mtDNA analysis reveals the ongoing speciation on Greek populations of Microtus (Terricola) thomasi (Arvicolidae,Rodentia)

The present article extends our previous work on the phylogenetic history of Microtus (Terricola) thomasi, analysing cytochrome b, 12S rRNA, 16S rRNA and the control region in 65 Greek populations. The analysis revealed three clades: one grouping the populations from Peloponnisos (Southern Greece); the second, the populations from Agios (Ag.) Stefanos and Evvoia island (Central East Greece); and the third, all the remaining populations with no geographical substructure. Genetic distances were low for most populations, with only the populations of Evvoia and Ag. Stefanos being substantially distant. Thus, although this species has a recent colonization history and probably descends from a highly polymorphic ancestor, a monophyletic and highly differentiated lineage is formed in Greece and is distributed in Ag. Stefanos and Evvoia. Molecular differentiation, distinct geographical distribution and restriction of gene flow between this lineage and the rest of the Greek populations provide evidence for its probable subspecific status, Microtus (Tericola) thomasi atticus. A possible mechanism leading the differentiation process of the proposed subspecies is suggested, based on the displacement of this species in central Greece by its congeneric, probably better‐fitted Microtus (Microtus) guentheri and the subsequent separation of Ag. Stefanos and Evvoia from the remaining Greek populations. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 117–130.     

Comparative cytogenetic study of two sister species of Iberian ground voles, Microtus (Terricola) duodecimcostatus and M. (T.) lusitanicus (rodentia, cricetidae)

The two Iberian species of pine voles, Microtus (Terricola) duodecimcostatus and M. (T.) lusitanicus of the subfamily Arvicolinae (Cricetidae, Rodentia), were compared after G- and C-banding and chromosomal mapping of ribosomal RNA genes (rDNA), telomeric repeats, and satellite DNA Msat-160. Notwithstanding their close relationship (one sister group in phylogenetic analyses) and sharing of the diploid and fundamental chromosome numbers, the 2 species show notable differences in the sex chromosome morphology, the number and distribution of rDNA sites, constitutive heterochromatin and satDNA patterns. The only telomeric repeats showed normal, all-telomeric, distribution in karyotypes of both species. The data are discussed with regard to interspecific and intrageneric variation of the analyzed characters and the chromosomal evolution in the genus Microtus.     

Evolutionary history of two allopatric Terricola species (Arvicolinae, Rodentia) from molecular, morphological, and palaeontological data

To investigate the phylogenetic and phylogeographical relationships of arvicolines, we use several Western European ground voles. More particularly, our study is focused on Microtus ( Terricola ) savii and M. ( T. ) pyrenaicus . These two allopatric species are usually considered as having originated from the same ancestor, possibly M . ( T. ) mariaclaudiae . We propose molecular and morphological approaches: nucleotidic data from the mitochondrial cytochrome b and 12S rRNA genes and global morphological analyses from the first lower molar. Four other Terricola species ( multiplex , lusitanicus , duodecimcostatus , subterraneus ) were added to the data set for both analyses, and two other vole species ( Clethrionomys glareolus and Chionomys nivalis ) as outgroup to the molecular analysis, and five fossil populations to the morphological one. Palaeontological data are also widely taken into account. Both molecular and morphological analyses indicate that intra- Terricola relationships reflect the present-day geographical distribution of our data set species. Our results show that M. ( T. ) savii and M. ( T. ) pyrenaicus are from separate speciation events leading to two different biogeographical groups, respectively the Alpine–Italian group and the French–Iberian group, the latter being much more homogeneous. These speciation events could be related to Quaternary climatic changes, which induced southward migration, leading first to M. ( T. ) savii and second to M. ( T. ) pyrenaicus . The classical hypothesis of a geographical speciation for these two taxa from M. ( T. ) mariaclaudiae is invalid. However, the morphological data suggest a potential phylogenetic relationship between M. ( T. ) mariaclaudiae (ancestor) and M. ( T. ) pyrenaicus (descendant).  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 309–323.     

