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Baykina E.M.,State Darwin Museum | Schwarzhans W.W.,Universitetsparken 15
Swiss Journal of Palaeontology | Year: 2017

Karaganops is established as a new fossil genus of the family Clupeidae, subfamily Clupeinae, to encompass the nominal species Sardinella perrata Daniltshenko 1970 from the Karaganian of southern Russia. Karaganops belongs to the group of genera characterized by the presence of two elongated rays at the end of the anal fin, which further includes Sardinella, Clupeonella, Sardina, Sardinops, and Harengula. It differs from Sardinella amongst others in the absence of tooth plates on the palatines and pterygoids, the presence of seven branchiostegal rays (vs. six), the presence of two epurals (vs. three) and the presence of nine pelvic fin rays (vs. 7–8). Otoliths found in situ further support the separation of Karaganops from Sardinella and related genera and add evidence for a second, otolith-based species—Karaganops komochtitziensis (Strashimirov 1985)—from the Middle Sarmatian of Bulgaria. Karaganops is thought to represent an endemic clupeid genus of the Eastern Paratethys that evolved during the mid-Miocene Karaganian ecological crisis in the basin. An otolith found in situ in a skeleton of Sarmatella doljeana (Kramberger 1883) from the Early Sarmatian of Belgrade, Serbia and an otolith interpreted to be associated with another fossil clupeid genus and species—Sarmatella tsurevica (Baykina 2012)—from the Early Sarmatian of the Caucasus facilitates correlation with previously described otolith-based species. It demonstrates that Sarmatella may not represent a primary endemic of the Paratethys and that its fossil record reaches back in time to Early Miocene and possibly Late Oligocene in the North Sea Basin. © 2016, Akademie der Naturwissenschaften Schweiz (SCNAT).


Zink R.M.,University of Minnesota | Ericson P.G.P.,Swedish Museum of Natural History | Fadeev I.V.,State Darwin Museum
Journal of Biogeography | Year: 2010

Aim: Boreal forest bird species appear to be divided into lineages endemic to each northern continent, in contrast to Holarctic species living in open habitats. For example, the three-toed woodpecker (Picoides tridactylus) and the winter wren (Troglodytes troglodytes) have divergent Nearctic and Palaearctic mitochondrial DNA clades. Furthermore, in these species, the next closest relative of the Nearctic/Palaearctic sister lineages is the Nearctic clade, suggesting that the Palaearctic may have been colonized from the Nearctic. The aim of this study is to test this pattern of intercontinental divergence and colonization in another Holarctic boreal forest resident - the pine grosbeak (Pinicola enucleator). Location: The Holarctic. Methods: We sequenced the mitochondrial ND2 gene and Z-specific intron 9 of the ACO1 gene for 74 pine grosbeaks collected across the Holarctic. The sequences were used to reconstruct the phylogeographical history of this species using maximum likelihood analysis. Results: We discovered two distinct mitochondrial and Z-specific lineages in the Nearctic and one in the Palaearctic. The two Nearctic mtDNA lineages, one in the northern boreal forest and one in south-western mountain forest, were more closely related to each other than either was to the Palaearctic clade. Two Nearctic Z-chromosome clades were sympatric in the boreal and south-western mountain forests. Unlike the topology of the mtDNA tree, the relationship among the Z-chromosome clades was the same as in the three-toed woodpecker and winter wren [Nearctic (Nearctic, Palaearctic)]. The Palaearctic Z-chromosome clade had much lower genetic diversity and a single-peak mismatch distribution with a mean < 25% of that for either Nearctic region, both of which had ragged mismatch distributions. Main conclusions: Our data suggest that, similar to the other boreal forest species, the pine grosbeak has divergent lineages in each northern continent and could have colonized the Palaearctic from the Nearctic. Compared with many Holarctic birds inhabiting open habitats, boreal forest species appear to be more differentiated, possibly because the boreal forests of the Nearctic and Palaearctic have been isolated since the Pliocene (3.5 Ma). © 2009 Blackwell Publishing Ltd.


