The uplift of Qinghai-Xizang (Tibet) Plateau and the vicariance speciation of glyptosternoid fishes (Siluriformes: Sisoridae)

Based on the phylogenetic and biogeographical studies of the glyptosternoid fishes in Qinghai-Tibet area, the following hypothesis is proposed: the speciation of this group has a direct relationship with the three uplift intervals of the Qinghai-Tibet Plateau. This process was explained by the theory of vicariance of biogeography. The ancestor of this group was similar to Bagarus and/or Glyptothorax, which still have a wide distribution. At the moment when the Tethys sea closed, the Indian tectonic plate collided with the Eurasian tectonic plate, so the Glyptothorax-like and Bagarus-like ancestors entered Eurasia and gradually became widely distributed. After the Pleistocene, with the enforced colliding, the gradual uplift of the Qinghai-Tibet Plateau brought about the current water environment, and the Glyptosternoids were generated from Glyptotho-rax-like fish under this environment. The present Glyptosternum, distributed across the Himalayas is the ancestor of Glyptosternoids. In the three uplift inte     

The uplift of the Qinghai–Tibet Plateau and glacial oscillations triggered the diversification of Tetraogallus (Galliformes,Phasianidae)

The Qinghai–Tibet Plateau (QTP) plays an important role in avian diversification. To reveal the relationship between the QTP uplift and avian diversification since the Late Cenozoic, here, we analyzed the phylogenetic relationship and biogeographical pattern of the genus Tetraogallus (Galliformes, Phasianidae) and the probable factors of speciation in the period of the QTP uplift inferred from concatenated data of four nuclear and five mitochondrial genes using the method of the Bayesian inference. Phylogenetic analysis indicated that T. himalayensis had a close relationship with T. altaicus and conflicted with the previous taxonomy of dark‐bellied and white‐bellied groups. The molecular clock showed that the speciation of Tetraogallus was profoundly affected by the uplift of the QTP and glacial oscillations. Biogeographic analysis suggested that the extant snowcocks originated from the QTP, and the QTP uplift and glacial oscillations triggered the diversification of Tetraogallus ancestor. Specifically, the uplift of the mountain provided a prerequisite for the colonization of snowcocks Tetraogallus as a result of the collision between the Indian and the Arab plates and the Eurasian plate, in which ecological isolation (the glacial and interglacial periods alternate) and geographical barrier had accelerated the Tetraogallus diversification process. Interestingly, we discovered hybrids between T. tibetanus and T. himalayensis for the first time and suggested that T. tibetanus and T. himalayensis hybridized after a second contact during the glacial period. Here, we proposed that the hybrid offspring was the ancestor of the T. altaicus. In conclusion, the uplift of QTP and glacial oscillations triggered the snowcocks colonization, and then, isolation and introgression hybridization promoted diversification.     

Biogeography and molecular phylogeny of the genus Schizothorax (Teleostei: Cyprinidae) in China inferred from cytochrome b sequences

Aim To test a vicariant speciation hypothesis derived from geological evidence of large‐scale changes in drainage patterns in the late Miocene that affected the drainages in the south‐eastern Tibetan Plateau. Location The Tibetan Plateau and adjacent areas. Methods The cytochrome b DNA sequences of 30 species of the genus Schizothorax from nine different river systems were analysed. These DNA sequences were analysed using parsimony, maximum likelihood and Bayesian methods. The approximately unbiased and Shimodaira–Hasegawa tests were applied to evaluate the statistical significance of the shortest trees relative to alternative hypotheses. Dates of divergences between lineages were estimated using the nonparametric rate smoothing method, and confidence intervals of dates were obtained by parametric bootstrapping. Results The phylogenetic relationships recovered from molecular data were inconsistent with traditional taxonomy, but apparently reflected geographical associations with rivers. Within the genus Schizothorax, we observed a divergence between the lineages from the Irrawaddy–Lhuit and Tsangpo–Parlung rivers, and tentatively dated this vicariant event back to the late Miocene (7.3–6.8 Ma). We also observed approximately simultaneous geographical splits within drainages of the south‐eastern Tibetan Plateau, the Irrawaddy, the Yangtze and the Mekong–Salween rivers in the late Miocene (7.1–6.2 Ma). Main conclusions Our molecular evidence tentatively highlights the importance of palaeoriver connections and the uplift of the Tibetan Plateau in understanding the evolution of the genus Schizothorax. Molecular estimates of divergence times allowed us to date these vicariant scenarios back to the late Miocene, which agrees with geological suggestions for the separation of these drainages caused by tectonic uplift in south‐eastern Tibet. Our results indicated the substantial role of vicariant‐based speciation in shaping the current distribution pattern of the genus Schizothorax.     

