The larva of Rhyacophila ferox Graf, 2006 (Trichoptera: Rhyacophilidae) from the Eastern Alps (Carinthia,Austria)

The hitherto unknown larva of Rhyacophila ferox Graf, 2006, is described and discussed in the context of contemporary Rhyacophilidae keys. In addition, zoogeographical and ecological notes are included.     

Origin and seasonality of subfossil caprine dung from the discovery site of the Iceman (Eastern Alps)

The discovery of a Neolithic glacier mummy (dated to 3300–3100 cal b.c.) on a remote site of an Alpine pass at 3,200 m in the Ötztal Mountains is still puzzling. In the initial phase of the Iceman research, four hypotheses were suggested to explain the find in its entirety. The speculations vary from a hunter or warrior to a shaman, a miner or a shepherd. None of these proposals is accepted or corroborated by archaeological findings, but on the basis of palynological investigations conducted in the vicinity of the discovery site the assumption that the Iceman was involved in an early form of transhumance has now gained general acceptance. Concerning this assumption we present in this paper a recent study conducted on about a hundred caprine (sheep/goat or ibex/chamois) dung pellets recovered from the find spot of the Iceman and which were dated from 5400 to 2000 cal b.c. The approach was to determine through plant remains from these faeces whether they were droppings derived from animals grazing in anthropogenic habitats at low altitudes or in alpine grasslands. The former case would suggest they were livestock, the latter game. The results showed that all droppings derive from animals grazing at high altitudes.     

Sympatric diploid and hexaploid cytotypes of Senecio carniolicus (Asteraceae) in the Eastern Alps are separated along an altitudinal gradient

We explored the fine-scale distribution of cytotypes of the mountain plant Senecio carniolicus along an altitudinal transect in the Eastern Alps. Cytotypes showed a statistically significant altitudinal segregation with diploids exclusively found in the upper part of the transect, whereas diploids and hexaploids co-occurred in the lower range. Analysis of accompanying plant assemblages revealed significant differences between cytotypes along the entire transect but not within the lower part only, where both cytotypes co-occur. This suggests the presence of ecological differentiation between cytotypes with the diploid possessing the broader ecological niche. No tetraploids were detected, indicating the presence of strong crossing barriers. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

An upper Tuvalian (Triassic) platform-basin system in the Julian Alps: the start-up of the Dolomia Principale (Southern Alps,Italy)

Summary East of Cave del Predil (formerly Raibl), a platform-basin transition of the Dolomia Principale (Hauptdolomit) is spectacularly exposed at a seismic scale. Therefore, the eastern margin of the vast domain of the Dolomia Principale, facing the Slovenian Basin, is documented. Despite of strong dolomitization of the massive margin, some sedimentary structures and fossils have been recognized. Corals seem to be very rare and sponges to be absent, whereas serpulids and marine phreatic cements seem to have been the main components of the framework. Interior platform bedded dolomites lap off the massive margin. Clinoforms interfinger with upper Tuvalian basinal deposits (Carnitza Formation). This setting documents the start-up of the Dolomia Principale during late Tuvalian time. Moreover, this margin of the Dolomia Principale is the more ancient so far pointed out. It testifies to the recovery of a rimmed platform after the late Julian-early Tuvalian crisis.     

Facies and biota of Anisian to Carnian carbonate platforms in the Northern Calcareous Alps (Tyrol and Bavaria)

Summary During the Middle and early Late Triassic carbonate ramps and rimmed platforms developed at the northwestern margin of the Tethys ocean. In the Northern Calcareous Alps, Anisian stacked homoclinal ramps evolved through a transitional stage with distally steepened ramps to huge rimmed platforms of Late Ladinian to Early Carnian age. Middle Triassic to early Late Triassic facies and biota of basin, slope and platform depositional systems are described. Special emphasis is given to foraminifers, sponges, microproblematic organisms and algae. The Ladinian to early Carnian reef associations are characterized by the abundance of segmented sponges, microproblematica, biogenic crusts and synsedimentary cements. Among the foraminifers, recifal forms likeHydrania dulloi andCucurbita infundibuliformis (Carnian in age) are reported from the Northern Calcareous Alps for the first time. Some sphinctozoid sponges likeParavesicocaulis concentricus were known until now only from the Hungarian and Russian Triassic.     

