The phylogeographical and management implications of genetic population structure in the imperiled snuffbox mussel, Epioblasma triquetra (Bivalvia: Unionidae)

Seven populations of the imperiled snuffbox mussel, Epioblasma triquetra , were sampled from across the central basin of North America. Samples were genotyped using 15 microsatellite DNA loci, and maternal history was inferred using mitochondrial DNA (mtDNA) cytochrome  c oxidase subunit-I (COI) sequences. Populations in the Clinch and St Francis rivers were quite distinct in their mtDNA. The population in the St Francis River had a unique, fixed haplotype. Among a suite of haplotypes, the population in the Clinch River had two unique haplotypes of common ancestry. The other populations were dominated by a common haplotype, which also occurred in the Clinch River population. Analysis of DNA microsatellites revealed much greater divergences and showed significant genetic structure between populations in the formerly glaciated regions. Divergence has occurred between the populations, as evidenced by moderate to high fixation indices ( F _ST and R _ST values) and nearly perfect assignment tests. These results indicate the occurrence of three glacial refugia for E. triquetra : the Tennessee River, rivers south of the Ozark Crest, and the lower Ohio River drainage near the confluence with the Mississippi. Populations in the lower Ohio River were likely to be responsible for the postglacial reinvasion into formerly glaciated regions, and into the upper Tennessee River drainage. The population of the St Francis River may constitute a distinct taxonomic entity. Conservation efforts, if necessary for this imperiled species, should not mix populations.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 371–384.     

Discordant patterns of morphological and molecular change in broadtail madtoms (genus Noturus)

Two morphologically distinct forms of an undescribed madtom catfish (Noturus sp.) occur in the rivers and lakes of southeastern USA. 'Lake' broadtail madtoms are endemic to Lake Waccamaw and are probably related to nearby 'river' broadtail populations. To investigate phylogenetic relationships, we surveyed mitochondrial DNA (mtDNA) sequence variation in 'lake' and 'river' broadtails and other members of the genus Noturus. Mitochondrial rDNA data suggest a sister group relationship between broadtail madtoms and N. insignis, not N. leptacanthus as posited previously. Population-level analyses using additional mtDNA characters (NADH dehydrogenase subunit 2 (ND2) and cytochrome b (Cytb)) identified two highly divergent genetic lineages within broadtail madtoms that do not correspond to the morphological designations 'river' and 'lake'.     

Intraspecific phylogeography of the slender madtom: the complex evolutionary history of the Central Highlands of the United States

A number of different biogeographical studies of the Central Highlands of the United States have yielded conflicting area cladograms. We estimate the mtDNA phylogeny of populations of the slender madtom, Noturus exilis, a clear-water stream catfish. The goal is to compare population relationships to those reported in previous studies that used upland, stream-dwelling vertebrates. A region of the NADH dehydrogenase subunit 4 gene, along with adjacent tRNAs, was sequenced for population samples from 21 different Central Highlands rivers. Sequence difference among 39 haplotypes ranged from 0.1% to 4.8%. Most haplotypes were restricted to specific rivers and mapped well onto geography. Slender madtoms from different drainages contained mostly monophyletic groups of haplotypes genetically divergent from haplotypes found in other drainages, although a few haplotypes were found in well-separated drainages. The area cladogram for the slender madtom was not similar to any of the other cladograms for other species and species groups from the area. We discuss a variety of methodological and biological reasons for the discordance, and suggest that some of the discrepancies may be resolved by the sequencing of multiple genes per species. We recommend that more, and more extensive, intraspecific phylogeography studies should be conducted for species living in the Central Highlands rivers.     

