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Bioapatite in mammalian teeth is readily preserved in continental sediments and represents a very important archive for reconstructions of environment and climate evolution. This project provides a comprehensive data base of major, minor and trace element and isotope tracers for tooth apatite using a variety of microanalytical techniques. The aim is to identify specific sedimentary environments and to improve our understanding on the interaction between internal metabolic processes during tooth formation and external nutritional control and secondary alteration effects. Here, we use the electron microprobe to determine the major and minor element contents of fossil and modern molar enamel, cement and dentin from Hippopotamids. Most of the studied specimens are from different ecosystems in Eastern Africa, representing modern and fossil lacustrine (Lake Kikorongo, Lake Albert, and Lake Malawi) and modern fluvial environments of the Nile River system. Secondary alteration effects - in particular FeO, MnO, SO3 and F concentrations – are 2 to 10 times higher in fossil than in modern enamel; the secondary enrichment of these components in fossil dentin and cement is even higher. In modern and fossil enamel, along sections perpendicular to the enamel-dentin junction (EDJ) or along cervix-apex profiles, P2O5 and CaO contents and the CaO/P2O5 ratios are very constant (StdDev ∼1%). Linear regression analysis reveals tight control of the MgO (R2∼0.6), Na2O and Cl variation (for both R2>0.84) along EDJ-outer enamel rim profiles, despite large concentration variations (40% to 300%) across the enamel. These minor elements show well defined distribution patterns in enamel, similar in all specimens regardless of their age and origin, as the concentration of MgO and Na2O decrease from the enamel-dentin junction (EDJ) towards the outer rim, whereas Cl displays the opposite trend. Fossil enamel from Hippopotamids which lived in the saline Lake Kikorongo have a much higher MgO/Na2O ratio (∼1.11) than those from the Neogene fossils of Lake Albert (MgO/Na2O∼0.4), which was a large fresh water lake like those in the western Branch of the East African Rift System today. Similarly, the MgO/Na2O ratio in modern enamel from the White Nile River (∼0.36), which has a Precambrian catchment of dominantly granites and gneisses and passes through several saline zones, is higher than that from the Blue Nile River, whose catchment is the Neogene volcanic Ethiopian Highland (MgO/Na2O∼0.22). Thus, particularly MgO/Na2O might be a sensitive fingerprint for environments where river and lake water have suffered strong evaporation. Enamel formation in mammals takes place at successive mineralization fronts within a confined chamber where ion and molecule transport is controlled by the surrounding enamel organ. During the secretion and maturation phases the epithelium generates different fluid composition, which in principle, should determine the final composition of enamel apatite. This is supported by co-linear relationships between MgO, Cl and Na2O which can be interpreted as binary mixing lines. However, if maturation starts after secretion is completed, the observed element distribution can only be explained by equilibration of existing and addition of new apatite during maturation. It appears the initial enamel crystallites precipitating during secretion and the newly formed bioapatite crystals during maturation equilibrate with a continuously evolving fluid. During crystallization of bioapatite the enamel fluid becomes continuously depleted in MgO and Na2O, but enriched in Cl which results in the formation of MgO, and Na2O-rich, but Cl-poor bioapatite near the EDJ and MgO- and Na2O-poor, but Cl-rich bioapatite at the outer enamel rim. The linkage between lake and river water compositions, bioavailability of elements for plants, animal nutrition and tooth formation is complex and multifaceted. The quality and limits of the MgO/Na2O and other proxies have to be established with systematic investigations relating chemical distribution patterns to sedimentary environment and to growth structures developing as secretion and maturation proceed during tooth formation.
For reconstructing environmental change in terrestrial realms the geochemistry of fossil bioapatite in bones and teeth is among the most promising applications. This study demonstrates that alkaline earth elements in enamel of Hippopotamids, in particular Ba and Sr are tracers for water provenance and hydrochemistry. The studied specimens are molar teeth from Hippopotamids found in modern and fossil lacustrine settings of the Western Branch of the East African Rift system (Lake Kikorongo, Lake Albert, and Lake Malawi) and from modern fluvial environments of the Nile River.
Concentrations in enamel vary by ca. two orders of magnitude for Ba (120–9336 μg g−1) as well as for Sr (9–2150 μg g−1). Concentration variations in enamel are partly induced during post-mortem alteration and during amelogenesis, but the major contribution originates from the variable water chemistry in the habitats of the Hippopotamids which is dominated by the lithologies and weathering processes in the watershed areas. Amelogenesis causes a distinct distribution of Ba and Sr in modern and fossil enamel, in that element concentrations increase along profiles from the outer rim towards the enamel-dentin junction by a factor of 1.3–1.5. These elements are well correlated with MgO and Na2O in single specimens, thus suggesting that their distribution is determined by a common, single process. Presuming that the shape of the tooth is established at the end of the secretion process and apatite composition is in equilibrium with the enamel fluid, the maturation process can be modeled by closed system Rayleigh crystallization.
