Crustal structure and reflectivity of the Swiss Alps from 3-Dimensional seismic modeling. 2. Penninic Nappes

Surface geological mapping, laboratory measurements of rock properties, and seismic reflection data are integrated through three-dimensional seismic modeling to determine the likely cause of upper crustal reflections and to elucidate the deep structure of the Penninic Alps in eastern Switzerland. Results indicate that the principal upper crustal reflections recorded on the south end of Swiss seismic line NFP20-EAST can be explained by the subsurface geometry of stacked basement nappes. In addition, modeling results provide improvements to structural maps based solely on surface trends and suggest the presence of previously unrecognized rock units in the subsurface. Construction of the initial model is based upon extrapolation of plunging surface. structures; velocities and densities are established by laboratory measurements of corresponding rock units. Iterative modification produces a best fit model that refines the definition of the subsurface geometry of major structures. We conclude that most reflections from the upper 20 km can be ascribed to the presence of sedimentary cover rocks (especially carbonates) and ophiolites juxtaposed against crystalline basement nappes. Thus, in this area, reflections appear to be principally due to first-order lithologic contrasts. This study also demonstrates not only the importance of three-dimensional effects (sideswipe) in interpreting seismic data, but also that these effects can be considered quantitatively through three-dimensional modeling.

Introduction of snow and geomorphic disturbance variables into predictive models of alpine plant distribution in the Western Swiss Alps

Indirect topographic variables have been used successfully as surrogates for disturbance processes in plant species distribution models (SDM) in mountain environments. However, no SDM studies have directly tested the performance of disturbance variables. In this study, we developed two disturbance variables: a geomorphic index (GEO) and an index of snow redistribution by wind (SNOW). These were developed in order to assess how they improved both the fit and predictive power of presenceabsence SDM based on commonly used topoclimatic (TC) variables for 91 plants in the Western Swiss Alps. The individual contribution of the disturbance variables was compared to TC variables. Maps of models were prepared to spatially test the effect of disturbance variables. On average, disturbance variables significantly improved the fit but not the predictive power of the TC models and their individual contribution was weak (5.6% for GEO and 3.3% for SNOW). However their maximum individual contribution was important (24.7% and 20.7%). Finally, maps including disturbance variables (i) were significantly divergent from TC models in terms of predicted suitable surfaces and connectivity between potential habitats, and (ii) were interpreted as more ecologically relevant. Disturbance variables did not improve the transferability of models at the local scale in a complex mountain system, and the performance and contribution of these variables were highly species-specific. However, improved spatial projections and change in connectivity are important issues when preparing projections under climate change because the future range size of the species will determine the sensitivity to changing conditions.

Large-scale DNA sequencing in taxonomy and conservation : a case study with the mayfly family Heptageniidae (Ephemeroptera) in the Alps and Madagascar

