Tectonically controlled late cretaceous terrestrial to neritic deposition (Northern Calcareous Alps, Tyrol, Austria)

Summary The Turonian to Santonian terrestrial to neritic succession (Lower Gosau Subgroup) in the Northern Calcareous Alps of the eastern part of the Tyrol, Austria, provides an example for deposition on a compartmentalized, narrow, microtidal to low-mesotidal, wave-dominated, mixed siliciclastic-carbonate shelf. The shelf was situated in front of a mainland with a relatively high, articulated relief, and underwent distinct changes in facies architecture mainly as a result of tectonism. The investigated succession was deposited above a deeply incised, articulated truncation surface that formed when the Eo-Alpine orogen, including the area of the future Northern Calcareous Alps, was uplifted and subaerially eroded. Distinct facies associations were deposited from (1) alluvial fans and fan deltas, (2) rivers, (3) siliciclastic lagoonal to freshwater marsh environments, (4) areally/temporally limited carbonate lagoons, (5) transgressive shores, (6) siliciclastic shelf environments, and (7) an aggrading carbonate shelf. During the Turonian to Coniacian, the combination of high rates of both subsidence and sediment accumulation, and a narrow shelf that was compartmentalized with respect to (a) morphology of the substratum, (b) fluviatile input of siliciclastics and contemporaneous input of carbonate clasts from fan deltas, (c) deposition of shallow-water carbonates, and (d) water energy and-depth gave rise to an exceptionally wide spectrum of facies as a distinguishing feature of the succession. With the exception of facies association 7, which formed only once, depositional sequences in the Turonian to Coniacian interval contain all of the facies associations 1 to 6. During Turonian to Coniacian times, the shelf was microtidal to low-mesotidal, and was dominated by waves, storm waves and storm-induced currents. In vegetated marshes, schizohaline to freshwater marl lakes existed. Transgressions occurred onto fan deltas and in association with estuaries, or in association with gravelly to rocky shores. The transgressive successions, including successions deposited from transgressive rocky carbonate shores, are overlain by regressive successions of shelf carbonates or shelf siliciclastics. Deposition of shallow-water carbonates generally occurred within lagoons and over short intervals of time. A „catch-up” succession of shelf carbonates about 100 m thick accumulated only in an area protected from siliciclastic input. In its preserved parts, the Turonian to Coniacian succession does not record deposition adjacent to major active faults. Lateral changes in thickness result mainly from onlap onto the articulated basal truncation surface. Subsidence most probably was controlled by major detachment faults outside the outcrop area, and/or was distributed over a wide area in association with secondary faults above the major detachments. During Coniacian to Early Santonian times, both the older substratum and the overlying Turonian-Coniacian succession were subaerially exposed, faulted and deeply eroded. The following Early Santonian transgression ensued with rocky carbonate shores ahead of a sandy, narrow shoreface-inner shelf environment and a deeper shelf with intermittentlydysaerobic mud. The transgression was associated with the influx of cooler and/or nutrient-rich waters, and heralds an overall deepening. Still during the Early Santonian, the deepening was interrupted by another phase of subaerial exposure. Subsequently, a short phase of shelf deposition was terminated by deepening into bathyal depths.     

The ultrastructure of the midgut epithelium in millipedes (Myriapoda,Diplopoda)

The midgut epithelia of the millipedes Polyxenus lagurus, Archispirostreptus gigas and Julus scandinavius were analyzed under light and transmission electron microscopies. In order to detect the proliferation of regenerative cells, labeling with BrdU and antibodies against phosphohistone H3 were employed. A tube-shaped midgut of three millipedes examined spreads along the entire length of the middle region of the body. The epithelium is composed of digestive, secretory and regenerative cells. The digestive cells are responsible for the accumulation of metals and the reserve material as well as the synthesis of substances, which are then secreted into the midgut lumen. The secretions are of three types – merocrine, apocrine and microapocrine. The oval or pear-like shaped secretory cells do not come into contact with the midgut lumen and represent the closed type of secretory cells. They possess many electron-dense granules (J. scandinavius) or electron-dense granules and electron-lucent vesicles (A. gigas, P. lagurus), which are accompanied by cisterns of the rough endoplasmic reticulum. The regenerative cells are distributed individually among the basal regions of the digestive cells. The proliferation and differentiation of regenerative cells into the digestive cells occurred in J. scandinavius and A. gigas, while these processes were not observed in P. lagurus. As a result of the mitotic division of regenerative cells, one of the newly formed cells fulfills the role of a regenerative cell, while the second one differentiates into a digestive cell. We concluded that regenerative cells play the role of unipotent midgut stem cells.     