Palaeoecology of solitary corals in soft-substrate habitats: the example of Cunnolites (upper Santonian, Eastern Alps)

The upper Santonian Hofergraben Member (Eastern Alps) provides an example of a soft‐substrate habitat suited mainly for solitary corals (Cunnolites), for colonial forms of solitary coral‐like shape (Placosmilia, Diploctenium), and for colonial corals of high sediment resistance (e.g. Actinacis, Pachygyra). The Hofergraben Member consists mainly of silty‐sandy marls of wave‐dominated, low‐energy shore zone to shallow neritic environments. Substrates of soft to firm mud supported level‐bottoms of non‐rudist bivalves, gastropods, solitary corals, colonial corals, rudists, echinoids, and benthic foraminifera. Boring and/or encrustation of fossils overall are scarce. In the marls, Cunnolites is common to abundant. Both a cupolate shape and a lightweight construction of the skeleton aided the coral to keep afloat soft substrata. Cunnolites taphocoenoses are strongly dominated by small specimens (about 1–3 cm in diameter). Cunnolites was immobile and mostly died early in life upon, either, smothering during high‐energy events, rapid sedimentation associated with river plumes, or by toppling and burial induced by burrowing. Comparatively few large survivor specimens may show overgrowth margins interpreted as records of partial mortality from episodic sedimentation or tilting on unstable substrate. Scattered pits and scalloped surfaces on large Cunnolites may have been produced, in some cases at least, by predators (durophagous fish?). Post‐mortem, large Cunnolites provided benthic islands to corals, epifaunal bivalves and bryozoans. In a single documented case of probable in vivo contact of Cunnolites with the colonial coral Actinastraea, the latter prevailed.     

Robust sequential Factor Analysis of geochemical data from the Permian-Triassic Gartnerkofel-1 core (Southern Alps): the geochemical response to changing paleo-oceanographic conditions in shallow-marine carbonate platforms

The Gartnerkofel-1 core provides a high-quality multi-element dataset that characterizes an Upper Permian to Lower Triassic shallow-marine carbonate sequence (Bellerophon and Werfen Formations) of the Carnic Alps (Southern Alps). Based on the well-known sedimentological evolution, robust sequential Factor Analysis is explored as a multivariate statistical technique to understand geochemical processes in carbonate platforms. The results demonstrate that 93% of the whole-rock compositional variability of the Gartnerkofel-1 core can be explained by the detrital input that is diluted by the carbonate production and the early diagenetic redox state. Two stages of anoxia, one at the Permian/Triassic boundary and one in the Mazzin Member of the Werfen Formation, are related to indicative factor scores. The factor scores within this interval suggest an enhanced dolomitization of shales and marls, a mobilization of manganese, and an accumulation of syndiagenetically precipitated pyrite.     

Late Triassic sedimentary evolution of Slovenian Basin (eastern Southern Alps): description and correlation of the Slatnik Formation

Successions of the Slovenian Basin structurally belong to the easternmost Southern Alps. During the Late Triassic, they were part of the Adriatic continental margin. Norian–Rhaetian successions of the Slovenian Basin are characterized mainly by dolomite of the Bača Dolomite Formation. However, in the northern part of the basin, Late Triassic limestone is preserved above Bača Dolomite Formation and is formalized as the Slatnik Formation. It is composed of hemipelagic limestone alternating with resedimented limestones. The succession documents an upward progradation of the slope environment composed of three high-frequency cycles. Most prominent progradation is referred to the second, i.e., Early Rhaetian cycle. The Slatnik Formation ends with thin-bedded hemipelagic limestone that records the end-Triassic productivity crisis, or rapid sea-level fall. The overlying resedimented limestones of the Early Jurassic Krikov Formation, document the recovery of production and shedding from the adjacent carbonate platform.     