Les larves aquatiques d'éphémères (Ephemeroptera) colonisent toutes les eaux douces du monde et sont couramment utilisées comme bio-indicateurs de la qualité de l'eau. Le genre Rhithrogena (Heptageniidae) est le deuxième plus diversifié chez les éphémères, et plusieurs espèces européennes ont une distribution restreinte dans des environnements alpins sensibles. Les espèces de Rhithrogena ont été classées en "groupes d'espèces" faciles à identifier. Cependant, malgré leur importance écologique et en terme de conservation, beaucoup d'espèces présentent des différences morphologiques ambiguës, suggérant que lataxonomie actuelle ne refléterait pas correctement leur diversité évolutive. De plus, aucune information sur leurs relations, leur origine, le taux de spéciation ou les mécanismes ayant provoqué leur remarquable diversification dans les Alpes n'est disponible. Nous avons d'abord examiné le statut spécifique d'environ 50% des espèces européennes de Rhithrogena en utilisant un large échantillonnage de populations alpines incluant 22 localités typiques, ainsi qu'une analyse basée sur le modèle général mixte de Yule et de coalescence (GMYC) appliqué à un gène mitochondrial standard (coxl) et à un gène nucléaire développé spécifiquement pour cette étude. Nous avons observé un regroupement significatif des séquences coxl en 31 espèces potentielles, et nos résultats ont fortement suggéré la présence d'espèces cryptiques et de fractionnements taxonomiques excessifs chez les Rhithrogena. Nos analyses phylogénétiques ont démontré la monophylie de quatre des six groupes d'espèces reconnus présents dans notre échantillonnage. La taxonomie ADN développée dans cette étude pose les bases d'une future révision de ce genre important mais cryptique en Europe. Puis nous avons mené une étude phylogénétique multi-gènes entre les espèces européennes de Rhithrogena. Les données provenant de trois gènes nucléaires et de deux gènes mitochondriaux ont été largement concordantes, et les relations entre les espèces bien résolues au sein de la plupart des groupes d'espèces dans une analyse combinant tous les gènes. En l'absence de points de calibration extérieurs tels que des fossiles, nous avons appliqué à nos données mitochondriales une horloge moléculaire standard pour les insectes, suggérant une origine des Rhithrogena alpins à la limite Oligocène / Miocène. Nos résultats ont montré le rôle prépondérant qu'ont joué les glaciations du quaternaire dans leur diversification, favorisant la spéciation d'au moins la moitié des espèces actuelle dans les Alpes. La biodiversité et le taux d'endémisme à Madagascar, notamment au niveau de la faune des eaux douces, sont parmi les plus extraordinaires et les plus menacés au monde. On pense que beaucoup d'espèces d'éphémères sont restreintes à un seul bassin versant (microendémisme) dans les zones forestières, ce qui les rendrait particulièrement sensibles à la réduction et à la dégradation de leur habitat. Mis à part deux espèces décrites, Afronurus matitensis et Compsoneuria josettae, les Heptageniidae sont pratiquement inconnus à Madagascar. Les deux genres ont une distribution discontinue en Afrique, à Madagascar et en Asie du Sud-Est, et leur taxonomie complexe est régulièrement révisée. L'approche standard pour comprendre leur diversité, leur endémisme et leur origine requerrait un échantillonnage étendu sur plusieurs continents et des années de travaux taxonomiques. Pour accélérer le processus, nous avons utilisé des collections de musées ainsi que des individus fraîchement collectés, et appliqué une approche combinant taxonomie ADN et phylogénie. L'analyses GMYC du gène coxl a délimité 14 espèces potentielles à Madagascar, dont 70% vraisemblablement microendémiques. Une analyse phylogénique incluant des espèces africaines et asiatiques portant sur deux gènes mitochondriaux et quatre gènes nucléaires a montré que les Heptageniidae malgaches sont monophylétiques et groupe frère des Compsoneuria africains. L'existence de cette lignée unique, ainsi qu'un taux élevé de microendémisme, mettent en évidence leur importance en terme de conservation. Nos résultats soulignent également le rôle important que peuvent jouer les collections de musées dans les études moléculaires et en conservation. - Aquatic nymphs of mayflies (Ephemeroptera) colonize all types of freshwaters throughout the world and are extensively used as bio-indicators of water quality. Rhithrogena (Heptageniidae) is the second most species-rich genus of mayflies, and several European species have restricted distributions in sensitive Alpine environments and therefore are of conservation interest. The European Rhithrogena species are arranged into "species groups" that are easily identifiable. However, despite their ecological and conservation importance, ambiguous morphological differences among many species suggest that the current taxonomy may not accurately reflect their evolutionary diversity. Moreover, no information about their relationships, origin, timing of speciation and mechanisms promoting their successful diversification in the Alps is available. We first examined the species status of ca. 50% of European Rhithrogena diversity using a widespread sampling scheme of Alpine species that included 22 type localities, general mixed Yule- coalescent (GMYC) model analysis of one standard mitochondrial (coxl) and one newly developed nuclear marker. We observed significant clustering of coxl into 31 GMYC species, and our results strongly suggest the presence of both cryptic diversity and taxonomic oversplitting in Rhithrogena. Phylogenetic analyses recovered four of the six recognized species groups in our samples as monophyletic. The DNA taxonomy developed here lays the groundwork for a future revision of this important but cryptic genus in Europe. Then we conducted a species-level, multiple-gene phylogenetic study of European Rhithrogena. Data from three nuclear and two mitochondrial loci were broadly congruent, and species-level relationships were well resolved within most species groups in a combined analysis. In the absence of external calibration points like fossils, we applied a standard insect molecular clock hypothesis to our mitochondrial data, suggesting an origin of Alpine Rhithrogena in the Oligocene / Miocene boundary. Our results highlighted the preponderant role that quaternary glaciations played in their diversification, promoting speciation of at least half of the current diversity in the Alps. Madagascar's biodiversity and endemism are among the most extraordinary and endangered in the world. This includes the island's freshwater biodiversity, although detailed knowledge of the diversity, endemism, and biogeographic origin of freshwater invertebrates is lacking. Many mayfly species are thought to be restricted to single river basins (microendemic species) in forested areas, making them particularly sensitive to habitat reduction and degradation. The Heptageniidae are practically unknown in Madagascar except for two described species, Afronurus matitensis and Compsoneuria josettae. Both genera have a disjunct distribution in Africa, Madagascar and Southeast Asia, and a complex taxonomic status still in flux. The standard approach to understanding their diversity, endemism, and origin would require extensive field sampling on several continents and years of taxonomic work. Here we circumvent this using museum collections and freshly collected individuals in a combined approach of DNA taxonomy and phylogeny. The cox/-based GMYC analysis revealed 14 putative species on Madagascar, 70% of which potentially microendemics. A phylogenetic analysis that included African and Asian species and data from two mitochondrial and four nuclear loci indicated the Malagasy Heptageniidae are monophyletic and sister to African Compsoneuria. The observed monophyly and high microendemism highlight their conservation importance. Our results also underline the important role that museum collections can play in molecular studies, especially in critically endangered biodiversity hotspots like Madagascar.