Dispersal vs. vicariance in the Mediterranean: historical biogeography of the Palearctic Pachydeminae (Coleoptera, Scarabaeoidea)

Aim The geological evolution of the Mediterranean region is largely the result of the Tertiary collision of the African and Eurasian Plates, but also a mosaic of migrating island arcs, fragmenting tectonic belts, and extending back‐arc basins. Such complex paleogeography has resulted in a ‘reticulate’ biogeographical history, in which Mediterranean biotas repeatedly fragmented and merged as dispersal barriers appeared and disappeared through time. In this study, dispersal‐vicariance analysis (DIVA) is used to assess the relative role played by dispersal and vicariance in shaping distribution patterns in the beetle subfamily Pachydeminae Reitter, 1902 (Scarabaeoidea), an example of east–west Mediterranean disjunction. Location The Mediterranean region, including North Africa, the western Mediterranean, Balkans–Anatolia, Middle East, Caucasus, the Iranian Plateau, and Central Asia. Methods A phylogenetic hypothesis of the Palearctic genera of Pachydeminae in conjunction with distributional data was analysed using DIVA. This method reconstructs the ancestral distribution in a given phylogeny based on the vicariance model, while allowing dispersal and extinction to occur. Unlike other methods, DIVA does not enforce area relationships to conform to a hierarchical ‘area cladogram’, so it can be used to reconstruct ‘reticulate’ biogeographical scenarios. Results Optimal reconstructions, requiring 23 dispersal events, suggest that the ancestor of Pachydeminae was originally present in the south‐east Mediterranean region. Basal splitting within the subfamily was caused by vicariance events related to the late Tertiary collision of the African microplates Apulia and Arabia with Eurasia, and the resultant arise of successive dispersal barriers (e.g. the Red Sea, the Zagros Mountains). Subsequent diversification in Pachydeminae involved multiple speciation events within the Middle East and Iran–Afghanistan regions, which gave rise to the least speciose genera of Pachydeminae (e.g. Otoclinius Brenske, 1896). Finally, the presence of Pachydeminae in the western Mediterranean region seems to be the result of a recent dispersal event. The ancestor of the Iberian genera Ceramida Baraud, 1987 and Elaphocera Gené, 1836 probably dispersed from the Middle East to the Iberian Peninsula across North Africa and the Gibraltar Strait during the ‘Messinian salinity crisis’ at the end of the Miocene. Main conclusions Although the basal diversification of Pachydeminae around the Mediterranean appears to be related to vicariance events linked to the geological formation of the Mediterranean Basin, dispersal has also played a very important role. Nearly 38% of the speciation events in the phylogeny resulted from dispersal to a new area followed by allopatric speciation between lineages. Relationships between western and eastern Mediterranean disjuncts are usually explained by dispersal through Central Europe. The biogeographical history of the Pachydeminae corroborates other biogeographical studies that consider North Africa to be an alternative dispersal route by which Mediterranean taxa could have achieved circum‐Mediterranean distributions.     