Morphological and molecular variation in the least madtom Noturus hildebrandi (Siluriformes: Ictaluridae), a Mississippi Embayment endemic: evidence for a cryptic lineage in the Hatchie River

Sheared principal component analysis of 40 morphometric characteristics measured for 146 individuals and relative frequencies of pigmentation patterns scored for 980 individuals of the least madtom Noturus hildebrandi, a diminutive catfish endemic to eastern lowland drainages of the Mississippi Embayment region of North America, suggested a clinal pattern of morphological variation extending across the range from north to south. DNA sequence data representing 90 individuals from the mitochondrial gene cytochrome b (cytb) analysed using Bayesian phylogenetic methods recovered four major haplotype clades, suggestive of a high degree of isolation by drainage. Individual gene trees of cytb and four additional nuclear loci as well as trees based on concatenated datasets of these genes consistently recovered a cryptic lineage of individuals from the Hatchie River drainage that is morphologically indistinguishable from surrounding populations. Gene‐tree analyses failed to recover a monophyletic N. hildebrandi with respect to Noturus baileyi. A coalescence‐based species tree analysis, however, did recover N. hildebrandi monophyly with high support, suggesting that relationships reflected in individual gene trees and concatenated datasets are in part artefacts of incomplete lineage sorting or an ancient introgressive event. Results are consistent with the hypothesis of an ancient connection between the Hatchie and Tennessee River systems. Current subspecific designations are of limited utility as they reflect morphological variation and are not entirely consistent with phylogeny. Discrepancies between the pattern of variation observed in the morphological and molecular data may be explained by recent local adaptation to individual stream conditions that masks deeper evolutionary divergences.     

Phylogeography and post-glacial colonization patterns of the rainbow darter, Etheostoma caeruleum (Teleostei: Percidae)

Aim To examine the effects of historical climate change and drainage isolation on the distribution of mitochondrial DNA cytochrome b genetic variation within the rainbow darter, Etheostoma caeruleum (Percidae: Etheostomatinae). Location Eastern North American streams including tributaries to the Mississippi River, Great Lakes, Potomac River and Hudson Bay drainages. Methods Parsimony analyses, Bayesian analyses and haplotype networks of mitochondrial DNA sequences. Results Four major clades were recovered from sampled populations of E. caeruleum. Three of four clades are distributed in the western portion of the species’ range (primarily west of the Mississippi River). Samples from this region do not form a monophyletic group, and sequences often vary greatly between samples from adjacent stream systems (up to 7.2% divergence). A basal clade includes samples from the White River system in the Ozark Highlands. The northern Ozarks–upper Midwest clade includes samples from Missouri River tributaries and the upper Midwest (Hudson Bay, upper Mississippi River, and western Lake Michigan drainage). The eastern clade is composed of individuals from the Ohio River, Great Lakes and Potomac River. The Mississippi River corridor clade includes samples from middle and lower Mississippi River tributaries. Main conclusions The four major clades of E. caeruleum are deep allopatric lineages with well‐defined boundaries and have additional phylogeographical structure within each clade. The Ozark Highlands have the greatest levels of diversity relative to distributional area, with marked cytochrome b subdivisions between adjacent stream systems. Samples from previously glaciated areas do not have a subset of the cytochrome b diversity found in unglaciated areas, but four separate source areas are identified based on phylogenetic analyses. Dispersal into previously glaciated areas followed several known glacial outlets and, based on sequence divergence between populations, may have occurred during different glacial or interglacial stages. The disjunct distribution and cytochrome b pattern of E. caeruleum in the Mississippi River corridor clade is consistent with late Pleistocene and Recent changes in the course and characteristics of the middle and lower Mississippi River. Phylogeographical boundaries between clades of E. caeruleum correspond to independent sources of biogeographical information and provide insight into historical stream drainage relationships, post‐glacial colonization and drainage isolation patterns.     

Molecules,morphology, missing data and the phylogenetic position of a recently extinct madtom catfish (Actinopterygii: Ictaluridae)

Taxa missing large amounts of data pose challenges that may hinder the recovery of a well‐resolved, accurate phylogeny and leave questions surrounding their phylogenetic position. Systematists commonly have to contend with one or two species in a group for which there is little or no material available suitable for recovering molecular data. It is unclear whether these taxa can be better placed using analyses based on morphological data only, or should be included in broader analyses based on both morphological and molecular data. The extinct madtom catfish Noturus trautmani is known from few specimens for which molecular data are unavailable. We included this taxon in parsimony and Bayesian analyses of relationships of madtom catfishes based on a combination of morphological and molecular data. Results indicate that using a combination of morphological and molecular data does a better job at providing a phylogenetic placement for N. trautmani than morphology alone, even though it is missing all of its molecular characters. We provide a novel hypothesis of relationships among Noturus species and recommendations for classification within the group. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 60–75.     

