Long-distance wolf recolonization of France and Switzerland inferred from non-invasive genetic sampling over a period of 10 years

In the early 1900s, the wolf (Canis lupus) was extirpated from France and Switzerland. There is growing evidence that the species is presently recolonizing these countries in the western Alps. By sequencing the mitochondrial DNA (mtDNA) control region of various samples mainly collected in the field (scats, hairs, regurgitates, blood or tissue; n = 292), we could (1) develop a non-invasive method enabling the unambiguous attribution of these samples to wolf, fox (Vulpes vulpes) or dog (Canis familiaris), among others; (2) demonstrate that Italian, French and Swiss wolves share the same mtDNA haplotype, a haplotype that has never been found in any other wolf population world-wide. Combined together, field and genetic data collected over 10 years corroborate the scenario of a natural expansion of wolves from the Italian source population. Furthermore, such a genetic approach is of conservation significance, since it has important consequences for management decisions. This first long-term report using non-invasive sampling demonstrates that long-distance dispersers are common, supporting the hypothesis that individuals may often attempt to colonize far from their native pack, even in the absence of suitable corridors across habitats characterized by intense human activities.

Mixing of Rhône River water in Lake Geneva (Switzerland-France) inferred from stable hydrogen and oxygen isotope profiles

Depth profiles were sampled at different locations throughout Lake Geneva on a monthly and seasonal basis over the course of 2 years and analysed for their stable hydrogen and oxygen isotope compositions. The isotopic compositions indicate an isotopic stratification in the metalimnion during summer and fall. This is related to mixing of Rhône River water, which in summer is dominated by snow and glacier melt waters, and lake water, with the latter having a homogenous isotopic composition. The observed interflow layer is 7-15 m thick and can be traced by the distinct stable isotope composition of the water for about 55 km throughout the lake as well as into shallow bay regions. Depth of the interflow layer close to the Rhône River mouth is similar to those previously described based on echo-soundings and turbidity profiles of sediment dispersion. In contrast to previous descriptions of the interflow within Lake Geneva, the stable isotope compositions allow for direct, natural tracing of the Rhône River water even in cases where the turbidity and conductivity measurements do not indicate such an interflow. In addition, the method allows for a quantification of the Rhône River and lake water in the interflow with the fraction of Rhône River water within the interflow estimated to be up to 37% in summer. The isotopic composition further indicates different vertical mixing processes within the two lake basins of Lake Geneva, related to the density gradients and local stability within the water column. The method may be applicable to other lakes in catchments with large differences in the topography as water that originates from high altitudes or glaciers has a distinct oxygen and hydrogen isotope composition compared to other sources of water originating at lower altitudes and/or from direct precipitation over the lake. Stable isotope measurements thus improve the understanding of the circulation of water within the lake, which is fundamental for an evaluation of the water residence times, dissolved pollutant and nutrient transport as well as oxygenation.

Geological structure, recharge processes and underground drainage of a glacierised karst aquifer system, Tsanfleuron-Sanetsch, Swiss Alps

The relationships between stratigraphic and tectonic setting, recharge processes and underground drainage of the glacierised karst aquifer system `Tsanfleuron-Sanetsch' in the Swiss Alps have been studied by means of various methods, particularly tracer tests (19 injections). The area belongs to the Helvetic nappes and consists of Jurassic to Palaeogene sedimentary rocks. Strata are folded and form a regional anticlinorium. Cretaceous Urgonian limestone constitutes the main karst aquifer, overlain by a retreating glacier in its upper part. Polished limestone surfaces are exposed between the glacier front and the end moraine of 1855/1860 (Little Ice Age); typical alpine karrenfields can be observed further below. Results show that (1) large parts of the area are drained by the Glarey spring, which is used as a drinking water source, while marginal parts belong to the catchments of other springs; (2) groundwater flow towards the Glarey spring occurs in the main aquifer, parallel to stratification, while flow towards another spring crosses the entire stratigraphic sequence, consisting of about 800 m of marl and limestone, along deep faults that were probably enlarged by mass movements; (3) the variability of glacial meltwater production influences the shape of the tracer breakthrough curves and, consequently, flow and transport in the aquifer.

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.

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.

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.

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.