Molecular phylogeny of the speciose vole genus Microtus (Arvicolinae, Rodentia) inferred from mitochondrial DNA sequences

Voles of the genus Microtus represent one of the most speciose mammalian genera in the Holarctic. We established a molecular phylogeny for Microtus to resolve contentious issues of systematic relationships and evolutionary history in this genus. A total of 81 specimens representing ten Microtus species endemic to Europe as well as eight Eurasian, six Asian and one Holarctic species were sequenced for the entire cytochrome b gene (1140 bp). A further 25 sequences were retrieved from GenBank, providing data on an additional 23, mainly Nearctic, Microtus species. Phylogenetic analysis of these 48 species generated four well-supported monophyletic lineages. The genus Chionomys, snow voles, formed a distinct and well-supported lineage separate from the genus Microtus. The subgenus Microtus formed the strongest supported lineage with two sublineages displaying a close relationship between the arvalis species group (common voles) and the socialis species group (social voles). Monophyly of the Palearctic pitymyid voles, subgenus Terricola, was supported, and this subgenus was also subdivided into two monophyletic species groups. Together, these groupings clarify long-standing taxonomic uncertainties in Microtus. In addition, the "Asian" and the Nearctic lineages reported previously were identified although the latter group was not supported. However, relationships among the main Microtus branches were not resolved, suggesting a rapid and potentially simultaneous radiation of a widespread ancestor early in the history of the genus. This and subsequent radiations discernible in the cytochrome b phylogeny, show the considerable potential of Microtus for analysis of historical and ecological determinants of speciation in small mammals. It is evident that speciation is an ongoing process in the genus and that the molecular data provides a vital insight into current species limits as well as cladogenic events of the past.     

Biochronological implications of the Arvicolidae (Rodentia, Mammalia) from the Lower Pleistocene hominid-bearing level of Trinchera Dolina 6 (TD6, Atapuerca, Spain).

Level TD6 of the Trinchera Dolina Section in the railway cutting of the Sierre de Atapuerca (Trinchera del Ferrocarril) has yielded a rich small mammal assemblage (26 species) in association with fossil human remains of Homo antecessor. The arvicolids of TD6 are identified as: Mimomys savini, Microtus seseae, Stenocranius gregaloides, Terricola arvalidens, Iberomys huescarensis, Allophaiomys chalinei, and Pliomys episcopalis. The rodent association also includes large rodents (i.e., Castor fiber, Marmota sp., and Hystrix refossa) and the small Allocricetus sp., Eliomys helleri, Micromys minutus, and Apodemus aff. flavicollis. The small vertebrate remains also include Insectivora (Beremendia fissidens, Sorex sp, Neomys sp., Crocidura sp., Galemys sp., Talpa sp., Erinaceus sp.), Chiroptera (Miniopterus schreibersii, Myotis sp., Rhinolophus sp.), and Lagomorpha (Oryctolagus sp., Lepus sp.), as well as lizards, birds and amphibians. The H. antecessor remains are derived from a 15 cm thick layer at the top of TD6 (TD6-T36-43), where A. chalinei, H. refossa and Marmota sp. do not occur. The paleomagnetic Matuyama/Brunhes boundary is found in the overlying level TD7 of the Gran Dolina Section. On the basis of the arvicolids, TD6 can be referred to the Biharian biochron. The Matuyama/Brunhes boundary is fixed in the late Biharian (Microtus-Mimomys rodent Superzone). The species M. savini (without M. pusillus), as well as the evolutionary stage of Microtus s.l., are characteristic of the Late Biharian. The evolutionary level of the species M. savini, T. arvalidens, S. gregaloides indicates that TD6 is older than West Runton (type Cromerian). In the Trinchera Dolina Section we are able to calibrate, for the first time, the evolutionary level of important biochronological markers with magnetostratigraphy. We propose that a radiation of Microtus s.l., along with the first appearance of primitive S. gregaloides, T. arvalidens and Iberomys, took place just before the Matuyama/Brunhes boundary. These species can be considered as characteristic elements of early Pleistocene faunas.     

