73 resultados para Lower Middle Pleistocene

em Université de Lausanne, Switzerland


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Abstact : Pollinic contribution to a Pleistocene study of the Lake Geneva region (Switzerland) Palaeoenvironment of the last 800,000 years. Although the "Plateau Romand" (tableland of French-speaking Switzerland) has been marked by the repetitive action of different Quaternary glacial thrusts, numerous preserved outcrops here and there, thanks to favourable topographical conditions, provide sufficient pollinic content to reconstitute the significant botanical fluctuations and allow bio- and chrono-stratigraphical correlations with the long European sequences. The present study examines around ten Pleistocene deposits rich in pollens, whose presumed ages cover a range of 800,000 years. - The Early and Early-Middle Pleistocene are examined at Ecoteaux (VD), undeniably the oldest site in this study. The "Lower Formation of Ecoteaux", whose low pollinic content reveals an environment of the cold desert type, shows an inverse palaeomagnetic remanence. An age earlier than 780,000 years BP (Matuyama period or earlier) is proposed. The "Upper Formation of Ecoteaux" contains over eight major fluctuations in the vegetal covering, of which at least four temporal phases and one climatic optimum. The presence of botanical genera, relics of the Tertiary in particular Pterocarya and Carya, associated with the vegetal dynamic made up of short oscillations, contrives to link this formation to the Cromerian Complex. The thermomeres recorded could correspond to one or several interglacial periods from Middle Pleistocene. Possible correlations with marine isotopic stages 15, 17 and 19 are discussed. - The Middle and Late Pleistocene is approached through the sediments of Onnens (VD), Creux d'Enfer (FR), Port-Valais (VS), La Dénériaz (VD) and Cortaillod (NE). Each site provides a temperate interglacial flora, whose pollinic content is evaluated according to the different conceivable bio-stratigraphic correlations. - Finally the Würm is studied thanks to the outcrop of Marly (FR), together with the excavations at Villars-sous-Yens (VD), Versoix (GE) and Ollon (VD). Together, these observations tend to show that the west of the Swiss Plateau is a zone of transition between the meridian vegetation of the lower Rhone valley and that of beyond the Jura. At the outlet of the Rhone glacier, and on the borders of the prealpine and Jurassian glaciers, the region is situated at the crossroads of post-glacial botanical migrations, between the major axis of the Rhone valley and the Eastern refuge zones; this situation confers particular importance on it. Although believed to be deeply scraped by ice, one notices today that the "Plateau Romand" harbours overtwenty interglacial and interstadial sites. The abrasive action of the Rhone glacier has thus been more modest than previously estimated. The effective extension of glacial tongues, together with atmospheric circulation, are doubtless varying from one interglacial to another. Finally, the speeds of glacial movements, and the unbelievable vegetal vitality, are important parameters, capable of leading to deposits, very early enriched in pollens. RÉSUMÉS : Contribution pollinique à l'étude du Pléistocène de la région lémanique (Suisse) - Paléoenvironnement des derniers 800'000 ans Bien que le Plateau romand soit marqué par l'action répétitive des différentes poussées glaciaires quaternaires, de nombreux affleurements préservés ça et là, à la faveur de conditions topographiques favorables, livrent un contenu pollinique suffisant pour reconstituer des fluctuations botaniques significatives et autoriser des corrélations bioet chronostratigraphiques avec les longues séquences européennes. Le présent travail étudie une dizaine de dépôts pléistocènes riches en pollens, dont les âges présumés balaient un éventail de plus de 800'000 ans. - Le Pléistocène Ancien et Moyen inférieur, sont examinés à Ecoteaux (VD); indéniablement le site le plus ancien de cette étude. La "Formation inférieure d'Ecoteaux", dont le maigre contenu pollinique traduit un environnement de type désertique froid, présente une rémanence magnétique inverse. Un âge antérieur à 780'000 ans BP (période de Matuyama ou antérieure) est proposé. La "Formation supérieure d'Ecoteaux" comprend plus de huit fluctuations majeures du couvert végétal, parmi lesquelles on compte au moins quatre épisodes tempérés et un optimum climatique. La présence de genres botaniques reliques du Tertiaire, en particulier Pterocarya et Carya, associée à la dynamique végétale faite de courtes oscillations, concourent à lier cette formation au Complexe Cromérien. Les thermomères enregistrées pourraient correspondre à un- ou plusieurs interglaciaires du Pléistocène moyen. Les corrélations possibles avec les stades isotopiques marins, MIS 15, 17 et 19 sont discutées. - Le Pléistocène Moyen et Récent est abordé à travers les sédiments d'Onnens (VD), du Creux d'Enfer (FR), de Port-Valais (VS), de La Dénériaz (VD) et de Cortaillod (NE). Chaque site livre une flore tempérée interglaciaire, dont le contenu pollinique est évalué en fonction des différentes corrélations bio-stratigraphiques envisageables. - Enfin le Würm est étudié grâce à l'affleurement de Marly (FR), accompagné des forages de Villars-sous-Yens (VD), Versoix (GE) et Ollon (VD). L'ensemble de ces observations tend à montrer que l'ouest du Plateau suisse est une zone charnière entre la végétation méridionale de la basse vallée du Rhône et celle d'au-delà du Jura. Au débouché du glacier du Rhône et aux confins des glaciers préalpins et jurassiens, cette région est située au carrefour des migrations botaniques post-glaciaires, entre l'axe majeur de la vallée du Rhône et les zones refuges de l'Est. Cette situation lui confère une importance toute particulière. Alors qu'on pensait le Plateau romand profondément raboté par les glaces, on constate aujourd'hui qu'il recèle plus d'une vingtaine de sites interglaciaires et interstadiaires. L'action abrasive du glacier rhodanien a donc été plus modeste qu'envisagée jusqu'ici. L'extension effective des langues glaciaires, tout comme la circulation atmosphérique, diffèrent sans doute d'un interglaciaire à l'autre. Enfin la vitesse des mouvements glaciaires et l'incroyable vitalité végétale sont des paramètres importants, susceptibles d'avoir entraîné des dépôts, très précocement enrichis en pollens.