Formica paralugubris (Hymenoptera, Formicidae) in the Italian Alps from new data and old data revisited.

We provide evidence that Formica paralugubris Seifert, 1996, a species of wood and recently described from Switzerland, is present in the Italian Alps. Until 1996, this species was confounded with F.lugubris Zetterstedt, 1838. We examine the wood and collection deposited at the University of Pavia (Italy) and collect new samples within the Italian Alps. Formica paralugubris seems to be more abundant than F.lugubris. Moreover, both species are found in sympatry in some localities

Alps, genes, and chromosomes: their role in the formation of species in the Sorex araneus group (Mammalia, Insectivora), as inferred from two hybrid zones.

During the Pleistocene glaciations, the Alps were an efficient barrier to gene flow between isolated populations, often leading to allopatric speciation. Afterwards, the Alps strongly influenced the post-glacial recolonization of Europe and represent a major suture zone between differentiated populations. Two hybrid zones in the Swiss and French Alps between genetically and chromosomally well-differentiated species-the Valais shrew, Sorex antinorii, and the common shrew, S. araneus-were studied karyotypically and by analyzing the distribution of seven microsatellite loci. In the center of the Haslital hybrid zone the two species coexist over a distance of 900 m. Hybrid karyotypes, among them the most complex known in Sorex, are rare. F-statistics based on microsatellite data revealed a strong heterozygote deficit only in the center of the zone, due to the sympatric distribution of the two species with little hybridization between them. Structuring within the species (both F(IS) and F(ST)) was low. An hierarchical analysis showed a high level of interspecific differentiation. Results were compared with those previously reported in another hybrid zone located at Les Houches in the French Alps. Genetic structuring within and between species was comparable in both hybrid zones, although chromosomal incompatibilities are more important in Haslital, where a linkage block of the race-specific chromosomes should additionally impede gene flow. Evidence for a more restricted gene flow in Haslital comes from the genetically intermediate hybrid karyotypes, whereas in Les Houches, hybrid karyotypes are genetically identical to individuals of the pure karyotypic races. Genic and chromosomal introgression was observed in Les Houches, but not in Haslital. The possible influence of a river, separating the two species at Les Houches, on gene flow is discussed.

New stratigraphic data from the Lower Penninic between the Adula nappe and the Gotthard massif and consequences for the tectonics and the paleogeography of the Central Alps