长江三角洲东缘晚新近纪沉积的孢粉与古环境研究

作者对长江三角洲东缘地区的南汇鹤鸣孔(Hm)、东海1井、高桥G2孔3口钻井晚新近纪地层的孢粉作了研究和对比,划分出了6个孢粉组合带和3个亚带,恢复了本地区植被演替、气候波动的6个阶段:第1阶段为稀疏的针叶、落叶阔叶混交林,反映气候凉冷稍湿;第2阶段为针叶阔叶混交林草地,反映气候温凉略湿;第3阶段为含常绿阔叶树的针叶阔叶混交林,反映气候温和略干;第4阶段为以常绿栎类、栲属、杨梅等为主的常绿阔叶林,反映气候热暖潮湿;第5阶段是以栎、松、禾本科为主的针叶、阔叶混交林,反映气候温暖略干;第6阶段是以落叶栎类、常绿栎类、松为主的落叶阔叶、常绿阔叶、针叶混交林草地,反映气候温暖湿润。这种气候波动与世界性气候变化相一致,为本地区的地层年代划分和对比提供了新的证据,为晚新近纪古植被、古气候、古环境的重建提供了重要的孢粉学资料。     

The Pleistocene subterranean volesTerricola (Rodentia) of Serbia and Montenegro

A general morphometrical analysis of the M₁ was conducted to identify the subterranean vole species found in Upper Pleistocene localities from Serbia and Montenegro, and to clarify the systematic position and the phylogenetic relationships between the different species in the Balkans. From the different localities studied, we can assign one population toMicrotus (Terricola) thomasi and the others to theM. (T.) subterraneus group. This study suggests thatM. (T.) grafi can be considered as a chronological sub-species ofM. (T.) subterraneus or as a different but phylogenetically very close species.     

Short-term climate and vegetation dynamics in Lena River Delta (northern Yakutia,Eastern Siberia) during early Eocene

Early Eocene climate and vegetation evolution in northern Yakutia (Eastern Siberia) are quantitatively studied for the first time based on paleobotanical records, using the coexistence approach (CA) for paleoclimate, the plant functional type approach (PFT) and the integrated plant record method (IPR). Paleobotanical data of this time-interval were obtained from 18 palynofloras of the coal beds outcropping on the bank of the Bykovskaya Channel in the area of Bulunkan Bay (Lena River Delta). The results indicate the persistence of warm temperate and at times possibly nearly tropical, overall humid climate conditions throughout the early Eocene, and a relatively strong seasonality in precipitation. The vegetation reconstructed is in agreement with the climate conditions determined by the CA. Analyses of the PFT diversity spectra indicate the prevalence of mixed warm temperate evergreen-deciduous forests. Based on the IPR method, plant biomes varied from mixed mesophytic forests during warmer intervals to broadleaved deciduous forest/mixed mesophytic forest ecotone during cooler episodes. The presence of mangroves in our early Eocene records is possibly related to hyperthermal events such as the Paleocene–Eocene Thermal Maximum.     

Molecular phylogeny of land and freshwater planarians (Tricladida, Platyhelminthes): from freshwater to land and back

The suborder Tricladida (phylum Platyhelminthes) comprises the well-known free-living flatworms, taxonomically grouped into three infraorders according to their ecology: Maricola (marine planarians), Paludicola (freshwater planarians), and Terricola (land planarians). Molecular analyses have demonstrated that the Paludicola are paraphyletic, the Terricola being the sister group of one of the three paludicolan families, the Dugesiidae. However, neither 18S rDNA nor COI based trees have been able to resolve the relationships among species of Terricola and Dugesiidae, particularly the monophyly of Terricola. Here, we present new molecular data including sequences of nuclear genes (18S rDNA, 28S rDNA) and a mitochondrial gene (COI) of a wider sample of dugesiid and terricolan species. The new sequences have been analyzed, together with those previously obtained, in independent and concatenated analyses using maximum likelihood and bayesian methods. The results show that, although some parts of the trees remain poorly resolved, they support a monophyletic origin for Terricola followed by a likely return of some species to freshwater habitats. Relationships within the monophyletic group of Dugesiidae are clearly resolved, and relationships among some terricolan subfamilies are also clearly established and point to the need for a thorough revision of Terricola taxonomy.     