Molecular phylogeny of north mediterranean freshwater barbs (genus Barbus: cyprinidae) inferred from cytochrome b sequences: biogeographic and systematic implications

We investigated phylogenetic relationships among north Mediterranean species of the genus Barbus using sequences of the cytochrome b gene. Our results indicate that the species belong to two major clades that are consistent with those previously defined from morphological features. The first clade includes species ranging from France to the Black Sea. In this clade, there is a well-supported monophyletic group of large-sized fluvio-lacustrine barbs; however, the monophyly of the small-sized rheophilic species is not clear. The second clade comprises species found in Spain, Greece, and Asia Minor and probably represents the oldest group present in the north Mediterranean rivers. In general, there is good concordance between geography and phylogenetic relationships. These results are compared to those from previous morphological- and allozyme-based studies and demonstrate widespread discordance and polyphyly in the traditional taxonomy of the genus Barbus. This study is one of the first reporting the phylogenetic and biogeographic relationships of a genus that is widely distributed in European rivers and contains species that are a major component of the European ichthyofauna.     

The phylogenetic relationships among Noturus catfishes (Siluriformes: Ictaluridae) as inferred from mitochondrial gene cytochrome b and nuclear recombination activating gene 2

Madtom catfishes of the genus Noturus are a well-known component of the North American ichthyofauna. Original nucleotide sequence data were collected from mitochondrial (cytochrome b) and nuclear (recombination activating gene 2) genes and used to estimate genetic variation and infer phylogenetic relationships among and within species of Noturus. Mitochondrial sequences were variable among species and several species were found to contain considerable genetic diversity. Relationships among members of the subgenus Rabida were resolved and in many cases well supported. Relationships among members of the subgenus Schilbeodes were poorly resolved. Previous phylogenetic hypotheses and the traditional classification (except the furiosus species group) were rejected in their explicit form according to the Kishino-Hasegawa and Shimodaira-Hasegawa tests of tree score difference.     

Historic speciation and recent colonization of Eurasian monkey gobies (Neogobius fluviatilis and N. pallasi) revealed by DNA sequences,microsatellites, and morphology

Aim Hidden diversity within an invasive ‘species’ can mask both invasion pathways and confound management goals. We assessed taxonomic status and population structure of the monkey goby Neogobius fluviatilis across Eurasia, comparing genetic variation across its native and invasive ranges. Location Native populations were analysed within the Black and Caspian Sea basins, including major river drainages (Dnieper, Dniester, Danube, Don and Volga rivers), along with introduced locations within the upper Danube and Vistula river systems. Methods DNA sequences and 10 nuclear microsatellite loci were analysed to test genetic diversity and divergence patterns of native and introduced populations; phylogenetic analysis of mtDNA cytochrome b and nuclear RAG‐1 sequences assessed taxonomic status of Black and Caspian Sea lineages. Multivariate analysis of morphology was used to corroborate phylogenetic patterns. Population genetic structure within each basin was evaluated with mtDNA and microsatellite data using F_ST analogues and Bayesian assignment tests. Results Phylogenetic analysis of mitochondrial and nuclear sequences discerned a pronounced genetic break between monkey gobies in the Black and Caspian Seas, indicating a long‐term species‐level separation dating to c. 3 million years. This pronounced separation further was confirmed from morphological and population genetic divergence. Bayesian inference showed congruent patterns of population structure within the Black Sea basin. Introduced populations in the Danube and Vistula River basins traced to north‐west Black Sea origins, a genetic expansion pattern matching that of other introduced Ponto‐Caspian gobiids. Main conclusions Both genetic and morphological data strongly supported two species of monkey gobies that were formerly identified as subspecies: N. fluviatilis in the Black Sea basin, Don and Volga Rivers, and the Kumo‐Manych Depression, and Neogobius pallasi in the Caspian Sea and Volga River delta. Genetic origins of introduced N. fluviatilis populations indicated a common invasion pathway shared with other introduced Ponto‐Caspian fishes and invertebrates.     