Patterns of diversification in the high Andean Ponderacris grasshoppers (Orthoptera: Acrididae: Melanoplinae)

The Andes, the world's longest mountain chain, harbours great taxonomic and ecological diversity. Despite their young age, the tropical Andes are highly diverse due to recent geological uplift. Speciation either followed the orogeny closely or occurred after the Andean uplift, as a result of subsequent climatic changes. Different scenarios have been proposed to explain the diversification of high Andean taxa. The Melanoplinae grasshopper Ponderacris Ronderos & Cigliano is endemic to the eastern slopes of the Andes of Peru and Bolivia, mostly distributed between 1000 and 4000 m above sea level. Diversification in several montane habitats of Bolivia and Peru allows tests via cladistic analysis of distinct possible geographic modes of speciation. Eight species are recognized, with three described here as new with revised diagnostic morphological characters provided: Ponderacris carlcarbonelli sp.n., P. chulumaniensis sp.n. and P. amboroensis sp.n. Cladistic analyses of 15 species (8 ingroup and 7 outgroup) and 38 morphological characters, under equal and implied weighting, confirm the monophyly of Ponderacris. Characters from the external morphology and colour pattern provided less phylogenetic information than did the male abdominal terminalia and phallic complex. Species distributed in the Peruvian Andes constituted a monophyletic group, whereas those from the Bolivian Andes formed a basal paraphyletic grade. Dispersal–vicariance analysis resulted in one ancestral distribution reconstruction indicating that the most recent common ancestor was distributed in the Lower Montane Yungas of Bolivia. Eleven dispersal and one vicariant events are postulated, with a South‐to‐North speciation pattern coincident with progressive Andean uplift. Vicariance could relate to fragmentation of montane forest during the dry intervals of the late Cenozoic. From the Bolivian area, ancestral Peruvian Ponderacris may have dispersed northward, coinciding with the rise of the Andes. Ten of 11 dispersal events occurred at terminal taxa and are likely to be recent. However, diversification of Ponderacris cannot be explained solely by the South‐to‐North speciation hypothesis, but may also include both vicariance and dispersal across barriers influenced by Pleistocene climatic cycles.     

Unravelling the evolutionary history of the polyploid complex Ranunculus parnassiifolius (Ranunculaceae)

Ranunculus L. represents the largest genus within Ranunculaceae, comprising more than 600 species with a worldwide distribution. However, there are still many gaps in our knowledge of the infrageneric taxonomy and evolution of Ranunculus. In this regard, intraspecific variation of the polyploid complex Ranunculus parnassiifolius remains under discussion. To reconstruct the biogeographical history of the polyploid complex R. parnassiifolius, 20 populations distributed throughout the Cantabrian Mountains, Pyrenees, and Alps were investigated. Phylogenetic studies were based on nuclear internal transcribed spacers (ITS) and plastid (rpl32‐trnL, rps16‐trnQ) sequence data, analysed using Bayesian approaches as well as the evolution of morphological characters. Additionally, biogeographical patterns were conducted using statistical dispersal–vicariance analysis. The analyses presented here support the recognition of two evolutionary independent units: R. cabrerensis sensu lato (s.l.) and R. parnassiifolius s.l. Furthermore gradual speciation depending on the biogeographical territory is proposed, and optimal reconstructions have probably favoured the ancestor of Ranunculus parnassiifolius as originating in the Iberian Peninsula. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 107 , 477–493.     

The phylogeny ofActaea (Ranunculaceae): A biogeographical approach

Phylogenetic analyses of the genusActaea were performed using morphological, ecological and biogeographical characters. Using solely morphological characters, the relationships of the three identified species-groups remain uncertain. Close biogeographical examination and comparison of the areas with ecological peculiarities as well as climate data gave important insight into the phylogeny ofActaea and the whole tribe. Consequently, the obtained biogeographical data were used for phylogenetic reconstructions. Both, from the point of view of morphological and biogeographical data,A. pachypoda andA. asiatica are the most ancestral species. They grow on the east sides of the continents, mainly in broad-leaved forests. In West Eurasia the apomorphicA. spicata andA. acuminata occur under similar climatic and ecological conditions, but these species are adapted to another climate rhythm. The most advanced species (A. erythrocarpa, A. rubra) are to be found in the boreal forests where they are widely distributed. This biogeographical approach revealed that the evolution of the species led to a gradual widening and shifting of their ecological constitutions.     