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We sequenced 1077 bp of the mitochondrial cytochrome b gene and 511 bp of the nuclear Apolipoprotein B gene in bicoloured shrew (Crocidura leucodon, Soricidae) populations ranging from France to Georgia. The aims of the study were to identify the main genetic clades within this species and the influence of Pleistocene climatic variations on the respective clades. The mitochondrial analyses revealed a European clade distributed from France eastwards to north-western Turkey and a Near East clade distributed from Georgia to Romania; the two clades separated during the Middle Pleistocene. We clearly identified a population expansion after a bottleneck for the European clade based on mitochondrial and nuclear sequencing data; this expansion was not observed for the eastern clade. We hypothesize that the western population was confined to a small Italo-Balkanic refugium, whereas the eastern population subsisted in several refugia along the southern coast of the Black Sea.

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In order to contribute to the debate about southern glacial refugia used by temperate species and more northern refugia used by boreal or cold-temperate species, we examined the phylogeography of a widespread snake species (Vipera berus) inhabiting Europe up to the Arctic Circle. The analysis of the mitochondrial DNA (mtDNA) sequence variation in 1043 bp of the cytochrome b gene and in 918 bp of the noncoding control region was performed with phylogenetic approaches. Our results suggest that both the duplicated control region and cytochrome b evolve at a similar rate in this species. Phylogenetic analysis showed that V. berus is divided into three major mitochondrial lineages, probably resulting from an Italian, a Balkan and a Northern (from France to Russia) refugial area in Eastern Europe, near the Carpathian Mountains. In addition, the Northern clade presents an important substructure, suggesting two sequential colonization events in Europe. First, the continent was colonized from the three main refugial areas mentioned above during the Lower-Mid Pleistocene. Second, recolonization of most of Europe most likely originated from several refugia located outside of the Mediterranean peninsulas (Carpathian region, east of the Carpathians, France and possibly Hungary) during the Mid-Late Pleistocene, while populations within the Italian and Balkan Peninsulas fluctuated only slightly in distribution range, with larger lowland populations during glacial times and with refugial mountain populations during interglacials, as in the present time. The phylogeographical structure revealed in our study suggests complex recolonization dynamics of the European continent by V. berus, characterized by latitudinal as well as altitudinal range shifts, driven by both climatic changes and competition with related species.