Middle to Late Holocene vegetation history of the Upper Engadine (Swiss Alps): the role of man and fire

To reconstruct the vegetation and fire history of the Upper Engadine, two continuous sediment cores from Lej da Champfèr and Lej da San Murezzan (Upper Engadine Valley, southeastern Switzerland) were analysed for pollen, plant macrofossils, charcoal and kerogen. The chronologies of the cores are based on 38 radiocarbon dates. Pollen and macrofossil data suggest a rapid afforestation with Betula, Pinus sylvestris, Pinus cembra, and Larix decidua after the retreat of the glaciers from the lake catchments 11,000 cal years ago. This vegetation type persisted until ca. 7300 cal b.p. (5350 b.c.) when Picea replaced Pinus cembra. Pollen indicative of human impact suggests that in this high-mountain region of the central Alps strong anthropogenic activities began during the Early Bronze Age (3900 cal b.p., 1950 b.c.). Local human settlements led to vegetational changes, promoting the expansion of Larix decidua and Alnus viridis. In the case of Larix, continuing land use and especially grazing after fire led to the formation of Larix meadows. The expansion of Alnus viridis was directly induced by fire, as evidenced by time-series analysis. Subsequently, the process of forest conversion into open landscapes continued for millennia and reached its maximum at the end of the Middle Ages at around 500 cal b.p. (a.d. 1450).     

Internal Disorder: Post-Mortem Burrows within the Tests of the Holasteroid Echinoid Echinocorys Leske (Upper Cretaceous to Paleocene)

A test of the Lower Paleocene holasteroid echinoid Echinocorys ex. gr. scutata Leske from Göynük, Turkey, preserves a Planolites beverleyensis (Billings) burrow close to the internal surface. Identification of such a specimen is fortuitous; only where the test has broken cleanly away from the internal mould can such burrows be exposed, if present at all. A second specimen, from the Upper Cretaceous of Norfolk, England, bears a sinuous boring identified as Planolites montanus Richter.     

Paleoenvironmental and diagenetic implications of δO and δC isotope ratios from the Upper Jurassic Plassen limestone (Northern Calcareous Alps, Austria)

The first δ¹⁸O and δ¹³C data from the Upper Jurassic of the Northern Calcareous Alps are presented. The interpretation of stable isotope ratios serves as an approach for paleoenvironmental and diagenetic studies of the Plassen carbonate platform, which cannot be obtained by paleontological methods and microfacies analyses alone. The studied part of the Plassen limestone is characterized by (1) lithoclast facies, also called ‘intraformational breccia’; the origin of lithoclasts was formerly unknown; (2) peloid facies; (3) bioclastic facies, composed of peloids, porostromate algae, green algae and red algae; and (4) oncoid facies. Two types of fracturing and four cement generations can be distinguished. Isotope ratios of the matrix, oncoids, three cement generations and whole rock samples revealed that (1) the studied section represents an open marine carbonate platform with high water circulation and high input of cool oceanic waters; (2) the platform was not affected by emersion and fresh water influence; normal marine conditions prevailed; (3) carbonate cements were precipitated in a closed diagenetic system, but burial diagenesis was absent; (4) both fabric-selective and non-fabric-selective fracturing occurred in a normal marine environment, affecting the formation of ‘intraformational breccias’.     

Limb bone histology and growth in Placerias hesternus (Therapsida: Anomodontia) from the Upper Triassic of North America

Abstract: Patterns of bone deposition are reported and deduced from mid‐shaft sections of 21 limb bones of the dicynodont Placerias hesternus from the Placerias Quarry (Upper Triassic), Arizona, USA. All sampled elements of P. hesternus have a large medullary cavity completely filled with bony trabeculae surrounded by dense cortical bone. Dense Haversian bone extends from the perimedullary region to at least the mid‐cortex in all sampled bones. Primary bone in the outer cortex of limb elements of P. hesternus is generally zonal fibrolamellar with a peripheral layer of parallel‐fibred bone. These data suggest periodic rapid osteogenesis followed by slower growth. Among dicynodonts, this strategy is most similar to growth previously reported in other Triassic (Lystrosaurus, Wadiasaurus) and some Permian taxa (Oudenodon, Tropidostoma). An external fundamental system (EFS), suggesting complete or near complete cessation of appositional growth, is present in the largest tibia. This is the first report of EFS in dicynodonts and may represent the attainment of maximum size in P. hesternus. Slow‐growing peripheral bone was observed in elements of varying size in our sample and may support a differential growth pattern between P. hesternus individuals from this locality. A complete growth series of P. hesternus, analysis of Placerias specimens from other localities, and further sampling of other Upper Triassic dicynodonts are needed to better understand a more complete picture of the growth and remodelling patterns that we have initially investigated.     