Gear effectiveness and size selectivity for five cryptic madtom species (Noturus spp.)

More than half of the diminutive North American catfish species known as madtoms (Noturus spp.) are considered imperiled due to range reductions from habitat fragmentation and degradation. In response, government agencies regularly conduct sampling that targets these cryptic fishes. Unfortunately, the effectiveness and biases associated with common sampling gears has rarely been quantified for non‐game fish, including madtoms. To improve sampling protocols for diverse madtom species, we: (a) quantified the detection probabilities and effort necessary to detect five species of madtom with relative certainty using complimentary sampling gears, and (b) described biases associated with size‐selectivity of the same gears when used to sample madtoms. Five madtom species that are native to Mississippi streams and have two distinct coloration patterns were selected: uniform pattern, Brown Madtom (Noturus phaeus) and Freckled Madtom (Noturus nocturnus); patterned, Piebald Madtom (Noturus gladiator), Least Madtom (Noturus hildebrandi), and Brindled Madtom (Noturus miurus). These species were captured using four standardized sampling gears (i.e., backpack electrofisher, seine, dipnet, and Gee‐style minnow trap) from 13 isolated stream reaches in Mississippi during baseflow conditions between August and November 2016. Each gear was randomly assigned to up to five transects at each site resulting in a maximum of 20 unique samples per stream reach. We estimated the detection probability of each species with each gear using presence–absence capture data. Cumulative detection probabilities were calculated for each species and gear combination to describe the number of samples necessary to achieve a detection probability of 0.95. Additionally, we pooled all capture data across sampling sites to compare madtom size distributions among gears. Electrofishing was generally the most efficient gear for detecting madtoms, but slightly overestimated fish size distributions. By seining, managers can expect to capture all species evaluated with a modest amount of effort; however, are likely to catch relatively small individuals. Seining was the most effective gear for only Piebald Madtom, a species petitioned for listing under the Endangered Species Act. To be relatively certain that Piebald Madtom are absent from a stream reach would require sampling ten 100 m transects by seining. In contrast, the same certainty would require more than twenty 100 m electrofishing transects. Minnow traps and dipnetting generally performed poorly and failed to detect some species at occupied sites. Biologists can now consider using each of these complimentary gears assessed when evaluating species status and population structure for diverse madtom species, while explicitly acknowledging the biases associated with each gear.     

INTRASPECIFIC PHYLOGEOGRAPHY OF NORTH AMERICAN HIGHLAND FISHES: A TEST OF THE PLEISTOCENE VICARIANCE HYPOTHESIS

The highland fish fauna of eastern North America consists of Appalachian and Ozark centers of endemism separated by the intervening Glacial Till Plains. Clades within these areas are more closely related phylogenetically to each other than to clades occurring in the intervening formerly glaciated region, suggesting that the Pleistocene glaciations fragmented a widespread highland region and its associated fauna. Alternatively, it is possible that these faunal assemblages predate the glaciations or that recent dispersals may have been more important than vicariance in determining faunal compositions. We examined the relationships among mitochondrial DNA (mtDNA) haplotypes within five clades of highland fishes, each with a distribution suggestive of a Pleistocene vicariance event. Darters of the subgenera Litocara and Odontopholis have distributions and mtDNA relationships that are consistent with the Pleistocene integration and burial of the Teays-Mahomet valley, a major drainage of the early Pleistocene. The distribution and mtDNA relationships among subspecies of Erimystax dissimilis are not consistent with Pleistocene vicariance, but relationships among Appalachian haplotypes are consistent with the late Pleistocene integration of the modern Ohio River system. Both Cottus carolinae and the Fundulus catenatus species group have representatives in the Mobile basin consistent with pre-Pleistocene divergences. Three haplotype clusters were found in C. carolinae, corresponding to the Appalachian, Ozark, and upper Kanawha River populations. However, Appalachian and Ozark F. catenatus populations are paraphyletic with respect to each other. This, coupled with a relatively low degree of sequence divergence, suggests that no long-term barriers to gene flow exist for C. carolinae and F. catenatus. These three distinct phylogeographic patterns indicate that Pleistocene vicariance is not the only explanation for the Appalachian-Ozark distribution of highland fish communities.     

