The Central Paratethys Sea-rise and demise of a Miocene European marine biodiversity hotspot.

The Miocene Climate Optimum (MCO, ~ 17-14 Ma) was a time of extraordinary marine biodiversity in the Circum-Mediterranean Region. This boom is best recorded in the deposits of the vanished Central Paratethys Sea, which covered large parts of central to southeastern Europe. This sea harbored an extraordinary tropical to subtropical biotic diversity. Here, we present a georeferenced dataset of 859 gastropod species and discuss geodynamics and climate as the main drivers to explain the changes in diversity. The tectonic reorganization around the Early/Middle Miocene boundary resulted in the formation of an archipelago-like landscape and favorable conditions of the MCO allowed the establishment of coral reefs. Both factors increased habitat heterogeneity, which boosted species richness. The subsequent cooling during the Middle Miocene Climate Transition (~ 14-13 Ma) caused a drastic decline in biodiversity of about 67%. Among the most severely hit groups were corallivorous gastropods, reflecting the loss of coral reefs. Deep-water faunas experienced a loss by 57% of the species due to changing patterns in circulation. The low sea level led to a biogeographic fragmentation reflected in higher turnover rates. The largest turnover occurred with the onset of the Sarmatian when bottom water dysoxia eradicated the deep-water fauna whilst surface waters-dwelling planktotrophic species underwent a crisis.

Automatic determination of 3D orientations of fossilized oyster shells from a densely packed Miocene shell bed.

Shell beds represent a useful source of information on various physical processes that cause the depositional condition. We present an automated method to calculate the 3D orientations of a large number of elongate and platy objects (fossilized oyster shells) on a sedimentary bedding plane, developed to support the interpretation of possible depositional patterns, imbrications, or impact of local faults. The study focusses on more than 1900 fossil oyster shells exposed in a densely packed Miocene shell bed. 3D data were acquired by terrestrial laser scanning on an area of 459 m2 with a resolution of 1 mm. Bivalve shells were manually defined as 3D-point clouds of a digital surface model and stored in an ArcGIS database. An individual shell coordinate system (ISCS) was virtually embedded into each shell and its orientation was determined relative to the coordinate system of the entire, tectonically tilted shell bed. Orientation is described by the rotation angles roll, pitch, and yaw in a Cartesian coordinate system. This method allows an efficient measurement and analysis of the orientation of thousands of specimens and is a major advantage compared to the traditional 2D approach, which measures only the azimuth (yaw) angles. The resulting data can variously be utilized for taphonomic analyses and the reconstruction of prevailing hydrodynamic regimes and depositional environments. For the first time, the influence of possible post-sedimentary vertical displacements can be quantified with high accuracy. Here, the effect of nearby fault lines-present in the reef-was tested on strongly tilted oyster shells, but it was found out that the fault lines did not have a statistically significant effect on the large tilt angles. Aside from the high reproducibility, a further advantage of the method is its non-destructive nature, which is especially suitable for geoparks and protected sites such as the studied shell bed.

Predictors of shell size in long-lived lake gastropods.

AIM: To investigate shell size variation among gastropod faunas of fossil and recent long-lived European lakes and discuss potential underlying processes. LOCATION: Twenty-three long-lived lakes of the Miocene to Recent of Europe. METHODS: Based on a dataset of 1412 species of both fossil and extant lacustrine gastropods, we assessed differences in shell size in terms of characteristics of the faunas (species richness, degree of endemism, differences in family composition) and the lakes (surface area, latitude and longitude of lake centroid, distance to closest neighbouring lake) using multiple and linear regression models. Because of a strong species-area relationship, we used resampling to determine whether any observed correlation is driven by that relationship. RESULTS: The regression models indicated size range expansion rather than unidirectional increase or decrease as the dominant pattern of size evolution. The multiple regression models for size range and maximum and minimum size were statistically significant, while the model with mean size was not. Individual contributions and linear regressions indicated species richness and lake surface area as best predictors for size changes. Resampling analysis revealed no significant effects of species richness on the observed patterns. The correlations are comparable across families of different size classes, suggesting a general pattern. MAIN CONCLUSIONS: Among the chosen variables, species richness and lake surface area are the most robust predictors of shell size in long-lived lake gastropods. Although the most outstanding and attractive examples for size evolution in lacustrine gastropods come from lakes with extensive durations, shell size appears to be independent of the duration of the lake as well as longevity of a species. The analogue of long-lived lakes as 'evolutionary islands' does not hold for developments of shell size because different sets of parameters predict size changes.