Drovetski S.V.,University of Tromsø | Rakovic M.,Natural History Museum Belgrade | Semenov G.,Russian Academy of Sciences | Fadeev I.V.,State Darwin Museum | Red'kin Y.A.,Moscow State University
PLoS ONE | Year: 2014

Phylogeographic studies of Holarctic birds are challenging because they involve vast geographic scale, complex glacial history, extensive phenotypic variation, and heterogeneous taxonomic treatment across countries, all of which require large sample sizes. Knowledge about the quality of phylogeographic information provided by different loci is crucial for study design. We use sequences of one mtDNA gene, one sex-linked intron, and one autosomal intron to elucidate large scale phylogeographic patterns in the Holarctic lark genus Eremophila. The mtDNA ND2 gene identified six geographically, ecologically, and phenotypically concordant clades in the Palearctic that diverged in the Early - Middle Pleistocene and suggested paraphyly of the horned lark (E. alpestris) with respect to the Temminck's lark (E. bilopha). In the Nearctic, ND2 identified five subclades which diverged in the Late Pleistocene. They overlapped geographically and were not concordant phenotypically or ecologically. Nuclear alleles provided little information on geographic structuring of genetic variation in horned larks beyond supporting the monophyly of Eremophila and paraphyly of the horned lark. Multilocus species trees based on two nuclear or all three loci provided poor support for haplogroups identified by mtDNA. The node ages calculated using mtDNA were consistent with the available paleontological data, whereas individual nuclear loci and multilocus species trees appeared to underestimate node ages. We argue that mtDNA is capable of discovering independent evolutionary units within avian taxa and can provide a reasonable phylogeographic hypothesis when geographic scale, geologic history, and phenotypic variation in the study system are too complex for proposing reasonable a priori hypotheses required for multilocus methods. Finally, we suggest splitting the currently recognized horned lark into five Palearctic and one Nearctic species. © 2014 Drovetski et al.


Drovetski S.V.,University of Tromsø | Semenov G.,Russian Academy of Sciences | Drovetskaya S.S.,19355 53rd Ave. NE | Fadeev I.V.,State Darwin Museum | And 2 more authors.
Ecology and Evolution | Year: 2013

Mountains host greater avian diversity than lowlands at the same latitude due to their greater diversity of habitats stratified along an elevation gradient. Here we test whether this greater ecological heterogeneity promotes sympatric speciation. We selected accentors (Prunellidae), an avian family associated with mountains of the Palearctic, as a model system. Accentors differ in their habitat/elevation preferences and south-central Siberia and Himalayan regions each host 6 of the 13 species in the family. We used sequences of the mtDNA ND2 gene and the intron 9 of the Z chromosome specific ACO1 gene to reconstruct a complete species-level phylogeny of Prunellidae. The tree based on joint analysis of both loci was used to reconstruct the family's biogeographic history and to date the diversification events. We also analyzed the relationship between the node age and sympatry, to determine the geographic mode of speciation in Prunellidae. Our data suggest a Miocene origin of Prunellidae in the Himalayan region. The major division between alpine species (subgenus Laiscopus) and species associated with shrubs (subgenus Prunella) and initial diversification events within the latter happened within the Himalayan region in the Miocene and Pliocene. Accentors colonized other parts of the Palearctic during the Pliocene-Pleistocene transition. This spread across the Palearctic resulted in rapid diversification of accentors. With only a single exception dating to 0.91 Ma, lineages younger than 1.5 Ma are allopatric. In contrast, sympatry values for older nodes are >0. There was no relationship between node age and range symmetry. Allopatric speciation (not to include peripatric) is the predominant geographic mode of speciation in Prunellidae despite the favorable conditions for ecological diversification in the mountains and range overlaps among species. In this study, we test whether greater diversity of habitats stratified along an elevation gradient in the mountains promotes sympatric speciation in birds. Using sequences of one mtDNA gene and one sex specific intron, we reconstructed a complete species-level phylogeny of Prunellidae, which we used to reconstruct the family's biogeographic history. Our data suggest that allopatric speciation (not to include peripatric) is the predominant geographic mode of speciation in Prunellidae despite the favorable conditions for ecological diversification in the mountains and range overlaps among species. © 2013 The Authors. Ecology and Evolution.