Late Pleistocene small mammals from the Wannenköpfe volcanoes (Neuwied Basin, western Germany) with remarks on the stratigraphic range of Arvicola terrestris

Redeposited loess sediments from a Pleistocene crater filling in the Neuwied Basin (western Germany) yielded a small mammal fauna mainly composed of arvicoline rodents. The occurrence of rare faunal elements such as Sorex minutissimus, Sicista subtilis, and a species of Microtus (Terricola) is remarkable. Based on the evolutionary level of Arvicola terrestris, the biostratigraphic age of the fossiliferous horizon is transitional between late Eemian and early Weichselian. This conclusion is partly supported by the generalized ecological requirements of a few dominant rodent species. The fauna contributes to the knowledge of this time period, which is poorly documented in western Germany. A new character of the m1 is proposed for the discrimination of Sicista betulina and S. subtilis. To avoid further confusion in the specific assignment of transitional Arvicola specimens from the late Middle Pleistocene to early Late Pleistocene, we propose the formal taxonomic boundary between chronospecies A. mosbachensis and A. terrestris to be shifted from SDQ value 100 to SDQ value 120.     

A robust molecular phylogeny of the Tricladida (Platyhelminthes: Seriata) with a discussion on morphological synapomorphies.

The suborder Tricladida (Platyhelminthes: Turbellaria, Seriata) comprises most well-known species of free-living flatworms. Four infraorders are recognized: (i) the Maricola (marine planarians); (ii) the Cavernicola (a group of primarily cavernicolan planarians); (iii) the Paludicola (freshwater planarians); and (iv) the Terricola (land planarians). The phylogenetic relationships among these infraorders have been analysed using morphological characters, but they remain uncertain. Here we analyse the phylogeny and classification of the Tricladida, with additional, independent, molecular data from complete sequences of 18S rDNA and 18S rRNA. We use maximum parsimony and neighbour-joining methods and the characterization of a unique gene duplication event involving the Terricola and the dugesiids to reconstruct the phylogeny. The results show that the Maricola is monophyletic and is the primitive sister group to the rest of the Tricladida (the Paludicola plus the Terricola). The Paludicola are paraphyletic since the Terricola and one paludicolan family, the Dugesiidae, share a more recent common ancestor than the dugesiids with other paludicolans (dendrocoelids and planariids). A reassessment of morphological evidence may confirm the apparent redundancy of the existing infraorders Paludicola and Terricola. In the meantime, we suggest replacing the Paludicola and Terricola with a new clade, the Continenticola, which comprises the families Dugesiidae, Planariidae, Dendrocoelidae and the Terricola.     

mtDNA variation and evaluation of phylogenetic relationships among karyotypically polymorphic populations of Microtus (Terricola) thomasi (Arvicolidae, Rodentia) from Greece

The Balkan endemic species Microtus (Terricola) thomasi – Thomas' vole – is widely distributed in the Greek mainland and presents regionally confined karyotypic forms. In order to test the significance of geographical isolation and/or karyological differences in microevolutionary processes at the intraspecific level we used restriction enzymes and partial sequencing to study the polymorphism of the mtDNA d -loop control region in ten Greek populations. An overall high rate of both inter- and intrapopulation polymorphism was observed. The mtDNA diversity was unevenly distributed so that most of the populations studied were very similar and closely related, and only a few of them were clearly differentiated from the others. The mtDNA analysis failed to reveal any clear variation pattern that could be correlated with karyotypic forms or geographical proximity of the populations studied. Instead, the mtDNA phylogeny gave evidence in support of the coexistence of non-polymorphic (2 n  = 44) and polymorphic (2 n  < 44) karyotypic forms within the ancestral Greek M. (T.) thomasi stock. Interestingly, only one population (Agios Stefanos) seemed to significantly diverge from all the others in both methods used, forming a well-supported paraphyletic clade. Allozymic studies have already reported similar results, corroborating the genetic peculiarity of this population.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 55–68.     