Phylogeny and biogeography Eupholidoptera Mařan (Orthoptera,Tettigoniidae): morphological speciation in correlation with the geographical evolution of the eastern Mediterranean

Recently, the systematics and biogeography of the Mediterranean biota have received much attention. This paper deals with Eupholidoptera Ma?an, a Mediterranean lineage of Tettigoniidae. The genus is restricted to the northern and eastern basin of the Mediterranean, with a significant number of species found on the Aegean islands. To produce a phylogeny and use it to make assumptions about the historical biogeography of Eupholidoptera, material of 46 species from several collections was studied. A phylogenetic analysis based mainly on morphological characters suggested two lineages in the genus: the E. chabrieri and the E. prasina groups. Based on the consistency between historical geographical events and branching events on the phylogenetic tree, Eupholidoptera is assumed to have evolved from an ancestor present in the Aegeid plate in the Mid‐Miocene. The division of the Aegeid plate into Anatolia and Greece in the Tortonian, the reoccurrence of terrestrial corridors between these mainlands in the Messinian, the regression of the Aegean area in the Pliocene and sea level changes in the Pleistocene are assumed to have been the main palaeogeographical events directing speciation in Eupholidoptera. As most of the species are allopatric, vicariance is suggested to be the main pattern. By combining the nature of the characters used in the phylogenetic analysis, the phylogenetic tree produced and the biogeographical assumptions, four tentative conclusions can be made: (i) radiation in the genus is a result of divergence in morphology; (ii) because the main character source is male genitalia, there has possibly been intensive sexual selection, which leads to morphological speciation; (iii) as the difference in temporal parameters of the song is prominent in sympatric/parapatric species pairs only, co‐occurrence is suggested to be the main reason driving divergence in the song; (iv) there seems to be a negative correlation between the size of the distribution range and the evolutionary rate in speciation; this may be the reason why the E. prasina group (restricted to a small part of the range of the genus) is more diverse than the E. chabrieri group, which is distributed over the entire range.     

A taxonomic review of the catfish identified as Glyptothorax zanaensis (Teleostei: Siluriformes: Sisoridae), with the descriptions of two new species

This study re‐examined the taxonomic status of the sisorid catfishes usually identified as Glyptothorax zanaensis using a combination of morphometric and molecular data. Our results resurrect Glyptothorax longinema from the synonymy of G. zanaensis, and we describe two previously unnamed species as Glyptothorax granosus sp. nov. and Glyptothorax fucatus sp. nov. All four species are diagnosed and described in detail. Truss‐based morphometrics combined with principal component analysis (PCA) detected three principal components (PCs) that can explain 86% of the total variation amongst species, which mainly reflect the characteristics of body depth, related depth, adhesive apparatus length, pectoral‐fin length, caudal peduncle length, and barbel lengths. We also generated a phylogenetic hypothesis of these species using concatenated mitochondrial cytochrome b and d ‐loop gene sequences. Molecular dating analysis revealed a rapid speciation of Glyptothorax in the south‐eastern corner of the Qinghai‐Xizang Plateau from the middle Pliocene to early Pleistocene. A key to identify the Glyptothorax species from the Salween River drainage is also provided. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 165 , 363–389.     

Uplift-driven diversification revealed by the historical biogeography of the cockroach Cryptocercus Scudder (Blattodea: Cryptocercidae) in eastern Asia

Cryptocercus Scudder is a genus of wingless cockroaches, which spend their lives feeding within rotting wood in old-growth montane forests. Their dispersal capability is likely to be limited because they depend on the succession of temperate forests, but their distribution exhibits intercontinental disjunctions. Although the natural history and conservation biology of the North American species of the genus have received much attention, the evolution of the eastern Asian lineages remains enigmatic. To resolve the geographic patterns and evolutionary history of Cryptocercus in eastern Asia, we analysed genetic data of species from China (27 species), Korea (one species) and the Russian Far East (one species). We performed phylogenetic analyses of seven genes to infer the evolutionary relationships among species of Asian Cryptocercus. We then used Bayesian molecular dating to estimate the evolutionary timescale of the genus. Ancestral geographic distributions were reconstructed in rasp using statistical dispersal-vicariance analysis and statistical dispersal-extinction-cladogenesis. Our maximum-likelihood and Bayesian phylogenetic trees supported four major lineages of Cryptocercus, revealing a clear geographical pattern. The divergence of American and Asian lineages was inferred to have occurred 74.8 Ma (95% credibility interval: 51.1–103.8 Ma), with the diversification of Asian taxa beginning at 30.7 Ma (95% credibility interval: 22.9–40.3 Ma). The most recent common ancestor of Asian Cryptocercus taxa was inferred to have had a broad distribution in Asia. The uplift of the Qinghai-Tibet Plateau in the Palaeogene and Neogene, along with climatic oscillations in the Quaternary, probably had substantial effects on the formation of the disjunction pattern in the Cryptocercus lineages found in the Hengduan Mountains and Qin-Daba Mountains of China. We propose that the distribution of Cryptocercus has been strongly influenced by habitat fragmentation and subsequent allopatric speciation.     

