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Evolutionary History of Moles in Western Europe: One Mole May Hide Another!
Abstract
If there’s one mammal all garden owners know well, without ever having actually seen one, it is the mole. A friendly character in childhood cartoons (The Mole by Zdenek Miler) or comics (Les Riquiquis from the French magazine Youpi j’ai compris), this animal is often hated by gardeners and lawn tenders who hardly appreciate the numerous molehills left behind by those that dig up the earth when making their underground tunnels. The methods put into practice to get rid of them in a more or less natural way, or to kill them, are therefore numerous: fritillary or Euphorbia bulbs, ultrasonic mole repellers, putting dog hair in the molehills, mole-killing chemical products, mole-traps, fumigation, etc. not to mention professional mole-trappers! There is in fact a whole network of professional mole-trappers who are experts in the art of using traditional mole traps, and have followed theoretical and practical training courses for one to three years.
Keywords : Geographical distribution; Mitochondrial DNA; Morphological analysis; Nuclear DNA; Species; Unexpected biodiversity
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A recent mitochondrial study showed that the common mole, Talpa europaea, is paraphyletic; which could be explained either by an ancient introgression of mtDNA from the Iberian blind mole T. occidentalis to T. europaea, or the existence of a differentiated taxonomic entity in Northern Spain that needs to be described. In this study we combined mitochondrial (Cytb) and nuclear (HDAC2) data to test these two alternative hypotheses. We also used ecological niche modeling to make inferences on the evolutionary history of these moles. Based on both mitochondrial and nuclear data and an extensive geographical sampling (399 sequenced individuals) we show that the populations of T. europaea from Spain and Southwestern France (south of the Loire River) are phylogenetically closer to T. occidentalis than to T. europaea. The Spanish- French lineage has a somewhat intermediate morphology between T. europaea and T. occidentalis, with some characters resembling more to T. occidentalis and others more to T. europaea. Moreover it also seems to have several distinctive characters, suggesting that it should be recognized as a new species. Within the three species we found a marked phylogeographical pattern, with several allopatric or parapatric lineages, dating from the Pleistocene. Our genetic data combined with species distribution models support the presence of several putative glacial refugia during glacial maxima for each species.
A mtDNA based study of the population genetics of moles recently captured in France allowed us to discover a new species, Talpa aquitania nov. sp. We are giving here a preliminary description of the new species. Its distribution covers an area lying south and west of the course of the Loire river in France and beyond the Pyrenees, a part of Northern Spain.
Our understanding of the effect of Pleistocene climatic changes on the biodiversity of European mammals mostly comes from phylogeographical studies of non-subterranean mammals, whereas the influence of glaciation cycles on subterranean mammals has received little attention. The lack of data raises the question of how and to what extent the current amount and distribution of genetic variation in subterranean mammals is the result of Pleistocene range contractions/expansions. The common mole (Talpa europaea) is a strictly subterranean mammal, widespread across Europe, and represents one of the best candidates for studying the influence of Quaternary climatic oscillation on subterranean mammals. Cytochrome b sequences, as obtained from a sampling covering the majority of the distribution area, were used to evaluate whether Pleistocene climate change influenced the evolution of T. europaea and left a trace in the genetic diversity comparable to that observed in non-subterranean small mammals. Subsequently, we investigated the occurrence of glacial refugia by comparing the results of phylogeographical analysis with species distribution modelling. We found three differentiated mitochondrial DNA lineages: two restricted to Spain and Italy and a third that was widespread across Europe. Phylogenetic inferences and the molecular clock suggest that the Spanish moles represent a highly divergent and ancient lineage, highlighting for the first time the paraphyly of T. europaea. Furthermore, our analyses suggest that the genetic break between the Italian and the European lineages predates the last glacial phase. Historical demography and spatial principal component analysis further suggest that the Last Glacial Maximum left a signature both in the Italian and in the European lineages. Genetic data combined with species distribution models support the presence of at least three putative glacial refugia in southern Europe (France, Balkan Peninsula and Black Sea) during the