Ultrastructural investigation of the ovary structure ofOphyiulus pilosus (Myriapoda,Diplopoda)

Summary Histological and ultrastructural investigations of diplopod ovary structure have revealed that oogonia and early meiotic oocytes develop only in the laterodorsal parts of the ovarian wall, where they form groups called germ nests. Euplasmic growth forces diplotene oocytes out of the ovarian wall and into the lumen of the ovary, which leads to the formation of ovarian sacs. Ovarian sacs constitute separate structural-functional units built of a centrally situated oocyte and the epithelial cover. Being turned with their basal parts to the surface of the oocyte and showing no signs of any synthetic nor secretory activity, the epithelial cells of the ovarian sac wall cannot be referred to as typical follicular cells. That is why oogenesis of diplopods must be regarded as solitary.     

The allometry of metabolism in southern African millipedes (Myriapoda: Diplopoda)

Abstract. We determined standard metabolic rate at 25°C in forty-eight species of millipede from southern Africa and compared these data with confident measures of standard metabolic rate previously published for other arthropod groups.Metabolic rate in millipedes was not significantly different from that in beetles, ants or spiders once body mass effects had been accounted for, but was significantly higher than that in ticks.The exponent for the mass scaling of metabolic rate did not vary significantly between the five arthropod orders.Our best estimate for the relationship between standard metabolic rate (μl O₂ h^-1) and body mass (mg) in non-tick arthropods was 0.86 mass^0.73.     

Notes on the life-history and ecology of Tachypodoiulus niger (Diplopoda, Iulidae) in Britain

In some iulids females can moult several times after first attaining maturity but males cannot. Males of Tachypodoiulus niger (Leach) are exceptional in that they can moult, but they lose their functional intromittent organs in the process, regaining them after a second moult. The process can be repeated, a series of functional males alternating with nonfunctional males. This extension of the life and the consequent increase in number of segments led Verhoeff (1928) to postulate that the ancestral millipede was short-bodied and that many-segmented forms were derived from it. We think that this extension of life is of ecological rather than phylogenetic significance–adapting the species to disperse widely to scattered habitats. In particular, the extension of life of males as well as of females may ensure a reasonable sex ratio in those areas where the species is least dense.
In Britain, Tachypodoiulus niger lays eggs in spring which reach the fourth and fifth stadia by their first winter and the seventh, eighth and ninth stadia by their second winter. Maturity is usually attained by males in the eighth stadium but occasional specimens mature at stadium seven and others defer maturity until the ninth. Both sexes can proceed to the fourteenth stadium but adults in Britain usually belong to two generations of two and three years old, divided mainly between the eighth, ninth and tenth stadia. Details are given of a large collection made by the late Dr Scott of animals taking refuge in his house at Henley-on-Thames. Animals in Britain appear to have a similar life history to those in Germany as described by Verhoeff, but effective comparison has depended on correcting and re-interpreting some of Verhoeff's data.     

The accessory lateral eye of a diplopod, Cylindroiulus truncorum (Silvestri, 1896) (Diplopoda: Julidae)

Using electron microscopy we describe an accessory lateral eye for Cylindroiulus, a diplopod. The accessory eye is situated at the cell body rind of the optic lobes, deep inside the head, and is composed of six R-cells; a dioptric apparatus is absent. Comparison reveals that many arthropods possess accessory lateral eyes in addition to the compound eyes or lateral ocelli. Their homology and distribution among the arthropod main lineages is discussed along with characters that may be useful for reconstructing phylogeny.     