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The 1:10,000 scale mapping of the southern part of the Aggtelek Plateau (Western Carpathians, Silica Nappe, NE Hungary) and the study of five sections revealed two Middle Triassic reef bodies. In the late Pelsonian the uniform Steinalm Platform was drowned and dissected due to the Reifling Event. A connection with the open sea was established, indicated by the appearance of gladigondolellid conodonts from the early Illyrian. Basins and highs were formed. In the NW part of the studied area lower - middle? Illyrian basinal carbonates were followed by a platform margin reef (early? - middle Illyrian; reef stage 1) developed on a morphological high. This is the oldest known Triassic platform margin reef within the Alpine-Carpathian region. The reef association is dominated by sphinctozoans and microproblematics. The fossils are characteristic of the Wetterstein - type reef communities. Differently from this in the SE part of the studied region a basin existed from the late Pelsonian until the early Ladinian. During the late Illyrian - early Ladinian, the reef prograded to the SE, and reef stage 2 was established. Meanwhile, on the NW part of the platform a lagoon was formed behind the reef. Based on our palaeontological study the stratigraphic range of Colospongia catenulata, Follicatena cautica, Solenolmia manon manon, Vesicocaulis oenipontanus must be extended down to the middle Illyrian. Synsedimentary tectonics were detected in the 1. Binodosus Subzone, 2. Trinodosus Zone - the most part of the Reitzi Zone, 3. Avisianum Subzone.

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Secular variations of the seawater carbon isotopic composition provide evidence for paleoceanographic and paleoclimatic changes and may serve for chemiostratigraphic correlations. The present study aimed to improve the current knowledge on the Upper Permian and Triassic segment of the Phanerozoic marine carbon isotope curve, whose Triassic part was poorly constrained by previous studies. Profiles of inorganic carbon isotopes are provided for sections from Himalaya (Salt Range, Kashmir, Spiti and Nepal), Oman and North Dobrogea (Romania) on the basis of whole-rock carbonate analysis. The data acquired, together with a literature compilation confirmed that most of the Upper Permian is characterized by high δ13C values (averaging +40/00) but failed to detect a positive excursion as suggested by recent compilations. In the light of these observations, the large drop in δ13C values associated with the end-Permian mass extinction appears to be driven by a sudden transfer of previously stocked 13C depleted carbon, rather than by the overturn of a Late Permian stratified ocean. The Triassic data-set outlines significant secular variations. The best documented is a carbon isotope positive excursion just across the Lower-Middle Triassic boundary, globally developed since it was detected in various paleogeographic settings. It is interpreted to reflect variations in surface ocean chemistry, possibly related to increased primary productivity, at times when the biotic recovery after the end-Permian mass-extinction began to accelerate significantly and when a sharp rise in seawater δ34S values occurred globally. Strontium isotope data obtained from well preserved biogenic phosphates allow a refinement of the Middle Triassic segment of the seawater strontium isotope curve and show a major inflexion point of the seawater strontium isotope curve also near the Lower Triassic - Middle Triassic boundary. These facts suggest that the transition from the Early to the Middle Triassic was a time of revolutionary global change which represented an important step in the evolution of Mesozoic marine environments. A tentative carbon isotope curve for the Upper Permian to Upper Triassic time interval is proposed. Its major features are: ? high but constant δ13C values during the Late Permian ? a sharp drop in δ13C values in the latest Permian ? subsequent recovery of δ13C values ? a short-lived positive excursion across the Early-Middle Triassic boundary ? a gradual rise in δ13C values starting in the Late Ladinian or in the Early Carnian It is foreseen that these fluctuations of the carbon isotope curve may serve as chronostratigraphic markers and further assist in the correlation of Permian and Triassic carbonate deposits.

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An earlier study revealed the strong phylogeographical structure of the lesser white-toothed shrew (Crocidura suaveolens group) within the northern Palaearctic. Here, we aim to reconstruct the colonization history of Mediterranean islands and to clarify the biogeography and phylogeographical relationships of the poorly documented Middle East region with the northern Palaearctic. We performed analyses on 998-bp-long haplotypes of the mitochondrial cytochrome b gene of 143 samples collected around the Mediterranean basin, including islands and the Middle East. The analyses suggest that the Cypriot shrew belongs to the rare group of relict insular Pleistocene mammal taxa that have survived to the present day. In contrast, the Cretan, Corsican and Menorcan populations were independently introduced from the Middle East during the Holocene. The phylogeographical structure of this temperate Palaearctic species within the Middle East appears to be complex and rich in diversity, probably reflecting fragmentation of the area by numerous mountain chains. Four deeply divergent clades of the C. suaveolens group occur in the area, meaning that a hypothetical contact zone remains to be located in central western Iran.