Euryhaline preferences of the decapod crab Mioplax socialis enabled it to survive during the Badenian/Sarmatian extinction (Miocene) in the Central Paratethys.

Although decapod crustaceans of the Central Paratethys were diverse during the Badenian (Langhian-Early Serravallian), a dramatic drop in their diversity occurred at the boundary with the Sarmatian. A crab Mioplax socialis is one of the few decapods reported from the Lower Sarmatian (Mohrensternia Zone) of the Paratethys. Until now, this species has been known from only a handful of specimens from Austria, Croatia and Bulgaria (Central Paratethys), and its systematics and ecology remain poorly known. Here, on the basis of new specimens from the Sarmatian tuffitic clays of the Stretava Formation (Skáros, Eastern Slovakia) we confirm that this species belongs to the subfamily Chasmocarcininae. The diagnostic characters of the male sternum that allow this classification are reported for the first time. The molluscan assemblage co-occurring with M. socialis demonstrate that this species tolerated conditions with variable salinity. Its tolerance of a broad range of salinity regimes may thus explain its survival across the Badenian-Sarmatian extinction event. Preservation of near-complete and fully articulated individuals of M. socialis suggests calm conditions and short residence times on the sediment-water interface.

Tectonics, climate, and the rise and demise of continental aquatic species richness hotspots.

Continental aquatic species richness hotspots are unevenly distributed across the planet. In present-day Europe, only two centers of biodiversity exist (Lake Ohrid on the Balkans and the Caspian Sea). During the Neogene, a wide variety of hotspots developed in a series of long-lived lakes. The mechanisms underlying the presence of richness hotspots in different geological periods have not been properly examined thus far. Based on Miocene to Recent gastropod distributions, we show that the existence and evolution of such hotspots in inland-water systems are tightly linked to the geodynamic history of the European continent. Both past and present hotspots are related to the formation and persistence of long-lived lake systems in geological basins or to isolation of existing inland basins and embayments from the marine realm. The faunal evolution within hotspots highly depends on warm climates and surface area. During the Quaternary icehouse climate and extensive glaciations, limnic biodiversity sustained a severe decline across the continent and most former hotspots disappeared. The Recent gastropod distribution is mainly a geologically young pattern formed after the Last Glacial Maximum (19 ky) and subsequent formation of postglacial lakes. The major hotspots today are related to long-lived lakes in preglacially formed, permanently subsiding geological basins.

Synopsis of valid species-group taxa for freshwater Gastropoda recorded from the European Neogene.

Here we present a complete list of all valid species-group taxa of freshwater gastropods reported from Miocene and Pliocene deposits in Europe. The last comparable work dates back to the 1920s and covered about 1,600 names. The extensive literature research underlying the present work revealed considerable changes in the taxonomic and systematic frameworks of Neogene freshwater gastropods and yielded a total number of 2,156 accepted taxa. Each taxon is accompanied by a full citation of its first description; where the information is available, page number and illustration reference are provided. First descriptions available as open-access full-text sources on the web were linked via hyperlink to the first page of the publication.

Replacement names and nomenclatural comments for problematic species-group names in Europe's Neogene freshwater Gastropoda. Part 2.

In the course of a new database project on Miocene to Recent freshwater gastropods of Europe, a great many of primary and secondary homonyms were revealed. Such nomenclatural issues need clarification in order to avoid misunderstandings and wrong statements about geographical distributions and temporal ranges. The following 16 new names are introduced to replace existing homonyms: Theodoxus militaris jurisicpolsakae nom. n., Viviparus stevanovici nom. n., Melanopsis haueri ripanjensis nom. n., Melanopsis wolfgangfischeri nom. n., Micromelania ramacanensis nom. n., Pseudamnicola welterschultesi nom. n., Muellerpalia haszprunari nom. n., Muellerpalia pseudovalvatoides nom. n., Lithoglyphus gozhiki nom. n., Valvata heidemariae willmanni nom. n., Radix macaleti nom. n., Gyraulus okrugljakensis nom. n., Gyraulus rasseri nom. n., Gyraulus vrapceanus nom. n., Planorbarius halavatsi nom. n., and Segmentina mosbachensis nom. n. Additionally, six cases of homonyms are discussed that are not replaced by new names, because they are considered junior synonyms.

Replacement names and nomenclatural comments for problematic species-group names in Europe's Neogene freshwater Gastropoda.