Occurrence and assemblage composition of millipedes (Myriapoda, Diplopoda) and terrestrial isopods (Crustacea, Isopoda, Oniscidea) in urban areas of Switzerland

Terrestrial isopods and millipedes, members of the invertebrate macro-decomposer guild, were collected through pitfall traps in three Swiss cities (Zurich, Lucerne, Lugano). A total of 7,198 individuals of 17 isopod species (7093 ind.), and 10 millipede species (105 ind.) were captured. Besides the Alpine endemic isopod (Trichoniscus alemannicus) and millipede (Cylindroiulus verhoeffi), urban assemblages were mainly composed of widespread, native European and even cosmopolitan species, which are frequent in anthropogenic areas. Overall species richness (isopods and millipedes combined) was similar in Zurich (17 species) and Lucerne (16), while only 13 species were sampled in Lugano. According to the Sørensen index of similarity, species composition of Zurich and Lucerne were more alike, while the one of Lugano was more distinct from the other two cities. This result can be explained by the spatial proximity of Zurich and Lucerne in the north of the Alps compared to Lugano, which is located more distantly and in the south of the Alps. Dominant isopods and millipedes in Zurich and Lucerne were found to be widespread synanthropic species in temperate Europe(Porcellio scaber, Trachelipus rathkii and Ophyiulus pilosus) while the dominant isopod in Lugano (Trachelipus razzautii) is a species with a north-eastern Mediterranean distribution. Our study reveals that the urban millipede and isopod fauna in Swiss cities mainly consists of widespread species, but species of narrower distribution (e.g. Trichoniscus alemannicus, Cylindroiulus verhoeffi) may also find suitable habitats in cities. Despite some signs of biotic homogenization, our study also found compositional differences of millipede and isopod assemblages between northern and southern cities that suggest geographical effects of the regional species pool.     

Gene expression suggests decoupled dorsal and ventral segmentation in the millipede Glomeris marginata (Myriapoda: Diplopoda)

Diplopods (millipedes) are known for their irregular body segmentation. Most importantly, the number of dorsal segmental cuticular plates (tergites) does not match the number of ventral structures (e.g., sternites). Controversial theories exist to explain the origin of this so-called diplosegmentation. We have studied the embryology of a representative diplopod, Glomeris marginata, and have analyzed the segmentation genes engrailed (en), hedgehog (hh), cubitus-interruptus (ci), and wingless (wg). We show that dorsal segments can be distinguished from ventral segments. They differ not only in number and developmental history, but also in gene expression patterns. engrailed, hedgehog, and cubitus-interruptus are expressed in both ventral and dorsal segments, but at different intrasegmental locations, whereas wingless is expressed only in the ventral segments, but not in the dorsal segments. Ventrally, the patterns are similar to what has been described from Drosophila and other arthropods, consistent with a conserved role of these genes in establishing parasegment boundaries. On the dorsal side, however, the gene expression patterns are different and inconsistent with a role in boundary formation between segments, but they suggest that these genes might function to establish the tergite borders. Our data suggest a profound and rather complete decoupling of dorsal and ventral segmentation leading to the dorsoventral discrepancies in the number of segmental elements. Based on gene expression, we propose a model that may resolve the hitherto controversial issue of the correlation between dorsal tergites and ventral leg pairs in basal diplopods (e.g., Glomeris) and is suggestive also for derived, ring-forming diplopods (e.g., Juliformia).     

Genesis of a soil seed bank on a primary succession in the Central Alps (Ötztal,Austria)

Soil seed bank and standing vegetation were investigated on the Rotmoos Glacier foreland (Ötztal, Tyrol, Austria) along the chronosequence (i.e. on the pioneer, early, and late successional stage) as well as on a subalpine pasture beyond the glacier foreland (old successional stage). We aimed to answer the following questions: (1) How large are soil seed banks along the successional gradient? (2) Do the seed banks reflect the actual standing vegetation or do they remember earlier successional stages or do they represent already the next successional stage?     