Allozyme diversity and the evolution ofCrepidiastrum (Compositae) on the Bonin (Ogasawara) Islands

The genusCrepidiastrum is distributed in East Asia and includes 7 species. In the Bonin Islands, three species ofCrepidiastrum occur, and all of them are endemic to the islands. For detecting the origin and speciation of these endemic species, electrophoretic studies have been done in three endemic species of the Bonin Islands as well as in the remaining four species ofCrepidiastrum, andYoungia denticulata which is considered to be closely related toCrepidiastrum. A total of 386 individuals were sampled from 14 populations. As a result, 17 loci of 10 enzyme systems were resolved and gene frequencies for each population were calculated. The genetic variability was low in island species, as reported in some oceanic island plants. Four groups were recognized in the dendrogram generated by the UPGMA method. The Bonin endemics were clustered together, suggesting a monophyletic origin.C. ameristophyllum andC. linguaefolium were found to be genetically very similar, and this may suggest recent and rapid speciation within the islands.     

Multilocus analysis of nucleotide variation and speciation in three closely related Populus (Salicaceae) species

Historical tectonism and climate oscillations can isolate and contract the geographical distributions of many plant species, and they are even known to trigger species divergence and ultimately speciation. Here, we estimated the nucleotide variation and speciation in three closely related Populus species, Populus tremuloides, P. tremula and P. davidiana, distributed in North America and Eurasia. We analysed the sequence variation in six single‐copy nuclear loci and three chloroplast (cpDNA) fragments in 497 individuals sampled from 33 populations of these three species across their geographic distributions. These three Populus species harboured relatively high levels of nucleotide diversity and showed high levels of nucleotide differentiation. Phylogenetic analysis revealed that P. tremuloides diverged earlier than the other two species. The cpDNA haplotype network result clearly illustrated the dispersal route from North America to eastern Asia and then into Europe. Molecular dating results confirmed that the divergence of these three species coincided with the sundering of the Bering land bridge in the late Miocene and a rapid uplift of the Qinghai‐Tibetan Plateau around the Miocene/Pliocene boundary. Vicariance‐driven successful allopatric speciation resulting from historical tectonism and climate oscillations most likely played roles in the formation of the disjunct distributions and divergence of these three Populus species.     

ITS sequences and speciation on far easternIndigofera (Leguminosae)

The internal transcribed spacer (ITS) regions of 18–26S nuclear ribosomal DNA was sequenced to address phylogenetic relationships and to measure the extent of differentiation among six species of the Far EasternIndigofera. ITS 1 had 230–240 base pairs (bp) long while ITS 2 had 211–213 bp long. The 5.8S rRNA coding region was 161 bp long. Sequence divergence calculated by Kimura's two parameter method between species ranged from 0.00 to 13.49%. A single most parsimonious tree was produced from 77 variable nucleotide sites, which had a consistency index of 0.97 and a retention index of 0.83. ITS sequence data suggested that the continental species ofI. kirilowii (2n=16) is diverged from the common ancestor of other species at first, and then the island species ofI. decora (2n=48) andI. venulosa and the Korean Peninsular species ofI. grandiflora (2n=16) andI. koreana (2n=32) are diverged from the ancestor. The molecular data supports that the speciation in the Far EasternIndigofera occurred with polyploidization from continental ancestor to peripheral peninsular and island species.     