Microevolutionary relationships between phylogeographical history,climate change and morphological variability in the common vole (Microtus arvalis) across France

Aim In this study, we analyse microevolutionary processes in common voles (Microtus arvalis) through the investigation of tooth morphological structure, in order to assess the relative impact of climate and phylogeographical history. Microevolutionary studies have shown that climate change may play a role in both population phylogeography and phenotypic differentiation. However, relatively little is known about the precise relationship between phylogeography and phenotypic variability and about how organisms respond to climate change. Location France, from sea level to the Alps (5 to > 2300 m a.s.l.). Methods This morphological analysis is based on first lower molar measurements from 16 geographically distinct common vole populations. Size and shape components are assessed separately. Population structure patterns are characterized using canonical variate analysis. We use phylogenetic analyses of two regions of mitochondrial DNA (the control region and the cytochrome b gene) to infer genetic structure. We calculate climate parameters from temperature and precipitation data. We investigate the influence of climate, geography and phylogeographical history on the phenotype using (1) multiple regression tests, (2) pairwise comparison of observation‐by‐variable matrices, and (3) a correlation method designed to compare three matrices. Results All populations were clearly structured, whatever the dataset. Neither size nor shape variation was correlated with climate parameters, but tooth shape was strongly correlated with both genetic structure and geographical distance. Main conclusions In French Microtus arvalis populations, molar shape differentiation is clearly associated with both phylogeographical history and geographical distance. Population phylogeographical history has a greater relevance than climate in accounting for variation in tooth morphology.     

Palynological evidence for Holocene environmental change in the Changjiang (Yangtze River) Delta, China

Holocene vegetation changes in response to climate fluctuations and human impacts are reviewed on the basis of pollen analyses from borehole cores taken from the Changjiang (Yangtze River) delta, China, and other previously reported data. During the earliest Holocene (10,930−9000 cal yr BP), the climate was warm and wet, allowing thermophilous hardwoods to occupy mid- and low-elevations surrounding a palaeo-Changjiang estuary. The climate became gradually cooler, and cool-tolerant conifers, grasses and ferns became dominant until 7600 cal yr BP, when the estuary became a delta. A mid-Holocene climatic optimum occurred between 7600 and 4800 cal yr BP, when evergreen and broadleaved deciduous trees flourished at mid- and low-elevations surrounding the delta front-prodelta. After this time, climate became cooler again until 1340 cal yr BP. During this period, evergreen and broadleaved deciduous trees were replaced by conifers and grasses inhabiting the inter/subtidal flat-delta front. This development of conifer-grassland vegetation was shortly interrupted between 3860 and 3200 cal yr BP when thermophilous tree cover increased and open vegetation with scattered conifers was reduced. Since 1340 cal yr BP, the vegetation has been similar to that at present under warm, wet conditions. Human impacts are recognized by the first appearance of Fagopyrum and a sudden increase in herb pollen at 4500 and 1340 cal yr BP, respectively.     

PHOTOSYNTHETIC SENSITIVITY TO TEMPERATURE IN POPULATIONS OF TWO C4 BOUTELOUA (POACEAE) SPECIES NATIVE TO DIFFERENT ALTITUDES

The relationships between photosynthesis, flowering, and growth temperatures were examined experimentally in four populations of the C₄ grass genus Bouteloua. Field-collected plants were grown under two temperature regimes, cool (20 C day/6 C night) and warm (30/16), representative of the extreme populations. Populations collected from the warm climates had significantly lower photosynthetic capacity when grown in the cool chamber relative to the warm chamber, while photosynthetic capacity in the cool climate populations did not differ between the growth conditions. Additionally, exposure to a 2-day cold temperature treatment (10/-2), representative of late-season frosts in high altitude sites, resulted in further reductions in photosynthesis in the warm climate plants, but not in the cool climate plants. This effect was greater for plants grown in the cool growth chamber. Flowering was reduced by 70% in the warm climate plants grown in the cool chamber, and was correlated with photosynthetic inhibition following the short-term cold temperature treatment. These results indicate that genetic differentiation for photosynthetic temperature sensitivity has occurred in the cool climate populations, and that long-term exposure to cool temperatures coupled with short-term relatively extreme low temperatures results in greater photosynthetic inhibition in nontolerant populations.     