Over the last 250 years of taxonomic descriptions of freshwater gastropods a large number of primary and secondary homonyms were produced. Several of them have now been uncovered in the course of a new database project. To overcome the associated nomenclatural problems we propose 10 replacement names: Theodoxus pseudodacicus nom. nov., Theodoxus stoicai nom. nov., Viviparus deleeuwi nom. nov., Viviparus lubenescuae nom. nov., Viviparus wesselinghi nom. nov., Melanopsis anistratenkoi nom. nov., Melanopsis gearyae nom. nov., Melanopsis magyari nom. nov., Melanopsis vrcinensis nom. nov., and Pyrgula rusti nom. nov. Additionally, we discuss taxa that might become secondary homonyms because of uncertain genus attributions. The genera Melanoptychia Neumayr, 1880 and Boistelia Cossmann, 1909 are synonymized with Melanopsis Férussac, 1807 in Férussac & Férussac, 1807 based on the lack of sufficient separation criteria. Involved combinations are expounded and recombined accordingly. The nomenclatural problems regarding Melanopsis costata Fuchs, 1870 (non Olivier, 1804) and Planorbis varians Fuchs, 1870 sensu Bandel (2010) are discussed.

High-resolution analysis of upper Miocene lake deposits: Evidence for the influence of Gleissberg-band solar forcing.

A high-resolution multi-proxy analysis was conducted on a 1.5-m-long core of Tortonian age (~ 10.5 Ma; Late Miocene) from Austria (Europe). The lake sediments were studied with a 1-cm resolution to detect all small-scale variations based on palynomorphs (pollen and dinoflagellate cysts), ostracod abundance, geochemistry (carbon and sulfur) and geophysics (magnetic susceptibility and natural gamma radiation). Based on an already established age model for a longer interval of the same core, this sequence can be limited to approx. two millennia of Late Miocene time with a resolution of ~ 13.7 years per sample. The previous study documented the presence of solar forcing, which was verified within various proxies on this 1.5-m core by a combination of REDFIT spectra and Gaussian filters. Significant repetitive signals ranged in two discrete intervals corresponding roughly to 55-82 and 110-123 years, fitting well within the lower and upper Gleissberg cycle ranges.Based on these results, the environmental changes along the 2000-year Late Miocene sequence are discussed. No major ecological turnovers are expected in this very short interval. Nonetheless, even within this brief time span, dinoflagellates document rapid changes between oligotrophic and eutrophic conditions, which are frequently coupled with lake stratification and dysoxic bottom waters. These phases prevented ostracods and molluscs from settling and promoted the activity of sulfur bacteria. The pollen record indicates rather stable wetland vegetation with a forested hinterland. Shifts in the pollen spectra can be mainly attributed to variations in transport mechanisms. These are represented by a few phases of fluvial input but mainly by changes in wind intensity and probably also wind direction. Such influence is most likely caused by solar cycles, leading to a change in source area for the input into the lake.Furthermore, these solar-induced variations seem to be modulated by longer solar cycles. The filtered data display comparable patterns and modulations, which seem to be forced by the 1000-year and 1500-year cycles. The 1000-year cycle modulated especially the lake surface proxies, whereas the 1500-year cycle is mainly reflected in hinterland proxies, indicating strong influence on transport mechanisms.

Paleomagnetic and geochronologic constraints on the geodynamic evolution of the Central Dinarides.

The geodynamic evolution of the Dinaride Mountains of southeastern Europe is relatively poorly understood, especially in comparison with the neighboring Alps and Carpathians. Here, we construct a new chronostratigraphy for the post-orogenic intra-montane basins of the Central Dinarides based on paleomagnetic and 40Ar/39Ar age data. A first phase of basin formation occurred in the late Oligocene. A second phase of basin formation took place between 18 and 13 Ma, concurrent with profound extension in the neighboring Pannonian Basin. Our paleomagnetic results further indicate that the Dinarides have not experienced any significant tectonic rotation since the late Oligocene. This implies that the Dinarides were decoupled from the adjacent Adria and the Tisza-Dacia Mega-Units that both underwent major rotation during the Miocene. The Dinaride orogen must consequently have accommodated significant shortening. This is corroborated by our AMS data that indicate post-Middle Miocene shortening in the frontal zone, wrenching in the central part of the orogen, and compression in the hinterland. A review of paleomagnetic data from the Adria plate, which plays a major role in the evolution of the Dinarides as well as the Alps, constrains rotation since the Early Cretaceous to 48 ± 10° counterclockwise and indicates 20° of this rotation took place since the Miocene. It also shows that Adria behaved as an independent plate from the Late Jurassic to the Eocene. From the Eocene onwards, coupling between Adria and Africa was stronger than between Adria and Europe. Adria continued to behave as an independent plate. The amount of rotation within the Adria-Dinarides collision zone increases with age and proximity of the sampled sediments to undeformed Adria. These results significantly improve our insight in the post-orogenic evolution of the Dinarides and resolve an apparent controversy between structural geological and paleomagnetic rotation estimates for the Dinarides as well as Adria.