Quantifying sediment storage in a high Alpine valley (Turtmanntal, Switzerland)

The determination of sediment storage is a critical parameter in sediment budget analyses. But, in many sediment budget studies the quantification of magnitude and time-scale of sediment storage is still the weakest part and often relies on crude estimations only, especially in large drainage basins (>100km2). We present a new approach to storage quantification in a meso-scale alpine catchment of the Swiss Alps (Turtmann Valley, 110km2). The quantification of depositional volumes was performed by combining geophysical surveys and geographic information system (GIS) modelling techniques. Mean thickness values of each landform type calculated from these data was used to estimate the sediment volume in the hanging valleys and the trough slopes. Sediment volume of the remaining subsystems was determined by modelling an assumed parabolic bedrock surface using digital elevation model (DEM) data. A total sediment volume of 781·3×106?1005·7×106m3 is deposited in the Turtmann Valley. Over 60% of this volume is stored in the 13 hanging valleys. Moraine landforms contain over 60% of the deposits in the hanging valleys followed by sediment stored on slopes (20%) and rock glaciers (15%). For the first time, a detailed quantification of different storage types was achieved in a catchment of this size. Sediment volumes have been used to calculate mean denudation rates for the different processes ranging from 0·1 to 2·6mm/a based on a time span of 10ka. As the quantification approach includes a number of assumptions and various sources of error the values given represent the order of magnitude of sediment storage that has to be expected in a catchment of this size.

Introgressive hybridization of threatened European tree frogs (Hyla arborea) by introduced H. intermedia in Western Switzerland

Hybridization by introduced taxa is a major threat to native species. Characterizing human introductions is thus one of the missions of conservation geneticists. Here we survey a declining population of the regionally endangered European tree frog (Hyla arborea) in the Grangettes natural reserve (Rhone valley, Western Switzerland), where previous evidence indicated human introduction of the Italian taxon H. intermedia. We combined fast-evolving mitochondrial and nuclear markers and an extended sampling to conduct population genetic analyses of the Grangettes and putative source areas. We show that the Grangettes population is a hybrid swarm, with all individuals featuring recent nuclear admixture and mitochondrial DNA of introduced H. intermedia, most likely of proximate south Alpine origin. In contrast, H. arborea and H. intermedia hardly introgress in their natural parapatric ranges, consistent with an advanced reproductive isolation. Thus, potential hybrid incompatibilities may account for the strong decline of this population, despite important conservation efforts. Although their hybrid nature makes them a priori unworthy of any protection, we propose specific measures to recover local H. arborea gene pool and preserve tree frogs in the Grangettes, the last population remaining from this heavily impacted part of the Alps.

Accessory phases as a tool to investigate magmatic underplating in polymetamorphic metasediments : an example from a fossil lower crust : Ivrea Verbano zne, NW Italy