Monophyly and phylogenetic relationships of the catfish genus Glyptothorax (Teleostei: Sisoridae) inferred from nuclear and mitochondrial gene sequences

GlyptothoraxBlyth (1860) is the most species-diverse and widely-distributed genus in the Sisoridae, but few studies have examined monophyly of the genus and phylogenetic relations within it. We used the nuclear RAG2 gene and mitochondrial COI and Cyt b genes from 50 of the approximately 70 species to examine monophyly of Glyptothorax and phylogenetic relationships within the genus. Molecular phylogenetic trees were constructed using maximum parsimony, maximum likelihood and Bayesian inference methods. All methods strongly supported monophyly of Glyptothorax, with Bagarius as its sister group. Both analyses of two- and three-gene datasets recovered nine major subclades of Glyptothorax, but some internal nodes remained poorly resolved. The phylogenetic relationships within the genus and existing taxonomic problems are discussed.     

Palaeobiogeography of the North Pacific toothed mysticetes (Cetacea,Aetiocetidae): a key to Oligocene cetacean distributional patterns

Biogeographical distributional patterns of cetaceans reflect dispersal events and colonization of the oceans from their ancestral area in the ancient Sea of Tethys ~53 Ma. Likewise, they reveal several vicariance events throughout the evolutionary history of this group. However, our understanding of how these processes took place and what biogeographical scenarios occurred among the different groups of cetaceans through time is limited. Consequently, this work focuses on explaining the distributional patterns of the well‐known North Pacific toothed mysticetes, Aetiocetidae, through the power of retrodiction offered by track analysis (panbiogeography) and cladistic biogeography, using the approach of evolutionary biogeography. Our results show that the distributional patterns of Aetiocetidae explain their endemism in the North Pacific, as well as indicating that their hypothetical ancestor probably colonized the Pacific from the Atlantic Ocean by a dispersal event (founder effect) via the Central American Seaway. Furthermore, their biogeographical history shows that the adaptive radiation (cladogenesis) of Aetiocetidae is result of peripatric speciation followed by sympatric speciation within a heterogeneous environment. Finally, the biogeographical framework of Aetiocetidae further supports the relevant role that the Pacific Ocean has played in the evolution of Oligocene cetaceans as a geographical area that promoted endemism, dispersal and colonization. At more local scales, environmental conditions further promoted increased diversity and disparity amongst Mysticeti.     

Molecular phylogeny and biogeography of the Qinghai-Tibet Plateau endemic Nannoglottis (Asteraceae)

All taxa endemic to the Qinghai-Tibet Plateau are hypothesized to have originated in situ or from immediately adjacent areas because of the relatively recent formation of the plateau since the Pliocene, followed by the large-scaled biota extinction and recession caused by the Quaternary ice sheet. However, identification of specific progenitors remains difficult for some endemics, especially some endemic genera. Nannoglottis, with about eight species endemic to this region, is one such genus. Past taxonomic treatments have suggested its relationships with four different tribes of Asteraceae. We intend to identify the closest relatives of Nannoglottis by evaluating the level of monophyly, tribal delimitation, and systematic position of the genus by using molecular data from ndhF gene, trnL-F, and ITS region sequences. We find that all sampled species of Nannoglottis are a well-defined monophyly. This supports all recent taxonomic treatments of Nannoglottis, in which all sampled species were placed in one broadly re-circumscribed genus. Nannoglottis is most closely related to the Astereae, but stands as an isolated genus as the first diverging lineage of the tribe, without close relatives. A tentative relationship was suggested for Nannoglottis and the next lineage of the tribe was based on the ITS topology, the "basal group," which consists of seven genera from the Southern Hemisphere. Such a relationship is supported by some commonly shared plesiomorphic morphological characters. Despite the very early divergence of Nannoglottis in the Astereae, the tribe must be regarded to have its origin in Southern Hemisphere rather than in Asia, because based on all morphological, molecular, biogeographical, and fossil data, the Asteraceae and its major lineages (tribes) are supposed to have originated in the former area. Long-distance dispersal using Southeast Asia as a steppingstone from Southern Hemisphere to the Qinghai-Tibet Plateau is the most likely explanation for this unusual biogeographic link of Nannoglottis. The 23-32-million-year divergence time between Nannoglottis and the other Astereae estimated by DNA sequences predated the formation of the plateau. This estimation is further favored by the fossil record of the Asteraceae and the possible time of origin of the Astereae. Nannoglottis seems to have reached the Qinghai-Tibet area in the Oligocene-Eocene and then re-diversified with the uplift of the plateau. The molecular infragenetic phylogeny of the genus identifies two distinct clades, which reject the earlier infrageneric classification based on the arrangement of the involucral bracts and the length of the ligules, but agree well with the habits and ecological preferences of its current species. The "alpine shrub" vs. "coniferous forest" divergence within Nannoglottis was estimated at about 3.4 million years ago when the plateau began its first large-scale uplifting and the coniferous vegetation began to appear. Most of the current species at the "coniferous forest" clade of the genus are estimated to have originated from 1.02 to 1.94 million years ago, when the second and third uprisings of the plateau occurred, the climate oscillated and the habitats were strongly changed. The assumed evolution, speciation diversity, and radiation of Nannoglottis based on molecular phylogeny and divergence times agree well with the known geological and paleobotanical histories of the Qinghai-Tibet Plateau.     