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Frailty prevalence in older adults has been reported but is largely unknown in middle-aged adults. We determined the prevalence of frailty indicators among middle-aged and older adults from a general Swiss population characterized by universal health insurance coverage and assessed the determinants of frailty with a special focus on socioeconomic status. Participants aged 50 and more from the population-based 2006-2010 Bus Santé study were included (N = 2,930). Four frailty indicators (weakness, shrinking, exhaustion, and low activity) were measured according to standard definitions. Multivariate logistic regressions were used to determine associations. Overall, 63.5%, 28.7%, and 7.8% participants presented no frailty indicators, one frailty indicator, and two or more frailty indicators, respectively. Among middle-aged participants (50-65 years), 75.1%, 22.2%, and 2.7% presented 0, 1, and 2 or more frailty indicators. The number of frailty indicators was positively associated with age, hypertension, and current smoking and negatively associated with male gender, body mass index, waist-to-hip ratio, and serum total cholesterol level. Lower income level but not education was associated with higher number of frailty indicators. Frailty indicators are frequently encountered in both older and middle-aged adults from the Swiss general population. Despite universal health insurance coverage, household income is independently associated with frailty.

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This article is a taxonomic study of the radiolarian species of the superfamilies Eptingiacea and Saturnaliacea occurring in the middle Carnian fauna from the Koseyahya section, near the town of Elbistan, southeastern Turkey. This fauna is characteristic of the Tetraporobrachia haeckeli Radiolarian Zone as defined in Austria and later found also in Turkey and Oman. It comes from an 8 m thick succession of clayey/cherty limestones from the lower part of the section. In addition, a few species from the late Ladinian and Carnian from Oman and the early Norian from Alaska have also been included in this study, in order to improve some generic diagnoses and to show the diversity and evolutionary trends of some genera. 32 radiolarian species of which 22 are new are described and illustrated, and assigned to 16 genera of which three are new (Capnuchospyris, Veleptingium, and Triassolaguncula). The diagnoses of some species, genera, subfamilies and families have been revised, and the family Eptingiidae has been raised to the rank of superfamily.

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OBJECTIVES: To determine whether the initial benefits of spinal cord stimulation (SCS) treatment for critical limb ischemia (CLI) persist over years. DESIGN: Analysis of data prospectively collected for every CLI patient receiving permanent SCS. Follow-up range 12 to 98 months (mean 46+/-23, median 50 months). POPULATION: 87 patients (28% stage III, 72%stage IV) with unreconstructable CLI due (83%) or not (17%) to atherosclerosis and with an initial sitting/supine transcutaneous pO2 gradient >15 mmHg. METHODS: Assessment of actuarial patient survival (PS), limb salvage (LS) and amputation-free patient survival (AFPS). Analysis of the impact of 15 risk factors on long-term outcomes using the Fischer's exact test for categorical variables and the t test for continuous variables. RESULTS: Follow-up was complete for patient and limb survival. A single non-atherosclerotic patient died during follow-up. Among atherosclerotic patients PS decreased from 88% at 1y, to 76% at 3y, 64% at 5y and 57% at 7y. LS reached 84% at 1y, 78% at 2y, 75% at 3y and remained stable thereafter. Diabetes was found to affect LS (p<0.05) and heart disease to reduce PS (p<0.01). AFPS was reduced in heart patients (p<0.01), diabetics (p<0.05) and in patients with previous stroke (p<0.05). CONCLUSIONS: In CLI patients the beneficial effects of SCS persist far beyond the first year of treatment and major amputation becomes infrequent after the second year.