The Ivrea and the Strona-Ceneri zones, NW italy and S Switzerland, offer the possibility to study the continental crust of the Southern Alps. Because of its high metamorphic degree and the abundant Permo- Carboniferous mafic intrusions, the Ivrea Zone is classically interpreted an exposed section trough the Permian lower crust. The present work is focused here on metasedimentary slices (septa) intercalated within Permian gabbro (mafic complex). In particular I studied the evolution of accessory phases such as rutile and zircon and the chemistry of the metasediments. The septa build an irregular and discontinuous band that cut obliquely the mafic complex from its deepest part (N) to its roof (S). The chemistry of the metasediments evolves along the band and the chemical evolution can be compared with that observed in the country-rock surrounding the mafic intrusion to the NE and overprinted by a main regional metamorphic event. This suggests that the degree of chemical depletion of the septa was mainly established during the same regional metamorphic event. Moreover it suggests that incorporation of the septa within the gabbro did not modify their original stratigraphie distribution within the crust. It implies that the mafic complex has been emplaced following a dynamic substantially different from the classic model of « gabbro glacier » (Quick et al., 1992; Quick et al., 1994). It is more likely that it has been emplaced by repeated injections of sills at different depths during a protracted period of time. Zircon trace elements and U-Pb ages suggest that regional metamorphism occurred 330-320Ma, the first sills in the deepest part of the Mafic Complex are injected at ~300Ma, the mafic magmas reached higher levels in the crust at 285Ma and the magmatic activity continued locally until 275Ma. The ages of detrital cores in zircons fix the maximal sedimentation age at ~370Ma, this age corresponds therefore with the maximal age of the incorporation of the Ivrea zone within the lower crust. I propose that the Ivrea zone has been accreted to the lower crust during the Hercynian orogeny sensu lato. The analysis of detrital ages suggests that the source terrains for the Ivrea zone and those for the Strona-Ceneri zone have a completely different Palaeozoic history. The systematic analysis of rutile in partially molten metasediments of the Ivrea zone reveals the occurrence of two generations. The two generations are characterized by a different chemistry and textural distribution. A first generation is formed during pro-grade metamorphism in the restitic counterpart. The second generation is formed in the melts during cooling at the same time that part of the first generation re-equilibrate. Re-equilibration of the first generation seems to be spatially controlled by the presence of fluids. Locally the second generation forms overgrowths on the first generation. Considered the different diffusivity of U and Pb in rutile, U heterogeneities have important implication for U-Pb dating of rutile. ID-TIMS and LA-ICPMS dating coupled with a careful textural investigation (SEM) suggest that rutile grains are characterized by multiple path along which Pb diffusion can occur: volume diffusion is an important process, but intragrain and subgrain boundaries provide additional high diffusivity pathways for Pb escape and reduce drastically the effective diffusion length. -- La zone d'Ivrea et la zone de Strona-Ceneri, en Italie nord-occidentale et Suisse méridionale, offrent la possibilité d'étudier la croûte continentale des Alpes du Sud. En raison du haut degré métamorphique et l'abondance d'intrusions mafiques d'âge Permo-Carbonifère [complexe mafique), la zone d'Ivrea est interprétée classiquement comme de la croûte inférieure permienne. Ce travail ce concentre sur des bandes metasédimentaires (septa) incorporées dans les magmas mafiques lors de l'intrusion. Les septa forment une bande irrégulière qui coupe obliquement le complexe mafique du bas (N) vers le haut (S). La chimie des septa évolue du bas vers le haut et l'évolution chimique se rapproche de l'évolution observé dans la roche encaissante l'intrusion affecté par un événement métamorphique régionale. Cette relation suggère que le degré d'appauvrissement chimique des septa a été établit principalement lors de l'événement métamorphique régional. De plus l'incorporation dans les gabbros n'a pas perturbée la distribution stratigraphique originelle des septa. Ces deux observations impliquent que le métamorphisme dans la roche encaissante précède la mise en place du gabbro et que cette dernière ne se fait pas selon le modèle classique (« gabbro glacier » de Quick et al., 1992, 1994), mais se fait plutôt par injections répétées de sills a différentes profondeurs. Les âges U-Pb et les éléments traces des zircons suggèrent que le métamorphisme régionale a eu lieu 330-320Ma, alors que les premiers sills dans la partie profonde du Mafic Complex s'injectent à ~300Ma, le magmatisme mafique atteigne des niveaux supérieurs à 285Ma et continue localement jusqu'à 270Ma. Les âges des coeurs détritiques des zircons permettent de fixer l'âge maximale de sédimentation à ~370Ma ce qui correspond donc à l'âge maximale de l'incorporation de la zone d'Ivrea dans la croûte inférieur. L'analyse systématique des rutiles, nous a permit de montrer l'existence de plusieurs générations qui ont une répartition texturale et une chimie différente. Une génération se forme lors de l'événement UHT dans les restites, une autre génération se forme dans les liquides lors du refroidissement, au même temps qu'une partie de la première génération se rééquilibre au niveau du Zr. Localement la deuxième génération peut former des surcroissances autour de la première génération. Dans ces cas, des fortes différences en uranium entre les deux générations ont des importantes implications pour la datation U-Pb sur rutile. Classiquement les ratios Pb/U dans le rutile sont interprétés comme indiquant l'âges du refroidissement du minéral sous une température à la quelle la diffusion du Pb dans le minéral n'est plus détectable et la diffusion à plus hautes températures est assumée se faire par «volume diffusion» dans le grain (Mezger et al., 1989). Par des datations ID-TIMS (sur grain entier) et LA-ICPMS (in-situ) et une analyse texturale (MEB) approfondie nous montrons que cette supposition est trop simpliste et que le rutile est repartie en sous-domaines. Chacun de ces domaines a ça propre longueur ou chemin de diffusion spécifique. Nous proposons donc une nouvelle approche plus cohérente pour l'interprétation des âges U-Pb sur rutile.