Small mammal (rodents and lagomorphs) European biogeography from the Late Oligocene to the mid Pliocene

Aim To analyse the fossil species assemblages of rodents and lagomorphs from the European Neogene in order to assess what factors control small mammal biogeography at a deep‐time evolutionary time‐scale. Location Western Europe: 626 fossil‐bearing localities located within 31 regions and distributed among 18 successive biochronological units ranging from c. 27 Ma (million years ago; Late Oligocene) to c. 3 Ma (mid Pliocene). Methods Taxonomically homogenized pooled regional assemblages are compared using the Raup and Crick index of faunal similarity; then, the inferred similarity matrices are visualized as neighbour‐joining trees and by projecting the statistically significant interregional similarities and dissimilarities onto palaeogeographical maps. The inferred biogeographical patterns are analysed and discussed in the light of known palaeogeographical and palaeoclimatic events. Results Successive time intervals with distinct biogeographical contexts are identified. Prior to c. 18 Ma (Late Oligocene and Early Miocene), a relative faunal homogeneity (high interregional connectivity) is observed all over Europe, a time when major geographical barriers and a weak climatic gradient are known. Then, from the beginning of the Middle Miocene onwards, the biogeography is marked by a significant decrease in interregional faunal affinities which matches a drastic global climatic degradation and leads, in the Late Miocene (c. 11 Ma), to a marked latitudinal pattern of small mammal distribution. In spite of a short rehomogenization around the Miocene/Pliocene boundary (6–4 Ma), the biogeography of small mammals in the mid Pliocene (c. 3 Ma) finally closely reflects the extant situation. Main conclusions The resulting biogeographical evolutionary scheme indicates that the extant endemic situation has deep historical roots corresponding to global tectonic and climatic events acting as primary drivers of long‐term changes. The correlation of biogeographical events with climatic changes emphasizes the prevalent role of the climate over geography in generating heterogeneous biogeographical patterns at the continental scale.     

Are the Northern Andes a species pump for Neotropical birds? Phylogenetics and biogeography of a clade of Neotropical tanagers (Aves: Thraupini)