Microfacies and depositional environment of an Upper Triassic intra-platform carbonate basin: the Fatric Unit of the West Carpathians (Slovakia)

Facies associations of the Rhaetian Fatra Formation from the Veká Fatra Mts. (West Carpathians) were deposited in a storm-dominated, shallow, intra-platform basin with dominant carbonate deposition and variable onshore peritidal and subtidal deposits, with 21 microfacies types supported by a cluster analysis. The deposits are formed by bivalves, gastropods, brachiopods, echinoderms, corals, foraminifers and red algae, ooids, intraclasts and peloids. A typical feature is the considerable variation in horizontal direction. The relative abundance and state of preservation of components as well as the fabric and geometric criteria of deposits can be correlated with depth/water energy-related environmental gradients. Four facies associations corresponding to four types of depositional settings were distinguished: a) peritidal, b) shoreface, above fair-weather wave base (FWWB), c) shallow subtidal, above normal storm wave base and d) above maximum storm wave base. The depositional environment can be characterized as a mosaic of low-relief peritidal flats and islands, shoreface banks and bars, and shallow subtidal depressions. The distribution and preservation of components were mainly controlled by the position of base level (FWWB), storm activity and differences in carbonate production between settings. Poorly or moderately diverse level-bottom macrobenthic assemblages are dominated by molluscs and brachiopods. The main site of patch-reef/biostrome carbonate production was located below the fair-weather wave base. Patch-reef/biostrome assemblages are poorly diverse and dominated by the branched scleractinian coral Retiophyllia, forming locally dm-scale autochthonous aggregations or more commonly parautochthonous assemblages with evidence of storm-reworking and substantial bioerosion by microborings and boring bivalves.Facies types and assemblages are comparable in some aspects to those known from the Upper Triassic of the Eastern and Southern Alps (Hochalm member of the Kössen Formation or Calcare di Zu Formation), pointing to similar intra-platform depositional conditions. The absence of large-scale patch-reefs and poor diversity of level-bottom and patch-reef/biostrome assemblages with abundance of eurytopic taxa indicate high-stress/unstable ecological conditions and more restricted position of the Fatric intra-platform setting from the open ocean than the intra-platform habitats in the Eastern or Southern Alps.     

Expansion of Brown Bears (Ursus arctos) into the Eastern Alps: A Spatially Explicit Population Model

We present a spatially explicit population model for analysing the expansion of brown bears (Ursus arctos) after the reintroduction program in central Austria. The model is based on field investigations into brown bears in Austria and Slovenia and on current knowledge of brown bears. The landscape of the eastern Alps is represented by a GIS-derived raster map defining local habitat suitability and five major spatial barriers to dispersal. The population model follows the fate of individual bears and simulates reproduction, dispersal, home range establishment, and mortality in annual time steps. We indirectly adjust unknown or uncertain model parameters with 10-year data on the number of females with cubs in central Austria and determine key variables of population dynamics, such as population sizes and growth rates within different population nuclei, dispersal distances, or mortality rates, for model parameterisations that reproduce the data on females with cubs. We estimated a current (1996–2000) growth rate of the population in Austria and adjacent parts of Italy of some 14%; a high proportion of this growth was due toimmigration from Slovenia. Consequently, the growth rate of the subpopulation in central Austria, which probably is isolated functionally (i.e., no exchange of females) from the nuclei along the Austrian–Slovenian border, yielded some 7%. This subpopulation may comprise seven residents, and we estimated for females a 33% risk of extinction during the 1992–2000 period. Validation and confirmation of our model results with data on bear densities that were not used for model construction and parameterisation supported our findings. The high female mortality rates, together with the vulnerability of the small population to chance events (i.e., demographic stochasticity), are the most pressing threat for the population in the eastern Alps. Our approach could be widely applied for analysing dynamics of rare and endangered species in which the paucity of data precludes an appraisal of the state of the population using standard methods.     