Biotic interactions have been considered as an important factor to be included in species distribution modelling, but little is known about how different types of interaction or different strategies for modelling affect model performance. This study compares different methods for including interspecific interactions in distribution models for bees, their brood parasites, and the plants they pollinate. Host–parasite interactions among bumble bees (genus Bombus: generalist pollinators and brood parasites) and specialist plant–pollinator interactions between Centris bees and Krameria flowers were used as case studies. We used 7 different modelling algorithms available in the BIOMOD R package. For Bombus, the inclusion of interacting species distributions generally increased model predictive accuracy. The improvement was better when the interacting species was included with its raw distribution rather than with its modeled suitability. However, incorporating the distributions of non-interacting species sometimes resulted in similarly increased model accuracy despite their being no significance of any interaction for the distribution. For the Centris-Krameria system the best strategy for modelling biotic interactions was to include the interacting species model-predicted values. However, the results were less consistent than those for Bombus species, and most models including biotic interactions showed no significant improvement over abiotic models. Our results are consistent with previous studies showing that biotic interactions can be important in structuring species distributions at regional scales. However, correlations between species distributions are not necessarily indicative of interactions. Therefore, choosing the correct biotic information, based on biological and ecological knowledge, is critical to improve the accuracy of species distribution models and forecast distribution change.
G- and C-band pattern and NOR's of Talpa occidentalis metaphase chromosomes are described. The C-band pattern shows that the chromosomal pair causing karyotypic differences in the genus Talpa presents their short arm totally heterochromatic. The implication of this fact in the karyotypic evolution of this genus and the taxonomic status of T. occidentalis are discussed.
A description is given of the last premolar and molars of T. europaea from The Netherlands and T. occidentalis of Spain. The metrical and morphological differences between the dentitions of the two species are small. A review is given of the literature of fossil Talpa in the Pleistocene of Europe. The suggestion that T. minor is ancestral to T. caeca is rejected. T. fossilis is tentatively considered a junior synonym of T. europaea.
Using both nuclear and mitochondrial sequences, we demonstrate high genetic differentiation in the genus Talpa and confirm the existence of cryptic species in the Caucasus and Anatolia, namely, T. talyschensis Vereschagin, 1945, T. ognevi Stroganov, 1948, and Talpa ex gr. levantis. Our data support four clades in the genus Talpa that showed strong geographical associations. The ‘europaea’ group includes six species from the western portion of the genus' range (T. europaea, T. occidentalis, T. romana, T. caeca, T. stankovici, and T. levantis s.l.); another three groups are distributed further east: the ‘caucasica’ group (Caucasus), the ‘davidiana’ group (eastern Anatolia and Elburz) and T. altaica (Siberia). The phylogenetic position of T. davidiana was highlighted for the first time. The order of basal branching remains controversial, which can be attributed to rapid diversification events. The molecular time estimates based on nuclear concatenation estimated the basal divergence of the crown Talpa during the latest Miocene. A putative scenario of Talpa radiation and issues of species delimitation are discussed. © 2015 The Linnean Society of London
Deals with the species of Talpidae (moles, shrew-moles and desmans) and Chrysochloridae (golden moles) families of Eurasia and North America, and the southern half of Africa, respectively. Particular attention is given to European mole Talpa europaea, star-nosed mole Condylura cristata of North America and the aquatic Pyrenean desman Galemys pyrenaicus. The introduction shows the range of species and their taxonomy and distribution. Subsequent chapters describe, with illustrations and figures, the structural and physiological adaptations for digging, including analyses of the energetic costs of digging and the construction of the tunnel system; feeding; life, death and reproduction; patterns of daily and annual activity; social organization, territories and communication; man/mole problems and mole control; and lastly, for desmans, swimming as a way of life. Appendices provide information on keeping moles in captivity and on mole ectoparasites. -J.W.Cooper
