three populations were determined. Ten new inversion sequences (gla B4, gla B5, gla B6, gla D5, gla E4, gla E5, gla E6, gla E7, gla F6, and gla F7) were detected in the species studied. Inversion including the centromeric region in IIIEF (gla E6 + gla F7) was pericentric. Several cases of gla B5 and gla B6 combination with gla B2 resulting in genotypic combination gla B2.5 and gla B2.6 were found. The dependence of the number of inversion sequences and genotype combinations from the content of heavy metal ions in sediments of the reservoirs was revealed. All populations were characterized by the prevalence of inversion sequence gla B2 in chromosome I.     

Divergence of the polytene chromosome banding sequences as a reflection of evolutionary rearrangements of the genome linear structure

Banding sequences of five chromosomal arms (A, C, D, E, and F), accounting for about 70% of the total genome size in 63 Chironomus species, were used as markers to analyze divergence patterns of the linear genome structure during the evolution. The number of chromosomal breakpoints between the pairs of banding sequences compared served as a measure of divergence. It was demonstrated that the greater the divergence between the species compared, the higher the number of chromosomal breakpoints and the smaller the size of the conserved chromosomal segments. A banding sequences comparison in sibling species demonstrated a lower number of chromosomal breakpoints; the breakpoint number was maximum in a comparison of the banding sequences in the subgenera Chironomus and Camptochironomus. The use of the number of chromosomal breakpoints as a genome divergence measure provided establishment of phylogenetic relationships between 63 Chironomus species and discrimination of sibling species groups and cytocomplexes on a phylogenetic tree.Translated from Genetika, Vol. 41, No. 2, 2005, pp. 187–195.Original Russian Text Copyright © 2005 by Gunderina, Kiknadze, Istomina, Gusev, Miroshnichenko.     

Molecular Organization of the X Chromosome in Different Species of the Obscura Group of Drosophila

Nine single copy regions located on the X chromosome have been mapped by in situ hybridization in six species of the obscura group of Drosophila. Three Palearctic species, D. subobscura, D. madeirensis and D. guanche, and three Nearctic species, D. pseudoobscura, D. persimilis and D. miranda, have been studied. Eight of the regions include known genes from D. melanogaster (Pgd, zeste, white, cut, vermilion, RNA polymerase II 215, forked and suppressor of forked) and the ninth region (lambda DsubF6) has not yet been characterized. In all six species, as in D. melanogaster, all probes hybridize to a single site. Established chromosomal arm homologies of Muller's element A are only partly supported by present results since two of the probes (Pgd and zeste) hybridize at the proximal end of the XR chromosomal arm in the three Nearctic species. In addition to the centric fusion of Muller's A (= XL) and D (= XR) elements, the metacentric X chromosome of the Nearctic species requires a pericentric inversion to account for this result. Previously proposed homologies of particular chromosomal regions of the A (= X) chromosome in the three species of the D. subobscura cluster and of the XL chromosomal arm in the three species of the D. pseudoobscura cluster are discussed in light of the present results. Location of the studied markers has changed drastically not only since the divergence between the melanogaster and obscura groups but also since the Palearctic and Nearctic species of the obscura group diverged.     

Evolutionary and climatic factors affecting tooth size in the red foxVulpes vulpes in the Holarctic

Research into the geographical pattern of tooth size in the red fox,Vulpes vulpes (Linnaeus, 1758) in the Holarctic was conducted on a sample of 3806 skulls belonging to 41 fox populations. The Nearctic was represented by 948 specimens (249 females, 359 males, 340 specimens of unknown sex) belonging to 13 populations, whereas the Palearctic was represented by 2858 red foxes (1034 females, 1256 males, 568 specimens of unknown sex) from 32 populations. In the Nearctic, the largest foxes live on Kodiak Island (V. v. harrimani) and the Kenai Peninsula (V. v. kenaiensis), while the smallest ones live in California (V. v. necator) and Georgia (V. v. fulvus). In the Palearctic, the largest foxes come from the Far East (V. v. jakutensis, V. v. beringiana, V. v. tobolica), while the smallest are from the southern borders of the Eurasian range (V. v. pusilla, V. v. barbara, V. v. arabica). In both the Palearctic and Nearctic, tooth size in the fox varies depending on the geo-climatic factors. The fox’s tooth size confirms the general basis of Bergmann’s rule. In the Palearctic, specimens with larger teeth occur in cooler habitats with greater seasonality. These are first and foremost Northern and Far Eastern populations. In the Nearctic, tooth size in red foxes depends on the temperature and humidity of their habitat. Competition within the species and between species has important impact on the variation and dimorphism of tooth size in the red fox. Both in the Nearctic and Palearctic, red foxes from regions of sympatric co-occurrence with other closely relatedVulpes species, are more sexually dimorphic in terms of tooth size than red foxes from allopatric regions. Analysis of morphological distance on the basis of the size of dental characteristics shows, that in the Palearctic, the foxes from India (V. v. pusilla), while in the Nearctic, the population from Kodiak Island (V. v. harrimani) are most distant from the remaining populations. Geographic barriers such as the Bering Strait, Parry Channel, Mackenzie River, Kolyma and Omolon River systems have had a critical impact on red fox evolution. The most likely place for the evolution and diversification of the phyletic lineVulpes vulpes seems to be the Middle East region.     

Patterns of inversion polymorphism in three species of the Drosophila melanogaster species group

In Drosophila, chromosomal polymorphism due to paracentric inversions is very common and constitutes an adaptive character. The degree of chromosomal variability varies in different species and also in different populations of the same species. Chromosomal polymorphism in Indian natural populations of three species, D. melaonogaster, D. ananassae and D. bipectinata which belong to the melanogaster species group has been studied and the quantitative data on frequency of inversions have been reported. Behaviour of chromosome inversions has also been studied in laboratory conditions. The present review summarises the work done on inversion polymorphism in Indian populations of three species which clearly demonstrates that these three species vary in their patterns of inversion polymorphism and have evolved different mechanisms for adjustment to their environments although they belong to the same species group.     

Evolution of Hawaiian Drosophilidae. II. Patterns and Rates of Chromosome Evolution in an Antopocerus Phylogeny

The phylogenetic relationships of seven species of the genus Antopocerus (Family Drosophilidae) have been determined by means of a study of the metaphase configurations and polytene chromosomes. Based on biogeographical, behavioral and cytogenetic information, A. longiseta from Molokai is tentatively identified as the primitive species of the genus. The metaphase karyotypes of all Antopocerus species are either five pairs of rod chromosomes and a pair of dots (5R1D), or six rods (6R). Heterochromatin additions converted the dots to rods. Chromosome breakpoints for inversions also are clustered at heterochromatic loci. The chromosome segments between heterochromatic loci may represent sets of functionally related loci, evolving as a unit. The rate of chromosomal inversion substitution is estimated in the origin of the taxon (probably a subgenus of Drosophila rather than a separate genus). It averages no greater than one substitution per 1,000 years, or one per 5,000 generations. The average genetic death rate per generation of one individual per hundred is required to achieve this substitution rate. The rate of inversion substitution during radiation of this taxon may be only 4.4 x 10^-3 times as fast as that present in forming the taxon. Alternatively, radiation may have required only 250,000 years if rates of substitution are the same as in the origination of the taxon. Average rates of substitution reflect genetic accidents, selection pressures and rates of adaptation to new niches, as well as the rate of encountering new niches. Rate of adaptation probably is much greater in this instance than rate of encountering new niches. Therefore, the average rate of evolution reflects more nearly biogeographic and ecological factors than genetic factors.