Aim We used mitochondrial DNA sequence data to reconstruct the phylogeny of a large clade of tanagers (Aves: Thraupini). We used the phylogeny of this Neotropical bird group to identify areas of vicariance, reconstruct ancestral zoogeographical areas and elevational distributions, and to investigate the correspondence of geological events to speciation events. Location The species investigated are found in 18 of the 22 zoogeographical regions of South America, Central America and the Caribbean islands; therefore, we were able to use the phylogeny to address the biogeographical history of the entire region. Methods Molecular sequence data were gathered from two mitochondrial markers (cytochrome b and ND2) and analysed using Bayesian and maximum‐likelihood approaches. Dispersal–vicariance analysis (DIVA) was used to reconstruct zoogeographical areas and elevational distributions. A Bayesian framework was also used to address changes in elevation during the evolutionary history of the group. Results Our phylogeny was similar to previous tanager phylogenies constructed using fewer species; however, we identified three genera that are not monophyletic and uncovered high levels of sequence divergence within some species. DIVA identified early diverging nodes as having a Northern Andean distribution, and the most recent common ancestor of the species included in this study occurred at high elevations. Most speciation events occurred either within highland areas or within lowland areas, with few exchanges occurring between the highlands and lowlands. The Northern Andes has been a source for lineages in other regions, with more dispersals out of this area relative to dispersals into this area. Most of the dispersals out of the Northern Andes were dispersals into the Central Andes; however, a few key dispersal events were identified out of the Andes and into other zoogeographical regions. Main conclusions The timing of diversification of these tanagers correlates well with the main uplift of the Northern Andes, with the highest rate of speciation occurring during this timeframe. Central American tanagers included in this study originated from South American lineages, and the timing of their dispersal into Central America coincides with or post‐dates the completion of the Panamanian isthmus.     

Plio-Pleistocene climatic oscilations, Holarctic biogeography and speciation in an avian subfamily

Aim In this paper, I discuss the temporal and spatial aspects of historical biogeography and speciation in a widely distributed Holarctic subfamily of birds (Tetraoninae). Location Northern Holarctic. Results Using dated fossils, I calibrated the molecular clock for the mitochondrial control region at 7.23 ± 1.58% nucleotide divergence (maximum likelihood corrected) per million years. The data suggest that grouse (Tetraoninae) originated in the Middle Pliocene, 6.3 Ma. Grouse apparently originated in the northern part of western Nearctic, and Palearctic was colonized independently three times, first by the ancestor of all grouse in the Middle Pliocene, then by the ancestor of forest (Falcipennis, Tetrao and Lyrurus) and prairie (Centrocercus, Dendragapus and Tympanuchus) grouse in the Late Pliocene, and finally by the ancestral Lagopus in the Early Pleistocene. Only once Nearctic was colonized from Palearctic by a common ancestor of forest grouse. Sympatry and range symmetry were positively correlated with molecular divergence. These correlations suggest that peripatric isolation was the predominant mode of speciation throughout grouse history. Main conclusions Speciation events in grouse were driven by climatic oscillations of the Pliocene and Pleistocene. Isolation of small peripheral populations from widely distributed ancestors was the dominant mode of speciation in grouse. Isolations during interglacials both across Beringia, and in southern mountain areas when boreal habitats were restricted to high elevations, suggest an important role for vicariance in grouse speciation.     

中国狼不同地理种群遗传多样性及系统进化分析

为了解中国狼不同地理种群遗传多样性及系统发育情况,从中国境内狼的主要分布区青海、新疆、内蒙古和吉林4个地区采集样品,用分子生物学技术手段成功地获得44个个体线粒体DNA控制区第一高变区(HVRⅠ)序列和40个线粒体Cyt b部分序列。线粒体控制区HVRⅠ共检测到51个变异位点,位点变异率为8.76%;线粒体Cyt b部分序列发现31个变异位点,位点变异率为5.33%,未见插入及缺失现象,变异类型全部为碱基置换。共定义了16个线粒体HVRⅠ单倍型,其中吉林与内蒙种群存在共享单倍型,估计这两地间种群亲缘关系较近。4个地理种群中新疆种群拥有较高的遗传多样性(0.94)。中国狼种群总体平均核苷酸多态性为2.27%,与世界其他国家地区相比,中国狼种群拥有相对较高的遗传多样性。通过线粒体HVRⅠ单倍型构建的系统进化树可以看出,中国狼在进化上分为2大支,其中位于青藏高原的青海种群独立为一支,推测其可能长期作为独立种群进化。基于青海种群与新疆,内蒙种群的线粒体Cyt b遗传距离,推测中国狼2个世系可能在更新世冰川时期青藏高原受地质作用急速隆起后出现分歧,分歧时间大约在1.1 MY前。