New stratigraphic data along a profile from the Helvetic Gotthard massif to the remnants of the North Penninic basin in eastern Ticino and Graubunden are presented. The stratigraphic record together with existing geochemical and structural data, motivate a new interpretation of the fossil European distal margin. We introduce a new group of Triassic facies, the North-Penninic-Triassic (NPT), which is characterised by the Ladinian ``dolomie bicolori''. The NPT was located in-between the Brianconnais carbonate platform and the Helvetic lands. The observed horizontal transition, coupled with the stratigraphic superposition of a Helvetic Liassic on a Briaconnais Triassic in the Luzzone-Terri nappe, links, prior to Jurassic rifting, the Brianconnais paleogeographic domain at the Helvetic margin, south of the Gotthard. Our observations suggest that the Jurassic rifting separated the Brianconnais domain from the Helvetic margin by complex and protracted extension. The syn-rift stratigraphic record in the Adula nappe and surroundings suggests the presence of a diffuse rising area with only moderately subsiding basins above a thinned continental and proto-oceanic crust. Strong subsidence occurred in a second phase following protracted extension and the resulting delamination of the rising area. The stratigraphic coherency in the Adula's Mesozoic questions the idea of a lithospheric m lange in the eclogitic Adula nappe, which is more likely to be a coherent alpine tectonic unit. The structural and stratigraphic observations in the Piz Terri-Lunschania zone suggest the activity of syn-rift detachments. During the alpine collision these faults are reactivated (and inverted) and played a major role in allowing the Adula subduction, the ``Penninic Thrust'' above it and in creating the structural complexity of the Central Alps. (C) 2012 Elsevier B.V. All rights reserved.

Reduced genetic diversity, increased isolation and multiple introductions of invasive giant hogweed in the western Swiss Alps.

Abstract The giant hogweed (Heracleum mantegazzianum) has successfully invaded 19 European countries as well as parts of North America. It has become a problematic species due to its ability to displace native flora and to cause public health hazards. Applying population genetics to species invasion can help reconstruct invasion history and may promote more efficient management practice. We thus analysed levels of genetic variation and population genetic structure of H. mantegazzianum in an invaded area of the western Swiss Alps as well as in its native range (the Caucasus), using eight nuclear microsatellite loci together with plastid DNA markers and sequences. On both nuclear and plastid genomes, native populations exhibited significantly higher levels of genetic diversity compared to invasive populations, confirming an important founder event during the invasion process. Invasive populations were also significantly more differentiated than native populations. Bayesian clustering analysis identified five clusters in the native range that corresponded to geographically and ecologically separated groups. In the invaded range, 10 clusters occurred. Unlike native populations, invasive clusters were characterized by a mosaic pattern in the landscape, possibly caused by anthropogenic dispersal of the species via roads and direct collection for ornamental purposes. Lastly, our analyses revealed four main divergent groups in the western Swiss Alps, likely as a consequence of multiple independent establishments of H. mantegazzianum.

Phylogeography and recolonization of the Swiss Alps by the Valais shrew (Sorex antinorii), inferred with autosomal and sex-specific markers.

Using one male-inherited, one female-inherited and eight biparentally inherited markers, we investigate the population genetic structure of the Valais shrew (Sorex antinorii) in the Swiss Alps. Bayesian analysis on autosomal microsatellites suggests a clear genetic differentiation between two groups of populations. This geographically based structure is consistent with two separate postglacial recolonization routes of the species into Switzerland from Italian refugia after the last Pleistocene glaciations. Sex-specific markers also confirm genetic structuring among western and eastern areas, since very few haplotypes for either Y chromosome or mtDNA genome are shared between the two regions. Overall, these results suggest that two already well-differentiated genetic lineages colonized the Swiss Alps and came into secondary contact in the Rhône Valley. Low level of admixture between the two lineages is likely explained by the mountainous landscape structure of lateral valleys orthogonal to the main Rhône valley.

Subduction of continental crust in the Western Alps

As a result of recent deep reflection and refraction seismology the crustal structure of the Western Alps is now quite well-defined. However, this raises the question of what is present below the Moho, such as a crustal eclogitic root. This study attempts to estimate the volume of this eclogitic root on the basis of palinspastic reconstructions. Even with a minimum estimate of the crustal material involved in the subduction processes which took place during the Alpine orogeny, a significant eclogitized crustal root must be present down to depths of around 100 km below the Po plain. A maximum estimate suggests that a large part of this root could now be recycled in the asthenosphere.

Structural analysis, clay mineralogy and K-Ar dating of fault gouges from Centovalli Line (Central Alps) for reconstruction of their recent activity