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This study describes major electrocardiogram (ECG) measurements and diagnoses in a population of African individuals; most reference data have been collected in Caucasian populations and evidence exists for interethnic differences in ECG findings. This study was conducted in the Seychelles islands (Indian Ocean) and included 709 black individuals (343 men and 366 women) aged 25 to 64 years randomly selected from the general population. Resting ECG were recorded by using a validated ECG unit equipped with a measurement and interpretation software (Cardiovit AT-6, Schiller, Switzerland). The epidemiology of 14 basic ECG measurements, 6 composite criteria for left ventricular hypertrophy and 19 specific ECG diagnoses including abnormal rhythms, conduction abnormalities, repolarization abnormalities, and myocardial infarction were examined. Substantial gender and age differences were found for several ECG parameters. Moreover, tracings recorded in African individuals of the Seychelles differed from those collected similarly in Caucasian populations in many respects. For instance, heart rate was approximately 5 beats per minute lower in the African individuals than in selected Caucasian populations, prevalence of first degree atrio-ventricular block was especially high (4.8%), and the average Sokolow-Lyon voltage was markedly higher in African individuals of the Seychelles compared with black and white Americans. The integrated interpretation software detected "old myocardial infarction" in 3.8% of men and 0% of women and "old myocardial infarction possible" in 6.1% and 3%, respectively. Cardiac infarction injury scores are also provided. In conclusion, the study provides reference values for ECG findings in a specific population of people of African descent and stresses the need to systematically consider gender, age, and ethnicity when interpreting ECG tracings in individuals.