Millennial-scale vegetation dynamics in an estuary at the onset of the Miocene Climate Optimum.

Pollen analyses have been proven to possess the possibility to decipher rapid vegetational and climate shifts in Neogene sedimentary records. Herein, a c. 21-kyr-long transgression-regression cycle from the Lower Austrian locality Stetten is analysed in detail to evaluate climatic benchmarks for the early phase of the Middle Miocene Climate Optimum and to estimate the pace of environmental change.Based on the Coexistence Approach, a very clear signal of seasonality can be reconstructed. A warm and wet summer season with c. 204-236 mm precipitation during the wettest month was opposed by a rather dry winter season with precipitation of c. 9-24 mm during the driest month. The mean annual temperature ranged between 15.7 and 20.8 °C, with about 9.6-13.3 °C during the cold season and 24.7-27.9 °C during the warmest month. In contrast, today's climate of this area, with an annual temperature of 9.8 °C and 660 mm rainfall, is characterized by the winter season (mean temperature: -1.4 °C, mean precipitation: 39 mm) and a summer mean temperature of 19.9 °C (mean precipitation: 84 mm).Different modes of environmental shifts shaped the composition of the vegetation. Within few millennia, marshes and salt marshes with abundant Cyperaceae rapidly graded into Taxodiaceae swamps. This quick but gradual process was interrupted by swift marine ingressions which took place on a decadal to centennial scale. The transgression is accompanied by blooms of dinoflagellates and of the green alga Prasinophyta and an increase in Abies and Picea. Afterwards, the retreat of the sea and the progradation of estuarine and wetland settings were a gradual progress again.Despite a clear sedimentological cyclicity, which is related to the 21-kyr precessional forcing, the climate data show little variation. This missing pattern might be due to the buffering of the precessional-related climate signal by the subtropical vegetation. Another explanation could be the method-inherent broad range of climate-parameter estimates that could cover small scale climatic changes.

Palaeoenvironmental evolution of Lake Gacko (Southern Bosnia and Herzegovina): Impact of the Middle Miocene Climatic Optimum on the Dinaride Lake System.

In the Early to Middle Miocene, a series of lakes, collectively termed the Dinaride Lake System (DLS), spread out across the north-western part of the Dinaride-Anatolian continental block. Its deposits, preserved in numerous intra-montane basins, allow a glimpse into the palaeoenvironmental, palaeobiogeographic and geodynamic evolution of the region. Lake Gacko, situated in southern Bosnia and Herzegovina, is one of the constituent lakes of the DLS, and its deposits are excellently exposed in the Gracanica open-cast coal-mine. A detailed study of the sedimentary succession that addresses facies, sediment petrography, geophysical properties, and fossil mollusc palaeoecology reveals repetitive changes in lake level. These are interpreted to reflect changes in the regional water budget. First-order chronologic constraints arise from the integration of radio-isotopic and palaeomagnetic data. (40)Ar/(39)Ar measurements on feldspar crystals from a tephra bed in the upper part of the sedimentary succession indicate a 15.31 ± 0.16 Ma age for this level. The reversed magnetic polarity signal that characterises the larger part of the investigated section correlates to chron C5Br of the Astronomically Tuned Neogene Timescale. Guided by these chronologic data and a detailed cyclostratigraphic analysis, the observed variations in lake-level, evident as two ~ 40-m and seven ~ 10-m scale transgression-regression cycles, are tuned to ~ 400-kyr and ~ 100-kyr eccentricity cycles. From the tuning, it can be inferred that the sediments in the Gacko Basin accumulated between ~ 15.8 and ~ 15.2 Ma. The economically valuable lignite accumulations in the lower part of the succession are interpreted to indicate the development of swamp forests in conjunction with lake-level falls corresponding to ~ 100-kyr eccentricity minima. Pedogenesis, rhizoliths and palustrine carbonate breccias in the upper part of the section reveal long-term aridity coinciding with a ~ 400-kyr eccentricity minimum. Eccentricity maxima are interpreted to trigger lake-level high-stands. These are accompanied by eutrophication events caused by enhanced denudation of the surrounding basement and increased detrital input into the basin. The presented age model proves that Lake Gacko arose during the Middle Miocene Climatic Optimum and that the optimum climatic conditions triggered the formation of this long-lived lake.