Between the cities of Domodossola and Locarno, the complex ``Centovalli Line'' tectonic zone of the Central Alps outlines deformation phases over a long period of time (probably starting similar to 30 Ma ago) and under variable P-T conditions. The last deformation phases developed gouge-bearing faults with a general E-W trend that crosscuts the roots of the Alpine Canavese zone and the Finero ultramafic body. Kinematic indicators show that the general motion was mainly dextral associated with back thrusting towards the S. The <2 mu m clay fractions of fault gouges from Centovalli Line consist mainly of illite, smectite and chlorite with varied illite-smectite, chlorite-smectite and chlorite-serpentine mixed-layers. Constrained with the illite crystallinity index, the thermal conditions induced by the tectonic activity show a gradual trend from anchizonal to diagenetic conditions. The <2 and <0.2 mu M clay fractions, and hydrothermal K-feldspar separates all provide K-Ar ages between 14.2 +/- 2.9 Ma and roughly 0 Ma, with major episodes at about 12,8, 6 and close to 0 Ma These ages set the recurrent tectonic activity and the associated fluid circulations between Upper Miocene and Recent. On the basis of the K-Ar ages and with a thermal gradient of 25-30 degrees C/km, the studied fault zones were located at a depth of 4-7 km. If they were active until now as observed in field, the exhumation was approximately 2.5-3.0 km for the last 12 Ma with a mean velocity of 0.4 mm/y. Comparison with available models on the recent Alpine evolution shows that the tectonic activity in the area relates to a continuum of the back-thrusting movements of the Canavese Line, and/or to several late-extensional phases of the Rhone-Simplon line. The Centovalli-Val Vigezzo zone therefore represents a major tectonic zone of the Central-Western Alps resulting from different interacting tectonic events. (C) 2011 Elsevier B.V. All rights reserved.

The thermal circuits of the Argentera Massif (western Alps, Italy): An example of low-enthalpy geothermal resources controlled by Neogene alpine tectonics

This work presents geochemistry and structural geology data concerning the low enthalpy geothermal circuits of the Argentera crystalline Massif in northwestern Italian Alps. I n this area some thermal springs (50-60 degreesC), located in the small Bagni di Vinadio village, discharge mixtures made up of a Na-Cl end-member and a Na-SO4 component. The latter is also discharged by the thermal springs of Terme di Valdieri located some kilometres apart within the same tectonic complex. Both end-members share the same meteoric origin and the same reservoir temperature, which is close to 150 degreesC. Explanations are thus required to understand how they reach the surface and how waters of the same origin and circulating in similar rocks can attain such different compositions. Sodium-sulphate waters discharged at both sites, likely represent the common interaction product of meteoric waters with the widespread granitic-migmatitic rocks of the Argentera Massif, whereas Na-CI waters originate through leaching of mineralised cataclastic rocks, which are rich in phyllosilicatic minerals and fluid inclusions, both acting as Cl- sources. Due to the relatively low inferred geothermal gradient of the region, -25C/km, meteoric waters have to descend to depths of 5.5-6 km to attain temperatures of similar to 150 degreesC. These relevant depths can be reached by descending meteoric waters, due to the recent extensional stress field, which allows the development of geothermal circulations at greater depths than in other sectors of the Alps by favouring a greater fractures aperture. The ascent of the thermal waters rakes place along brittle shear zones. In both sites, the thermal waters emerge at the bottoms of the valleys, close to either the lateral termination of a brittle shear zone at Terme di Valdieri, or a step-over between two en-echelon brittle shear zones at Bagni di Vinadio. These observations attest to a strong control operated on the location of outlet regions by both brittle tectonics and the minima in hydraulic potential inside the fractured massif.

Detection and characterization of hydraulically active fractures in a carbonate aquifer: Results from self-potential, temperature and fluid conductivity logging along a 260-m-deep borehole in the Combioula hydrothermal system in the southwestern Swiss Alps

A geophysical and geochemical study has been conducted in a fractured carbonate aquifer located at Combioula in the southwestern Swiss Alps with the objective to detect and characterize hydraulically active fractures along a 260-m-deep borehole. Hydrochemical analyses, borehole diameter, temperature and fluid electrical conductivity logging data were integrated in order to relate electrokinetic self-potential signals to groundwater flow inside the fracture network. The results show a generally good, albeit locally variable correlation of variations of the self-potential signals with variations in temperature, fluid electrical conductivity and borehole diameter. Together with the hydrochemical evidence, which was found to be critical for the interpretation of the self-potential data, these measurements not only made it possible to detect the hydraulically active fractures but also to characterize them as zones of fluid gain or fluid loss. The results complement the available information from the corresponding litholog and illustrate the potential of electrokinetic self-potential signals in conjunction with temperature, fluid electrical conductivity and hydrochemical analyses for the characterization of fractured aquifers, and thus may offer a perspective for an effective quantitative characterization of this increasingly important class of aquifers and geothermal reservoirs.