PubMed | State Darwin Museum, University of Tromsø, Moscow State University, Russian Academy of Sciences and Kirov City Zoological Museum
Type: Journal Article | Journal: PloS one | Year: 2015

When isolated but reproductively compatible populations expand geographically and meet, simulations predict asymmetric introgression of neutral loci from a local to invading taxon. Genetic introgression may affect phylogenetic reconstruction by obscuring topology and divergence estimates. We combined phylogenetic analysis of sequences from one mtDNA and 12 nuDNA loci with analysis of gene flow among 5 species of Pacific Locustella warblers to test for presence of genetic introgression and its effects on tree topology and divergence estimates. Our data showed that nuDNA introgression was substantial and asymmetrical among all members of superspecies groups whereas mtDNA showed no introgression except a single species pair where the invaders mtDNA was swept by mtDNA of the local species. This introgressive sweep of mtDNA had the opposite direction of the nuDNA introgression and resulted in the paraphyly of the local species mtDNA haplotypes with respect to those of the invader. he multilocus nuDNA species tree resolved all inter- and intraspecific relationships despite substantial introgression. However, the node ages on the species tree may be underestimated as suggested by the differences in node age estimates based on non-introgressing mtDNA and introgressing nuDNA. In turn, the introgressive sweep and strong purifying selection appear to elongate internal branches in the mtDNA gene tree.


Rubtsov A.S.,State Darwin Museum
Biology Bulletin | Year: 2015

Biological diversity is continuous as a result of evolutionary processes, whereas the language for its description is symbolic and, therefore, discrete. Thus, the main question of the “species problem” is as follows: do species really exist in nature or not? The answer of this question depends on the “system of coordinates” that is used for the description of species. Species inhabiting the same territory (“one-dimensional species”) are usually discrete. However, when we begin to describe closely related forms within their ranges (“two-dimensional species”), the determination of the interspecific border becomes problematic. Therefore, speciation can be defined as the process of establishing the sympatry between diverging taxa, and species can be defined as morphologically distinguishable groups of organisms that can coexist in the sympatry zone for a long time. The diversity of the existing species concepts can be reduced to the two main ones—biological and phylogenetic. The latter has a disadvantage compared to the former because it does not deal with the theoretical substantiation of the species reality. But within the framework of the concepts of biological species, the theoretical substantiation of species reality (the hypothesis of the reinforcement of isolating mechanisms in a hybrid zone) has no reliable empirical support too. Another substantial disadvantage is that the use of the biological concept in practice may lead to misinterpretation of the taxonomic status of forms at different stages of the speciation process. At the initial stages of divergence, taxa can be regarded as independent species (for example, ecological races of the red crossbill (Loxia curvirostra), whereas at the final stage of speciation, forms are regarded as conspecific due to their hybridization in the sympatry zone (the yellowhammer Emberiza citrinella and the pine bunting E. leucocephala). The “species problem” seems to be solvable using a synthesis of the biological and phylogenetic concepts of species. © 2015, Pleiades Publishing, Inc.


Rubtsov A.S.,State Darwin Museum | Opaev A.S.,RAS Severtsov Institute of Ecology
Biology Bulletin | Year: 2012

The yellowhammer and pine bunting represent a quite rare case of mass hybridization in a broad secondary contact zone. Previously, it was shown that the two forms did not differ by mitochondrial DNA (Irwin et al., 2009). This result allowed suggesting either a very close evolutionary relationship between these species or a consequence of borrowing the mitochondrial genome due to long-term hybridization. An attempt to reconstruct the phylogeny of these species was made on the basis of cluster analysis performed relative to the song characters and morphological features (male coloration and morphometric characters). The geographical variability of songs in yellowhammers and pine buntings was also investigated. The cladistic analysis included another 5 closely related bunting species: cirl (E. cirlus), chestnut-breasted (E. stewarti), ortolan (E. hortulana), grey-necked (E. buchanani), and Cretzschmar's bunting (E. caesia). Based on the results of our analysis, the yellowhammer and pine bunting show no sister relationship. They compose the smallest clusters: one with the cirl bunting and the other one with the chestnut-breasted bunting. Thus, in this paper we attempt to prove the hypothesis of a high divergence level between the species and mtDNA introgression from the pine bunting into the yellowhammer following long term hybridization. The analysis of song dialects agrees with the assumption that there existed an ancient hybrid zone between the yellowhammer and the pine bunting in Europe. © 2012 Pleiades Publishing, Ltd.