Evolution of the Helvetic shelf (Switzerland) during the Barremian-early Aptian : paleoenvironmental, paleogeographic and paleoceanographic controlling factors

The Early Cretaceous has experienced the development of large shallow-water carbonate platform in tropical and subtropical regions, favoured by exceptionally warm climatic conditions, optimal trophic conditions and a suitable tectonic and paleogeographic context. This period was also characterized by shorter intervals, in which the widespread deposition of marine sediments enriched in organic matter occurred ("oceanic anoxic episodes": OAE). This study focuses on the Barremian- Aptian interval, during which the Urgonian platform developed throughout the northern Tethyan passive margin. Due to the Alpine orogeny, sediments belonging to this platform - named locally Schrattenkalk Formation, are presently outcropping in the Helvetic Alps. This study aims to reconstruct the paleogeographic evolution of the Helvetic platform, and to define the environmental and oceanographic factors, which influenced its development. Several key episodes in the life of this platform have been identified: - The installation of the platform, covering hemipelagic sediments of the Drusberg Member, near the limit between the early and late Barremian. - The temporary change of carbonate production type during the basal Aptian, with the deposition of the Rawil Member. - And finally the definitive interruption of photozoan carbonate platform sedimentation in the study area, during the early Aptian. The sedimentological, biostratigraphical and chemostratigraphic (8I3C) data lead to the sequential subdivision of eleven sections and one core, located throughout the different Helvetic nappes of Switzerland. The sequence stratigraphie framework, initially defined for the Urgonian carbonate platform of the Vercors area (SE France), is confirmed in the Helvetic nappes, where the same number of sequences was observed. Many similarities between these two areas are put forward in this work. The sequence stratigraphie framework helped to highlight the installation of a bioclastic body, included in the Schrattenkalk Formation, since the middle Early Barremian (sequence B2). The age of the installation of the rudist-rich limestone, which corresponds to the Urgonian facies sensu stricto, is attributed to the late Barremian (maximum flooding surface of the sequence B3). This age coincides with the one determined in other northern Tethyan areas for the installation of the Urgonian platform. The results of this study show a strong tectonic control of the platform architecture, with the presence of syn-sedimentary faults in a perpendicular position to the progradation direction of the platform. The presence of these faults was highlighted by the study of the evolution of the microfacies distribution and by thickness variations in different areas. Sea level fluctuations also played an important role in the various life phases of the platform. Three major falls in sea level have been identified. A significant emersion of the proximal domain has been observed, involving an important drop of the relative sea level, leading to the exposure of the Drusberg Member hemipelagic series. A second major drop in sea level is identified near the Barremian-Aptian boundary, and a third is registered on the top of the Upper Schrattenkalk Member on the whole platform; it is associated with a karst affecting the underlying limestones to a depth of over 20 meters. This observation sheds new light on the conditions linked to the demise of Urgonian platform, which was strongly influenced by this phase of emersion.

Distribution and population genetic variation of cryptic species of the Alpine mayfly Baetis alpinus (Ephemeroptera: Baetidae) in the Central Alps.

BACKGROUND: Many species contain evolutionarily distinct groups that are genetically highly differentiated but morphologically difficult to distinguish (i.e., cryptic species). The presence of cryptic species poses significant challenges for the accurate assessment of biodiversity and, if unrecognized, may lead to erroneous inferences in many fields of biological research and conservation. RESULTS: We tested for cryptic genetic variation within the broadly distributed alpine mayfly Baetis alpinus across several major European drainages in the central Alps. Bayesian clustering and multivariate analyses of nuclear microsatellite loci, combined with phylogenetic analyses of mitochondrial DNA, were used to assess population genetic structure and diversity. We identified two genetically highly differentiated lineages (A and B) that had no obvious differences in regional distribution patterns, and occurred in local sympatry. Furthermore, the two lineages differed in relative abundance, overall levels of genetic diversity as well as patterns of population structure: lineage A was abundant, widely distributed and had a higher level of genetic variation, whereas lineage B was less abundant, more prevalent in spring-fed tributaries than glacier-fed streams and restricted to high elevations. Subsequent morphological analyses revealed that traits previously acknowledged as intraspecific variation of B. alpinus in fact segregated these two lineages. CONCLUSIONS: Taken together, our findings indicate that even common and apparently ecologically well-studied species may consist of reproductively isolated units, with distinct evolutionary histories and likely different ecology and evolutionary potential. These findings emphasize the need to investigate hidden diversity even in well-known species to allow for appropriate assessment of biological diversity and conservation measures.