A sequence stratigraphic approach to a Middle Triassic shelf-slope complex of the Ligurian Alps (Ligurian Briançonnais,Monte Carmo-Rialto unit,Italy)

The Mesozoic sedimentary cover belonging to the Monte Carmo-Rialto unit of the Ligurian Briançonnais domain is composed of Scythian clastics and Anisian to Carnian carbonate rocks over 300 m thick. This paper focuses on the stratigraphy of this carbonate complex, its environmental significance, and its evolution in light of dynamic stratigraphy. Our facies analysis of limestones and dolomites of the Triassic complex allowed us to reconstruct an environmental model. Data support a distally steepened carbonate ramp of Anisian age evolving to a more diversified Ladinian platform with an oolitic sand-bar belt separating the lagoon from the slope. The Monte Carmo-Rialto slope facies are the only witnesses of deep sedimentation in the Triassic terrains of the Ligurian Briançonnais domain, otherwise represented by shallow-water carbonate deposits. On the basis of facies succession, we have identified nine medium-scale cycles (third-order sequences) in the study area, comparable to those evidenced in the Briançonnais s.s. domain by the French authors. Small-scale cycles analysis evidenced mainly shallowing-upward trends in the examined sequences; although a few evidences of transgression-related deposits (deepening upward cycles) have been found at the base three sequences, they have been mostly obliterated by dolomitization and masked by local tectonics. For this reason, we can undoubtedly distinguish only the part of each sequence belonging to HST, while the TST, though present, still remains a partition that cannot be precisely characterized. In the same way, LSTs are not present in the Monte Carmo-Rialto unit, due to the original relative landward position of the examined area. Sequence stratigraphy analysis indicates different long-term dynamics for the two evolutionary stages of the Triassic Ligurian platform: a general landward backstepping to moderate progradation during the Early Anisian and true progradation during the latest Anisian and Ladinian. In addition, a good fit with the sequences proposed by the SEPM chart has been found, indicating a correspondence for the third-order sequences of the Middle Triassic.     

High abundance of Myriochele fragilis Nilsen & Holthe, 1985 (Polychaeta: Oweniidae) in the deep sea of the Eastern Mediterranean

An exceptionally high abundance of Myriochele fragilis Nilsen & Holthe, 1985 (Polychaeta: Oweniidae) is reported for the Ierapetra Basin in the Eastern Mediterranean Sea from depths of 4159 and 4260 m, representing the first record outside the Norwegian and Arctic Sea and extending the depth range by more than 1600 m. Specimens are described and compared to closely related species.     

Non-marine biota from the Lower Permian of the central Southern Alps (Orobic and Collio basins, N Italy): a key to the paleoenvironment

The Lower Permian in the central Southern Alps yields an important low-diversity fossil assemblage which was deposited in a varied continental setting, showing mainly alluvial fan to lacustrine and, locally, playa-like floodplain environments. The present study is not taxonomical and its objective is to make a first report of new invertebrate organisms and trackways discovered in the Orobic and Collio basins. Such a fossil record, which comprises freshwater jellyfishes, arthropod tracks, stromatolites, algae and other organisms, will further our knowledge about the local and regional geological history of the central-western Southern Alps and improve our understanding of Early Permian palaeoenvironments. On the whole, in both intramontane basins the ichnodiversity and fossil content decreases from a stratigraphic lower portion, mainly lacustrine and alluvial, towards an upper one, characterised by coarse alluvial deposits to floodplain fines. On the basis of both the already known palaeontological data from the two basins, mainly macroflora and tetrapod footprint associations, and the recently discovered taxa, we tried to make reliable palaeoenvironmental inferences and, possibly, hypothesise on a climatic change, which could have occurred during the Early Permian.     

A three- and two-dimensional documentation of structural elements in schwagerinids (superfamily Fusulinoidea) exemplified by silicified material from the Upper Carboniferous of the Carnic Alps (Austria/Italy): a comparison with verbeekinoideans and alveolinids

Silicified schwagerinids (superfamily Fusulinoidea v. Moeller 1878) from the Upper Carboniferous (Carnic Alps, Austria and Italy) were isolated from cemented carbonate rocks using hydrochloric acid. The shells show details of the wall texture and of internal structures in three dimensions which are illustrated with SEM pictures. Thin sections from hand specimens provided two-dimensional sections of the shell for comparison. The functional significance of fusulinoidean internal structures is discussed and compared with verbeekinoideans and alveolinids. Particular attention is paid on the disposition of the different openings within the shell and from the chamber lumen to the outside which reflects the direction of protoplasmic flow. Based on the knowledge of the nature of protoplasm ultrastructure in Recent foraminifera and its biological significance we draw some conclusions about the nature of protoplasm in fusulinoideans and its change within the Permian verbeekinoideans.