The analysis of a dense grid of high resolution seismic profiles collected offshore the present day Loire River estuary indicates the presence of a thick and complex Pleistocene coastal wedge between the coast and 50 meters water depth. Most of this coastal wedge is preserved in a fossil valley network starting 10 km off the coast on the "Precontinent Breton" topographies and wedging out progressively 50 km in the offshore where the shelf flattens. This system is comprised of three main valleys 30 km long, 40-60 m deep and 0.7 to 4 km wide each, in average, with a northern valley incompletely filled by sediment. These valleys are incised into Eocene (Ypresian-Bartonian) sedimentary rocks lying unconformably on the metamorphic and magmatic rocks of the South Armorican Massif basement. The coastal wedge is comprised of six seismic units. From the base of the valleys to the seafloor, these units are successively interpreted as (1) colluvial (U1) and braided river deposits (U2), overlain by restricted marine to estuarine sediments (U3), and (2) straight to meandering fluvial deposits (U4) giving rise vertically to floodout marine sediments (U5). The whole sediment pile is capped by open marine bioturbated mudstones (U6). This succession of seismic units is organised in two depositional sequences bounded by an unconformity of regional extent, which corresponds to a drastic change in the paleovalley fill architecture. The lower sequence fills up the southern and central valleys when the upper sequence fills up the northern valley network. Both sequences are sharply truncated by a ravinement surface at the base of the offshore shales formed during the Holocene marine transgression. The correlation of the observed depth of the incisions and transgression surfaces with the global sea-level curve provides an indirect estimate of the ages of the depositional sequences. The lower sequence is probably Saalian (130-200 Ka, MIS 6) and the upper one Weischelian (110-12 Ka, MIS 2-4 and 5a-d). The ages and the infill of these valleys are very close to the ones observed in the largest paleovalleys of the inner French Atlantic shelf (Gironde, English Channel) and consistent with the sedimentary record of the continental Loire River (stepped terraces). In the later case, the Saalian period corresponds to a sharp increase on the incision of the river that shaped the morphology and determined the location of the present-day Loire valley.
Nine sites along the downstream and middle section of the Loire river at Cordemais (Loire Atlantique, France) situated in the estuary to Saint-Nicolas-de-Bourgueil (Indre et Loire, France) were investigated. Interdisciplinary studies combined palynology, geography, archaeology, sedimentology and history, thus enabling us to reconstruct the evolution and the long-term response of the Loire valley ecosystem to natural variations and anthropogenic pressure from the Preboreal to the present in an integrated manner. The Atlantic marine transgression (between 7000 and 5000B.P.) caused the level of brackish water to increase at Oudon between 6740+205/–200 and 5010+115/–100 B.P. This phenomenon, which was the first of its kind to be detected near the central Loire region (approximately 80km from the current mouth of the Loire river), was accompanied by the development of subhalophile vegetation (Chenopodiaceae) and the appearance of dinoflagellate cysts. A regressive phase occurred during the Subboreal, about 4500 B.P., and led to the erosion of most of the estuarian sediments and to the disappearance of plant species linked to salinity at Oudon: peat deposits built up at most other sites. Human activities had an early effect; moderate deforestation took place at Champtoc about 6600B.P. as farming was already orientated towards rearing animals. However, possibly cultivated plants were present towards the middle and the end of the Neolithic period (wheat, rye, buckwheat, flax) at about 5600B.P. and chestnut and walnut were probably exploited in the Loire valley region at about 4600B.P. The Bronze Age seemed to mark a phase when societies settled down (planting of vineyards) and deforestation peaked from the Gallo-Roman period onwards. The textile industry (flax, hemp), in the context of crop rotation set up during the Iron Age, developed rapidly during the Middle Ages, whereas nowadays the rearing of animals is the dominant activity in the Loire Valley, following the introduction of maize into the region in 1950.