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Résumé Les Soricidae sont l'une des plus grandes familles de mammifères avec plus de 300 espèces décrites. Elle a été récemment divisée en trois sous-familles, les Soricidae, qui sont distribuées dans la région Holarctique, les Crocidurinae en Afrique et en Eurasie, et les Myosoricinae en Afrique. La diversité spécifique de cette famille a conduit à des interprétations taxonomiques multiples, qui sont à l'origine de polémiques entre spécialistes, et même les premiers résultats moléculaires ont été fortement contradictoires. Le but de cette thèse est donc d'appliquer des meilleures techniques sur des échantillons mieux ciblés, afin de résoudre les contradictions taxonomiques et comprendre l'histoire de cette famille. Par le biais de marqueurs génétiques mitochondriaux et nucléaires, j'ai étudié: (i) Les relations taxonomiques à différent niveaux hiérarchiques au sein des Soricidae, c'est-à dire, entre les sous-familles, tribus, et genres, ainsi qu'au sein de deux complexes d'espèces largement distribués, et d'une espèce européenne, le but étant d'établir la congruence entre les données génétiques et les interprétations morphologiques classiques. (ii) Les relations biogéographiques, soit l'origine potentielle des différentes sous-familles, tribus, et genres, le nombre d'échanges intercontinentaux, ainsi que la structure phylogéographique à un niveau (péri)-spécifique, afin d'établir l'histoire de la diversification de cette famille. Les analyses combinées d'ADN mitochondrial et nucléaire ont montré un rapport clair entre les taxa à un niveau taxonomique élevé, mettant en évidence les rapports entre les sous-familles, les tribus, et les genres. Bien que Myosorex constitue un groupe monophylétique distinct, sa définition en tant que sous-famille séparée ne peut pas être reconnue. Ainsi, nous proposons d'attribuer un niveau de tribu pour ce clade (inclus dans les Crocidurinae). Nous avons également montré l'inclusion du genre Anourosorex dans les Soricinae et non en position basale dans les Soricidae. Au sein des Crocidurinae, Suncus s'est révélé être paraphylétique, et le genre Diplomesodon devrait être considéré d'un point de vue génétique comme invalide, puisque il se trouve au sein du clade du genre Crocidura. À un niveau taxonomique plus bas, nous avons montré la monophylie de deux complexes d'espèces largement distribués, le groupe de C. suaveolens et de C. olivieri. Néanmoins à l'intérieur de ceux-ci, des différences majeures avec la classification morphologique se sont révélées. Par exemples, C. sibirica n'est pas une espèce valide, les analyses de phylogénie moléculaire ne montrant pas de variations génétiques entre celle-ci et un échantillon de la localité type de C. suaveolens. D'un point de vue biogéographique, les fluctuations climatiques et les activités tectoniques des 20 derniers millions d'années ont fortement influencé la diversité actuelle des Soricidae. À un niveau taxonomique élevé, l'apparition de connexions de terre temporaires entre le Vieux et le Nouveau Monde au Miocène moyen ont mené à plusieurs colonisations indépendantes de l'Amérique par les Soricinae. Celles-ci ónt conduit à une diversification d'une tribu (Notiosoricini), ainsi que de genres (par ex: Cryptotis, Blarina) et d'un sous-genre (Otisorex) endémique au Néarctique. Dans le Vieux Monde, les barrières entre l'Afrique et Eurasie étaient plus perméables, menant à plusieurs échanges bidirectionnels de Crocidurinae. La diversification des clades principaux s'est produite au Miocène, certains clades étant endémiques d'Afrique ou d'Eurasie, tandis que d'autres se sont diversifiés à travers le Vieux Monde. À un niveau spécifique ou péri-spécifique, la fluctuation climatique du Pliocène et les glaciations du Pléistocène ont fortement divisé les populations dans tout le Paléarctique, menant à des entités génétiques distinctes. En Europe, les populations du groupe de C. suaveolens ont été divisées en une lignée Sud-Ouest et une Sud-Est, alors qu'au Proche-Orient et au Moyen-Orient, la diversité de clades est plus importante. En conclusion, mes études ont révélé que du Miocène à nos jours, la diversification des Soricidae a été provoquée par la colonisation de nouveaux habitats (dispersion), ainsi que par l'isolement des populations par diverses barrières (vicariance). Abstract The Soricidae is one of the largest mammalian families with more than 300 species described. It has been recently divided into three subfamilies, the Soricinae, which are distributed in the Holartic region, the Crocidurinae in Africa and Eurasia, and the Myosoricinae in Africa. The specific diversity of this family have led to multiple systematic interpretations and controversies between authors. Fortunately, today, cytotaxonomic, allozymic and molecular studies have permitted to clarify some uncertainties. Nevertheless, the Soricidae remains still poorly known. In this thesis, we aim at understanding with the use of mitochondrial and nuclear markers: (i) the taxonomic relationships at different hierarchical levels within Soricidae, i.e., between the subfamilies, tribes, and genera, as well as within two largely distributed species complexes, and within a European species, the goal being to establish congruence between the genetic data and traditional morphological interpretations; (ii) the biogeographic relationships, especially the potential origin of the different subfamilies, tribes, and genera, the number of transcontinental exchanges, as well as the phylogeographic structure at a (peri)-specific level, in order to establish the history of the genetic diversification of this family. The combined analyses of mitochondrial and nuclear DNA highlight for the first time a clear relationship between taxa at a high taxonomical level, permitting to distinguish the relationships between subfamilies, tribes, and genera. Although Myosorex formed a distinct monophyletic group, its definition as a distinct sub-family cannot be advocated. Thus, we propose to attribute a tribe level for this Glade (included within the Crocidurinae). Additionally, this combination of genes pleads in favour of the inclusion of the genus Anourosorex within the Soricinae and not in a basal position within the Soricidae. Within the Crocidurinae, Suncus appeared to be paraphyletic, and Diplomesodon should be considered from a genetic point of view as invalid, and is presently considered as Crocidura. At a lower taxonomic level, we showed the monophyly of two widely distributed species complexes, the C. suaveolens group and the C. olivieri group. Nevertheless within those, we showed major differences compared to morphological classification. For examples, C. sibirica revealed to not be a valid species, the molecular phylogenetic analyses failed to evidence genetical variations between it and samples of the type locality of C. suaveolens. In a biogeographic point of view, the climatic fluctuations and the tectonic plate activities of the last 20 Myr have strongly influenced the actual diversity of the family. At a high taxonomic level, the successive land bridge connections between the Old and the New World, which occurred during the Middle Miocene, have led to several independent colonisations of America by Soricinae, and a subsequent diversification of endemic Nearctic's tribe (Notiosoricini), genera (e.g. Cryptotis, Blaring) and sub-genus (Otisorex) within the Soricinae. Within the Old World, the barriers between Africa and Eurasia were more permeable, leading to several bidirectional exchanges within the Crocidurinae. The diversification of major clades occurred through the Miocene, some clades being endemic to Africa or Eurasia, whereas others diversified through the Old World. At a species level or a peri-specific level, the Pliocene climatic fluctuation and the Pleistocene glaciations have strongly divided the populations throughout the Palaearctic, leading to well defined genetic entities. In Europe, populations of the C. suaveolens group were split in a classical south-western and south-eastern lineage. In contrast, the Near East and the Middle East reveal many differentiated clades. In conclusion, our studies revealed that, from the Miocene to present, the diversification and speciation events within the Soricidae were caused by natural colonisation of new habitats (dispersion) and isolation of populations by various barriers (vicariance).