Phylogeography of the bigeye chub Hybopsis amblops (Teleostei: Cypriniformes): early Pleistocene diversification and post‐glacial range expansion

The bigeye chub, Hybopsis amblops, is a member of the Central Highlands ichthyofauna of eastern North America. Phylogenetic analyses of the H. amblops species group based on a 1059 bp fragment of the mitochondrial DNA cytochrome b gene did not recover a monophyletic group. The inclusion of Hybopsis hypsinotus in the species complex is questionable. Within H. amblops, five strongly supported clades were identified; two clades containing haplotypes from the Ozark Highlands and three clades containing haplotypes from the Eastern Highlands and previously glaciated regions of the Ohio and Wabash River drainages. Estimates of the timing of divergence indicated that prior to the onset of glaciation, vicariant events separated populations east and west of the Mississippi River. East of the Mississippi River glacial cycles associated with the blocking and rerouting of the Teays River system caused populations to be pushed southward into refugia of the upper Ohio River. Following the most recent Wisconsinan glaciation, populations expanded northward into previously glaciated regions and southward into the Cumberland River drainage. In the Ozarks, west of the Mississippi River, isolation of clades appears to be maintained by the lack of stream capture events between the upper Arkansas and the White River systems and a barrier formed by the Arkansas River.     

Mitochondrial DNA divergence in the critically imperilled pygmy madtom,Noturus stanauli (Siluriformes: Ictaluridae)

The mitochondrial cytochrome b gene was sequenced for six individuals of the pygmy madtom, Noturus stanauli, a globally imperilled catfish, from both known localities in Tennessee (U.S.A.) separated by over 1055 river km. Phylogenetic and population genetic analyses revealed little divergence between these populations for this locus.     

Differences Found in the Macroinvertebrate Community Composition in the Presence or Absence of the Invasive Alien Crayfish,Orconectes hylas

Introductions of alien species into aquatic ecosystems have been well documented, including invasions of crayfish species; however, little is known about the effects of these introductions on macroinvertebrate communities. The woodland crayfish (Orconectes hylas (Faxon)) has been introduced into the St. Francis River watershed in southeast Missouri and has displaced populations of native crayfish. The effects of O. hylas on macroinvertebrate community composition were investigated in a fourth-order Ozark stream at two locations, one with the presence of O. hylas and one without. Significant differences between sites and across four sampling periods and two habitats were found in five categories of benthic macroinvertebrate metrics: species richness, percent/composition, dominance/diversity, functional feeding groups, and biotic indices. In most seasons and habitat combinations, the invaded site had significantly higher relative abundance of riffle beetles (Coleoptera: Elmidae), and significantly lower Missouri biotic index values, total taxa richness, and both richness and relative abundance of midges (Diptera: Chironomidae). Overall study results indicate that some macroinvertebrate community differences due to the O. hylas invasion were not consistent between seasons and habitats, suggesting that further research on spatial and temporal habitat use and feeding ecology of Ozark crayfish species is needed to improve our understanding of the effects of these invasions on aquatic communities.     

Phylogenetic relationships among holarctic populations of Chironomus entis and Chironomus plumosus in view of possible horizontal transfer of mitochondrial genes

In eight Holarctic populations of two typical chironomid sibling species of the plumosus group, Chrionomus entis and Chironomus plumosus, nucleotides sequences of mitochondrial (cytb) and nuclear (gb2b) gene regions were examined. The phylogenetic trees reflecting the evolutionary histories of the nuclear and mitochondrial markers exhibited significant differences. On the tree based on the nuclear gene sequences the populations clustered according to their species affiliation, whereas on the tree based on the mitochondrial gene sequences the populations were grouped according to their geographic position. This discrepancy is probably explained by mitochondrial gene flow between sympatric species with incomplete reproductive isolation (sibling species). Based on our results together with the earlier data on nuclear and mitochondrial gene sequences of some other species from the phylogenetic group plumosus, a scheme of phylogenetic relationships within this group is proposed. This scheme is in many ways different from the traditional view on the evolutionary relationships among species of the plumosus group.     