Rubtsov A.S.,State Darwin Museum
Zoologicheskii Zhurnal | Year: 2015

Biological diversity is continuous as a result of evolutionary processes, whereas the language for its description is symbolic and, therefore, discrete. So, the main question of the "species problem" is that: do species really exist in nature or not? The answer of this question depends on a "system of coordinates", which is used for the description of species. The species inhabiting the same territory ("one-dimension species") are usually discrete. However, when we begin to describe closely related forms within their ranges ("two-dimension species"), the determination of interspecific border becomes problematic. One can define speciation as a process of establishing the sympatry between diverging taxa and species, as morphologically distinguished groups of organisms that can coexist in the same territory for a long time. The diversity of the existing species concepts may be reduced to the two main types - biological and phylogenetic. The latter has a disadvantage as compared to the former because it is not deal with the problem of species reality. Within the framework of biological species concepts, the theoretical substantiation of species reality (the reinforcement hypothesis) has no reliable empiric support. Moreover, the practical use of biological concepts may lead to a misinterpretation of the speciation process. At the initial stages of diversification, forms can be described as independent species (for example, ecological races of the red crossbill, Loxia curvirostra), whereas at the final stage of speciation, they are regarded as conspecific ones due to their hybridization (yellowhammer Emberiza citrinella and pine bunting E. leucocephala). The "species problem" seems to be solved using the synthesis of biological and phylogenetic species concepts. © 2015, Russian Academy of Sciences. All rights reserved.


Baykina E.M.,State Darwin Museum | Schwarzhans W.W.,Universitetsparken 15
Swiss Journal of Palaeontology | Year: 2016

Moldavichthys switshenskae is established as a new fossil genus and species of the family Clupeidae, subfamily Alosinae, to encompass the specimens from the Sarmatian of Moldova, which hitherto have been recorded as Clupeonella humilis (v.Meyer, 1851). Specimens from the type area and strata (Ottnangian of the Western Paratethys) differ from the Moldovan ones in a greater number of vertebrae, the more posterior position of dorsal fin and a greater number of postventral scutes. Their generic allocation requires review. Moldavichthys resembles the recent genus Alosa, known from the northern Atlantic, NE Pacific, the Mediterranean and the Ponto-Caspian Sea. Moldavichthys differs from Alosa in the shape of the jaw and the opercular bones and the otolith morphology and is thought to represent an extinct Paratethyan endemic lineage. Otolith in situ was found in three skeletons of Moldavichthys switshenskae. They are unusually compact and compressed when compared to Recent Alosa otoliths, with the caveat however, that very little is known of otoliths from the extant Ponto-Caspian Alosa species. There are no isolated fossil otoliths known, which would correspond to M. switshenskae, although recently Alosa paulicrenata Bratishko, Schwarzhans & Reichenbacher 2015 has been described from the Konkian of Kazakhstan. A review of Sarmatian otoliths from Bulgaria has revealed a species, which we tentatively assign as Moldavichthys? gomotartziensis (Strashimirov, 1985). © 2016 Akademie der Naturwissenschaften Schweiz (SCNAT)


Rubtsov A.S.,State Darwin Museum | Opaev A.S.,RAS Severtsov Institute of Ecology
Zoologicheskii Zhurnal | Year: 2012

Yellowhammer and pine bunting represent a quite rare case of mass hybridization in the broad secondary contact zone. Previously, it was shown that the two forms did not differ by mitochondrial DNA (Irwin et al., 2009). This result allowed suggesting a very close evolutionary relationship between these species or to be considered as a consequence of borrowing the mitochondrial genome due to the long-term hybridization. An attempt to reconstruct phylogeny of these species was made on the basis of the cluster analysis performed irrelative of the song characteristics and morphological features (male coloration and morphometric characters). The geographical variability of songs in yellowhammers and pine buntings was also investigated. Five other closely related bunting species to the analyzed ones were included into the cladistic analysis: cirl (E. cirlus), chestnut-breasted (E. stewarti), ortolan (E. hortulana), grey-necked (E. buchanani), and Cretzschmar's bunting (E. caesia). Based on the results of the analysis, yellowhammer and pine bunting are not sister species. They compose the smallest clusters: one with the cirl bunting and the other one with the chestnut-breasted bunting. Thus, the hypothesis of the high divergence level of the species studied followed by the long-term hybridization between the two species is confirmed. The analysis of song dialects fits the assumption of an ancient hybrid zone between yellowhammer and pine bunting in Europe.

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