The expression of the end-Cenomanian oceanic anoxic event (OAE 2) in the Helvetic Alps

Upper Cenomanian pelagic sediments from the northern Alpine Helvetic fold-and-thrust belt (northern Tethyan margin) coeval with Oceanic Anoxic Event (OAE) 2 are characterized by the temporal persistence of micrite sedimentation and lack of organic carbon-rich layers. We studied an expanded section in the Chrummflueschlucht (east of Euthal, Switzerland), which encompasses the OAE 2 time interval. In order to identify the paleoceanographic and paleoenvironmental conditions during OAE 2 in this part of the northern Tethyan margin, and more specifically to trace eventual changes in nutrient levels and oxic conditions, we investigated the biostratigraphy (planktonic foraminifera), the bulk-rock mineralogy, and measured stable carbon- and oxygen-isotopes, total phosphorus (P) and redox-sensitive trace-element (RSTE) contents. We were able to determine – with some remaining uncertainties – the different planktonic foraminiferal biozones characteristic of the Cenomanian–Turonian boundary interval (Rotalipora cushmani, Whiteinella archaeocretacea and Helvetoglobotruncana helvetica zones). In the lower part of the section (R. cushmani total range zone), the bulk-rock δ13C record shows a long-term increase. Within sediments attributed to the W. archaeocretacea partial range zone, δ13C values reach a maximum of 3.3‰ (peak “a”). In the following the values decrease and increase again to arrive at a plateau with high δ13C values of around 3.1‰, which ends with a peak of 3.3‰ (peak “c”). At the top of the section, in sediments belonging to the H. helvetica total range zone, δ13C values decrease to post-OAE values of around 2.2‰. The last occurrence of R. cushmani is observed just above the positive δ13C shift characterizing OAE 2. P contents display small variations along the section with a long-term decreasing trend towards the top. Before the OAE 2 interval, P values show higher values and relatively good covariation with detrital input, indicating higher nutrient input before OAE 2. In sediments corresponding to the onset of the δ13C positive excursion, P content is marked by a sharp peak probably linked to a slowdown in sedimentation rates and/or the presence of a small hiatus, as is shown by the presence of glauconite and phosphatic grains. In the interval corresponding to OAE 2, P values remain low and increase slightly at the end of the positive shift in the δ13C record (in the H. helvetica total range zone). The average contents of RSTE (U, V, As, Co, Mo and Mn) remain low throughout the section and appreciable RSTE enrichments have not been observed for the sedimentary interval corresponding to OAE 2. No correlation is observed with stratigraphic trends in RSTE contents in organic-rich deeper-water sections. The presence of double-keeled planktonic foraminifera species during most of the Cenomanian/Turonian boundary event is another evidence of relatively well-oxygenated conditions in this part of the northern Tethyan outer shelf. Our results show that the Chrummflueschlucht section corresponds to one of the most complete section for the Cenomanian–Turonian boundary interval known from the Helvetic realm even if a small hiatus may be present at the onset of the δ13C record (peak “a”). The evolution of P contents suggests an increase in input of this nutritive element at the onset of OAE2. However, the trends in RSTE contents and the planktonic foraminifera assemblages show that the Helvetic realm has not been affected by strongly depleted oxygen conditions during OAE 2.

Recent Glacier Retreat Exceeds Internal Variability

Glacier fluctuations exclusively due to internal variations in the climate system are simulated using downscaled integrations of the ECHAM4/OPYC coupled general circulation model (GCM). A process-based modeling approach using a mass balance model of intermediate complexity and a dynamic ice flow model considering simple shearing flow and sliding are applied. Multimillennia records of glacier length fluctuations for Nigardsbreen (Norway) and Rhonegletscher (Switzerland) are simulated using autoregressive processes determined by statistically downscaled GCM experiments. Return periods and probabilities of specific glacier length changes using GCM integrations excluding external forcings such as solar irradiation changes, volcanic, or anthropogenic effects are analyzed and compared to historical glacier length records. Preindustrial fluctuations of the glaciers as far as observed or reconstructed, including their advance during the “Little Ice Age,” can be explained by internal variability in the climate system as represented by a GCM. However, fluctuations comparable to the present-day glacier retreat exceed any variation simulated by the GCM control experiments and must be caused by external forcing, with anthropogenic forcing being a likely candidate.