Molecular phylogeny of the genus Capoeta (Teleostei: Cyprinidae) in Anatolia,Turkey

Phylogeny of Capoeta genus distributed in Anatolia were carried out by analysing mitochondrial cytochrome b gene (1140 bp) sequences from 332 samples representing 59 populations of 15 species across their geographical distribution. Haplotype network and phylogenetic analysis (neighbor-joining, maximum-likelihood, maximum parsimony, and bayesian inference) of the 103 cytochrome b haplotypes detected in Capoeta species resulted in similar tree topologies including four distinct clades, in congruent with taxonomic classification of Capoeta based on morphological characteristics such as scale size, mouth shape, and body spotting. Based on cyt b nucleotide sequences, the present study suggests that four undescribed Capoeta species may exist in Anatolia freshwater; one species in the Kizilirmak River, the second species in the Dirgine River, the third species B. Menderes River, and the fourth species in the some Yesilirmak tributaries that run into the Black Sea Basin. Capoeta taxa distributed in the rivers of Anatolian freshwater basins are isolated from each other during middle Miocene (Serravallian)-late Pleistocene (Ionian) (about 13.75–0.41 million years). This suggests that distribution and presence of Capoeta species were shaped under paleogeographic conditions such as Pleistocene climate changes in Quarternary period as well as tectonic uplift and faulting, which probably has not changed up to now.     

Quantitative Phylogenomics of Within-Species Mitogenome Variation: Monte Carlo and Non-Parametric Analysis of Phylogeographic Structure among Discrete Transatlantic Breeding Areas of Harp Seals (Pagophilus groenlandicus)

Phylogenomic analysis of highly-resolved intraspecific phylogenies obtained from complete mitochondrial DNA genomes has had great success in clarifying relationships within and among human populations, but has found limited application in other wild species. Analytical challenges include assessment of random versus non-random phylogeographic distributions, and quantification of differences in tree topologies among populations. Harp Seals (Pagophilus groenlandicus Erxleben, 1777) have a biogeographic distribution based on four discrete trans-Atlantic breeding and whelping populations located on “fast ice” attached to land in the White Sea, Greenland Sea, the Labrador ice Front, and Southern Gulf of St Lawrence. This East to West distribution provides a set of a priori phylogeographic hypotheses. Outstanding biogeographic questions include the degree of genetic distinctiveness among these populations, in particular between the Greenland Sea and White Sea grounds. We obtained complete coding-region DNA sequences (15,825 bp) for 53 seals. Each seal has a unique mtDNA genome sequence, which differ by 6 ~ 107 substitutions. Six major clades / groups are detectable by parsimony, neighbor-joining, and Bayesian methods, all of which are found in breeding populations on either side of the Atlantic. The species coalescent is at 180 KYA; the most recent clade, which accounts for 66% of the diversity, reflects an expansion during the mid-Wisconsinan glaciation 40 ~ 60 KYA. F_ST is significant only between the White Sea and Greenland Sea or Ice Front populations. Hierarchal AMOVA of 2-, 3-, or 4-island models identifies small but significant Φ_SC among populations within groups, but not among groups. A novel Monte-Carlo simulation indicates that the observed distribution of individuals within breeding populations over the phylogenetic tree requires significantly fewer dispersal events than random expectation, consistent with island or a priori East to West 2- or 3-stepping-stone biogeographic models, but not a simple 1-step trans-Atlantic model. Plots of the cumulative pairwise sequence difference curves among seals in each of the four populations provide continuous proxies for phylogenetic diversification within each. Non-parametric Kolmogorov-Smirnov (K-S) tests of maximum pairwise differences between these curves indicates that the Greenland Sea population has a markedly younger phylogenetic structure than either the White Sea population or the two Northwest Atlantic populations, which are of intermediate age and homogeneous structure. The Monte Carlo and K-S assessments provide sensitive quantitative tests of within-species mitogenomic phylogeography. This is the first study to indicate that the White Sea and Greenland Sea populations have different population genetic histories. The analysis supports the hypothesis that Harp Seals comprises three genetically distinguishable breeding populations, in the White Sea, Greenland Sea, and Northwest Atlantic. Implications for an ice-dependent species during ongoing climate change are discussed.     