Anthracoporella mounds in the Late Carboniferous Auernig Group,Carnic Alps (Austria)

Summary A heretofore undocumented example of skeletal mounds formed by the dasycladacean algaAnthracoporella spectabilis is described from mixed carbonate-clastic cycles (Auernig cyclothems) of the Late Carboniferous (Gzhelian) Auernig Group of the central Carnic Alps in southern Austria. The massive mound facies forms biostromal reef mounds that are up to several m thick and extend laterally over more than 100 m. The mound facies is developed in the middle of bedded limestones, which are up to 16 m thick. These limestones formed during relative sea-level highstands when clastic influx was near zero. The mound facies is characterized by well developed baffler and binder guilds and does not show any horizontal or vertical zonation. Within the massive mound faciesAnthracoporella is frequently found in growth position forming bafflestones and wackestones composed of abundantAnthracoporella skeletons which toppled in situ or drifted slightly.Anthracoporella grew in such profusion that it dominated the available sea bottom living space, forming ‘algal meadows’ which acted as efficient sediment producers and bafflers. BecauseAnthracoporella could not provide a substantial reef framework, and could not withstand high water turbulence, the biostromal skeletal mounds accumulated in shallow, quiet water below the active wave base in water depths less than 30 m. The massive mound facies is under- and overlain by, and laterally grades into bedded, fossiliferous limestones of the intermound facies, composed mainly of different types of wackestones and packstones. Individual beds containAnthracoporella andArchaeolithophyllum missouriense in growth position, forming “micromounds’. Two stages of mound formation are recognized: (1) the stabilization stage when bioclastic wackestones accumulated, and (2) the skeletal mound stage when the sea-bottom was colonized byAnthracoporella and other members of the baffler and binder guilds, formingAnthracoporella bafflestones and wackestones of the mound facies. A slight drop in sea-level led to the termination of the mound growth and accumulation of organic debris, particularly calcareous algae, fusulinids, crinoids and bryozoans, forming well bedded limestones, which overlie the mound facies     

Palaeogeography and geodynamic evolution of the Gosau Group of the Northern Calcareous Alps (Late Cretaceous, Eastern Alps, Austria)

The sedimentation of the Late Cretaceous Gosau Group of the Northern Calcareous Alps records the complex geodynamic evolution of the northern part of the Austroalpine microplate. The tectonically controlled sedimentation and facies distribution were mainly governed by the oblique subduction of the Pennic-Ligurian ocean below the Austroalpine margin. Most of the Gosau successions comprise two distinct sedimentary complexes. The Lower Gosau Subgroup (upper Turonian-Campanian) is characterized by terrestrial to shallow-marine facies associations: alluvial fan and fan delta deposits, shallow-marine sandstones and sandy limestones, and storm-influenced nearshore and shelf deposits. During the early Santonian sea-level highstand, a great part of the Northern Calcareous Alps was covered by a shallow shelf sea. Palaeotransport directions point to source areas situated both to the north and to the south of the Northern Calcareous Alps.

The Upper Gosau Subgroup (upper Santonian-Eocene) comprises deep-water hemipelagic and turbiditic deposits. Rapid subsidence into bathyal to abyssal depths started diachronously from the northwest (Santonian) to the southeast (Maastrichtian) after a short phase of deformation and erosion. Abyssal deposits in the northern parts of the Northern Calcareus Alps and north-directed palaeotransport indicate a tilting of the slope towards the north. This subsidence pulse is attributed to an event of subcrustal tectonic erosion, probably triggered by the subduction of a topographic high of the South Penninic ocean. The relationships of the Gosau Group of the Austroalpine unit to Late Cretaceous deposits of the Carpathians and Southern Alps are discussed in the framework of a plate tectonic model.     

Millipedes faced with drought: the life cycle of a Mediterranean population of Ommatoiulus sabulosus (Linnaeus) (Diplopoda,Julida, Julidae)

Growth, development and life-cycle duration of the millipede Ommatoiulussabulosus (f.aimatopodus) were studied in a Mediterranean shrubland of southern France and compared with previous data from northwest Europe. Changes in the proportions of stadia during the course of the year were analysed in several generations. The results show that stadia VII and VIII are consistently reached after the first year of growth, and stadia IX and X after the second year. First reproduction may occur at the age of two years in males reaching maturity at stadium X, but not until the age of three in those reaching maturity at stadia XI and XII. Reproduction cannot occur until at least the age of three in females, which carry mature eggs from stadium XI onwards. In comparison with more northern populations, life-cycle duration is not shorter in the Mediterranean population but there are marked differences in its phenology: the breeding period is in autumn, so that juveniles of stadia II to VI are never faced with the summer drought, and larger individuals are mostly inactive in summer; moreover, all individuals moult once every winter. The results illustrate how julid millipedes of humid temperate regions could respond to higher temperatures and drier summer conditions in the context of climate change.     