First report of Siphonaptera infesting Microtus (Microtus) cabrerae (Rodentia-Muridae-Arvicolinae) in Cuenca,Spain and notes about the morphologic variability of Ctenophthalmus (Ctenophthalmus) apertus personatus (Insecta-Siphonaptera-Ctenophthalmidae)

The fleas infesting Microtus (Microtus) cabrerae from three different areas of Cuenca province (Spain) have been studied. It is the first time that on ectoparasitological study of this badly known rodent has been done. Four Siphonaptera species have been detected: Rhadinopsylla (Actenophthalmus) pentacantha, Peromyscopsylla spectabilis spectabilis, Nosopsyllus fasciatus and Ctenophthalmus (Ctenophthalmus) apertus personatus, which was the most abundant species (26 males and 31 females of a total of 28 males and 35 females). Considering the great morphologic variability within the male processus basimerus ventralis (p.b.v.) of segment IX of C. personatus subspecies, three morphotypes have been recognised. The male polymorphism detected, would be the result of both host confinement and genetic selection acting on the parasite. It should be pointed out that C. (C.) apertus personatus is not narrowly host-specific, therefore further studies are required to clarify this taxonomic situation.     

印度库茂恩喜马拉雅纳尼塔尔温带地区晚全新世古植被和古气候

根据从印度库茂恩（Kumaun)喜马拉雅纳尼塔尔（Naini Tal)温带地区获得的85cm钻井的孢粉资料,恢复了1720＋／－130年以来晚全新世古植被演变史,并推测了当地的古气候,当前孢粉资料说明了该地区当时发育有栎树混交林和波动的温暖润气候,孢粉组合序列的初始阶段,森林距沉积盆地较远,但是,由于气候变得更加温暖潮湿,森林也很快茂密起来,约1200年前,因气候恶化,森林变得稀疏,而在约124年前,气候再次转好,麦类及其他作物花粉资料证明农耕活动大约开始于1460年前左右。     

Islands and the evolution of Microtus arvalis (Microtinae)

The widespread European vole species Microtus arvalis (Pallas) occurs in the British Isles only on Guernsey and various of the Orkney islands. The traditional explanation for this peculiar distribution has been that the island races are the sole relicts of a previously cosmopolitan form. This idea is not supported by the present study, in which population samples are compared with voles from a number of localities from Continental Europe. Using the frequencies of non-metrical skull variants as genetical markers, it emerged that although the Guernsey population ( Microtus arvalis sarnius ) was virtually indistinguishable from typical German specimens, and therefore likely to be a relict from the time when Guernsey was connected to France, the Orkney populations ( Microtus arvalis orcadensis ) formed a distinct group with affinities more to south Europe than elsewhere. It seems probable that arvalis was brought to Orkney by some of the earliest human colonizers about 4000 years ago, and then radiated to the different islands. It is argued that this manner of peripheral differentiation may be an important method of evolutionary change.     

Comparative odontometrical analysis of the first lower molar in <Emphasis Type="Italic">Microtus (Terricola) multiplex</Emphasis> (Arvicolinae,Rodentia) from western Alps

The morphology of the first lower molar (M₁) of Microtus (Terricola) multiplex (Fatio, 1905) was compared amongst 15 populations from the Alps (Switzerland, Italy, France). M. multiplex orientalis from Trentino Alto Adige is close to the nominative subspecies M. multiplex multiplex from Ticino characterised by a great size, a not tilted pitymyan rhombus and an important development of the anterior part of the M₁. M. multiplex druentius from Ubaye mainly differs from the nominative subspecies by a smaller tooth size. Populations from Valle d’Aosta and Piemonte show on the whole a morphology intermediate between M. m. multiplex and M. m. druentius subspecies, however, the pitymyan rhombus is more tilted and the development of the anterior part more reduced in populations from Eastern and Central Piemonte. The Western populations (from Trièvès, Vercors, Royans and Chambaran) belonging to the subspecies M. m. niethammeri are the most differentiated with a small or median size of the M1, a reduced development of the anterior part and a very tilted pitymyan rhombus, particularly in the population from Chambaran. The populations from Matheysine and Grésivaudan are morphologically a link between M. m. druentius and M. m. niethammeri subspecies.