Evolution after the flood: phylogeography of the desert fish Utah Chub

The Bonneville Basin and upper Snake River drainage of western North America underwent extensive hydrological changes during the late Pleistocene, potentially influencing the geographic distribution and evolutionary trajectories of aquatic species that occupied this region. To test this hypothesis, I reconstructed the phylogeographic history of the desert fish Utah chub (Gila atraria) by examining 16 populations that span the natural distribution of this species across the Bonneville Basin and upper Snake River. I compared mitochondrial control region sequences (934 bp) among 77 individuals revealing 24 unique haplotypes. Geographic and phylogenetic relationships among haplotypes were explored using parsimony, maximum likelihood, nested clade analysis, and analysis of molecular variance. I found that G. atraria is composed of two distinct clades that represent an early Pleistocene split between the upper Snake River and Bonneville Basin. Within each of these clades, geographic structuring was highly concordant with the hydrological history of late Pleistocene Lake Bonneville and the upper Snake River, suggesting that glacial-induced shifts in climate and unpredictable geological events have played a major role in shaping genetic subdivision among populations. To examine the effects of vicariant events on phenotypic divergence among Utah chub populations, I mapped chub life histories to the control region haplotype network. I found a nonrandom association between haplotypes and life-history phenotypes. These results suggest that historical events responsible for population fragmentation may have also contributed to phenotypic shifts in life histories, both indirectly by limiting gene flow among populations and directly by altering the selective environments where populations persisted.     

On the diversity and taxonomic status of graylings (Thymallus Thymallidae) from the Lena River

Two forms of graylings were distinguished in the Lena River, which differ in body coloration, the shape and pattern of the dorsal fin, and biological characteristics. With respect to these traits, the forms are divided between the upper and lower Lena. The first inhabits most of the basin, from the upper courses to the lower reaches, whereas the second inhabits the Lena delta and the adjacent arctic rivers and their tributaries. Comparative analysis of meristic traits revealed their low appropriateness for diagnosing Lena lineages as well as grayling populations inhabiting other northern rivers of Siberia. The body and fin coloration in the upper Lena form significantly differ from that found in forms and subspecies of graylings from the basins of the other rivers in the ranges of Eastern Siberian, Kamchatka, and Alaska. The lower Lena form should be classified with the Eastern Siberian subspecies T. arcticus pallasii and is closer to the populations of graylings from the rivers Anabar and Yana, and the rivers of the Chukot Peninsula and Alaska, and the upper courses of the Missouri. These results are supported by analysis of the mitochondrial DNA of graylings throughout the Paleoarctic. Further studies of these lineages and phylogenetic relationships between Arctic lineages are necessary for the final determination of the taxonomic status of the upper Lena form.     

Phylogenetic Relationships among Holarctic Populations of Chironomus entis and Chironomus plumosus in View of Possible Horisontal Transfer of Mitochondrial Genes

In eight Holarctic populations of two typical chironomid sibling species of the plumosus group, Chironomus entisandChironomus plumosus, nucleotide sequences of mitochondrial (cytb) and nuclear (gb2b) gene regions were examined. The phylogenetic trees reflecting the evolutionary histories of the nuclear and mitochondrial markers exhibited significant differences. On the tree based on the nuclear gene sequences the populations clustered according to their species affiliation, whereas on the tree based on the mitochondrial gene sequences the populations were grouped according to their geographic position. This discrepancy is probably explained by mitochondrial gene flow between sympatric species with incomplete reproductive isolation (sibling species). Based on our results together with the earlier data on nuclear and mitochondrial gene sequences of some other species from the phylogenetic group plumosus, a scheme of phylogenetic relationships within this group is proposed. This scheme is in many ways different from the traditional view on the evolutionary relationships among species of the plumosus group.