Interactions of temperature and nutrient changes: effects on phytoplankton in the Piburger See (Tyrol,Austria)

1. Contemporary limnological and palaeolimnological data from Piburger See (Eastern Alps, Austria) allowed the reconstruction of its trophic state since the late 19th century and the assessment of changes in phytoplankton biomass and species composition in relation to selected environmental parameters. 2. A radiometrically dated sediment core from Piburger See was analysed for geochemical parameters, spheroidal carbonaceous particles (SCPs), bacterial and algal pigments, and diatoms. The low SCP sediment inventory assigns Piburger See to the ‘cleaner’ sites in Europe with respect to fossil‐fuel related air pollution. The sedimentary pigment and diatom record reveals moderate eutrophication during the 20th century, followed by a slow re‐oligotrophication since the mid‐1980s because of lake restoration starting in 1970. 3. Epilimnetic temperature for Piburger See was reconstructed using air temperature records. A pronounced temperature increase has been recorded during the mid‐1940s and since the late‐20th century, both promoting algal growth and changes in species composition (e.g. increase in centric diatoms and recent bloom of Asterionella formosa). 4. Climate scenarios project additional substantial warming for this mountain lake by the end of the 21st century which will be most pronounced during the growing season. The predicted change in lake water temperature and thermal dynamics represents a key driver for the trophic and ecological status of Piburger See in the future.     

Middle Triassic basin evolution and stratigraphy in the Carnic Alps (Austria)

Summary A local intraplatform basin developed in the Gartnerkofel-Zielkofel area of the Carnic Alps (southern Carinthia, Austria) during the Middle Triassic (Ladinian). This basin was filled with a transgressive basinal sequence composed of the Uggowitz Formation and overlying Buchenstein Formation. At the northwestern slope of the Gartnerkofel, the platform carbonates of the Schlern Dolomite interfinger with the Buchenstein Formation, causing the formation of two depositional sequences. The Uggowitz Formation consists of the Uggowitz Breccia and the Kühweg Member. Sediments of the Uggowitz Breccia were formed by different types of gravity induced processes. The Kühweg Member is a thin sequence of silt-and fine-grained sandstones which were deposited in a slope to basin margin environment by turbidity currents. The overlying Buchenstein Formation consists of hemipelagic to pelagic limestones of Fassanian age with intercalated pyroclastic rocks (Pietra verde). Nodular limestones were deposited under slow rates of accumulation during a relative sea-level highstand. The uppermost Buchenstein Formation is composed of hemipelagic limestone beds with intercalated graded calcarenites and breccias of platform-derived debris, showing characteristics features of a fore-reef slope of the prograding Schlern Dolomite. Uggowitz Formation and basal Buchenstein Formation are interpreted as a transgressive systems tract, nodular limestones from the middle part of the Buchenstein Formation mark an early highstand systems tract, forereef slope sediments of the upper Buchenstein Formation formed during the beginning regression of a late highstand systems tract, the basal part of the overlying Schlern Dolomite probably reflects a lowstand systems tract. The intercalated bedded limestone facies within the Schlern Dolomite is characterized by large, platform derived blocks, slump structures, breccia beds, graded calcarenites and hemipelagic limestones indicating a forereef slope environent. This intercalated facies belongs to the Buchenstein Formation and interfingers with the Schlern Dolomite. Conodonts from this intercalated slope facies point to Late Fassanian age. Therefore, the two Middle Triassic depositional sequences of the Gartnerkofel area can be correlated with the depositional sequences ‘Ladinian 1’ and ‘Ladinian 2’ of the Dolomites, proposed byDe Zanche et al. (1993). A brief comparison with the basinal sequences of similar age of the karawanken Mountains and the Carnia is presented.