Enamel from many Hippopotamid specimens has Sr/Ca and Ba/Ca which are typical for herbivores, but the compositions extend well into the levels of plants and carnivores. Within enamel from single specimens these element ratios covary and provide a specific fingerprint of the Hippopotamid habitat. All specimens together, however, define subparallel trends with different Ba/Sr ranging from 0.1 to 3. This ratio varies on spatial and temporal scales and traces provenance signals as well as the fractionation of the elements in the hydrological cycle. Thus, Sr concentrations and Ba/Sr in enamel differentiate between habitats having basaltic or Archean crustal rocks as the ultimate sources of Sr and Ba. The provenance signal is modulated by climate change. In Miocene to Pleistocene enamel from the Lake Albert region, Ba/Sr decreases systematically with time from about 2 to 0.5. This trend can be correlated with changes in climate from humid to arid in vegetation from C3 to C4 biomass as well as with increasing evaporation of the lake water. The most plausible explanation is that with time, Ba mobility decreased relative to that of Sr. This can arise if preferential adsorption of Ba to clay and Fe-oxide-hydroxide is related to increasing aridification. Additionally, weathering solutions and lake water can become increasingly alkaline and barite becomes stable. In this case, Ba will be preferentially deposited on the watershed of Lake Albert and rivers with low Ba/Sr will feed the habitats of the Hippopotamids.
This study demonstrates that alkaline earth elements in enamel of hippopotamids, in particular Ba and Sr, are tracers for water provenance and hydrochemistry in terrestrial settings. The studied specimens are permanent premolar and molar teeth found in modern and fossil lacustrine sediments of the Western Branch of the East African Rift system (Lake Kikorongo, Lake Albert, and Lake Malawi) and from modern fluvial environments of the Nile River.
Concentrations in enamel vary by two orders of magnitude for Ba (120–9336 μg g−1) as well as for Sr (9–2150 μg g−1). The variations are partially induced during post-mortem alteration and during amelogenesis, but the major contribution originates ultimately from the variable water chemistry in the habitats of the hippopotamids which is controlled by the lithologies and weathering processes in the watershed areas. Amelogenesis causes a distinct distribution of MgO, Ba and Sr in modern and fossil enamel, in that element concentrations increase along profiles from the outer rim towards the enamel–dentin junction by a factor of 1.3–1.9. These elements are well correlated in single specimens, thus suggesting that their distribution is determined by a common, single process, which can be described by closed system Rayleigh crystallization of bioapatite in vivo.
Enamel from most hippopotamid specimens has Sr/Ca and Ba/Ca which are typical for herbivores. However, Ba/Sr ranges from 0.1 to 3 and varies on spatial and temporal scales. Thus, Sr concentrations and Ba/Sr in enamel differentiate between habitats having basaltic mantle rocks or Archean crustal rocks as the ultimate sources of Sr and Ba. This provenance signal is modulated by climate change. In Miocene to Pleistocene enamel from the Lake Albert region, Ba/Sr decreases systematically with time from 2 to 0.5. This trend can be correlated with changes in climate from humid to arid, in vegetation from C3 to C4 biomass as well as with increasing evaporation of the lake water. The most plausible explanation is that Ba mobility decreased with increasing aridification due to preferential deposition with clay and Fe-oxide-hydroxide or barite on the watershed of Lake Albert.
Freshwater biodiversity has declined dramatically in Europe in recent decades. Because of massive habitat pollution and morphological degradation of water bodies, many once widespread species persist in small fractions of their original range. These range contractions are generally believed to be accompanied by loss of intraspecific genetic diversity, due to the reduction of effective population sizes and the extinction of regional genetic lineages. We aimed to assess the loss of genetic diversity and its significance for future potential reintroduction of the long-tailed mayfly Palingenia longicauda (Olivier), which experienced approximately 98% range loss during the past century. Analysis of 936 bp of mitochondrial DNA of 245 extant specimens across the current range revealed a surprisingly large number of haplotypes (87), and a high level of haplotype diversity (Hd = 0.875). In contrast, historic specimens (6) from the lost range (Rhine catchment) were not differentiated from the extant Rába population (F ST = 0.02, p = 0.61), despite considerable geographic distance separating the two rivers. These observations can be explained by an overlap of the current with the historic (Pleistocene) refugia of the species. Most likely, the massive recent range loss mainly affected the range which was occupied by rapid post-glacial dispersal. We conclude that massive range losses do not necessarily coincide with genetic impoverishment and that a species' history must be considered when estimating loss of genetic diversity. The assessment of spatial genetic structures and prior phylogeographic information seems essential to conserve once widespread species.
Background: Mammalian fossils from the Eppelsheim Formation (Dinotheriensande) have been a benchmark for Neogene vertebrate palaeontology since 200 years. Worldwide famous sites like Eppelsheim serve as key localities for biochronologic, palaeobiologic, environmental, and mammal community studies. So far the formation is considered to be of early Late Miocene age (~9.5 Ma, Vallesian), representing the oldest sediments of the Rhine River. The stratigraphic unity of the formation and of its fossil content was disputed at times, but persists unresolved.
Principal Findings: Here we investigate a new fossil sample from Sprendlingen, composed by over 300 mammalian specimens and silicified wood. The mammals comprise entirely Middle Miocene species, like cervids Dicrocerus elegans, Paradicrocerus elegantulus, and deinotheres Deinotherium bavaricum and D. levius. A stratigraphic evaluation of Miocene Central European deer and deinothere species proof the stratigraphic inhomogenity of the sample, and suggest late Middle Miocene (~12.5 Ma) reworking of early Middle Miocene (~15 Ma) sediments. This results agree with taxonomic and palaeoclimatic analysis of plant fossils from above and within the mammalian assemblage. Based on the new fossil sample and published data three biochronologic levels within the Dinotheriensand fauna can be differentiated, corresponding to early Middle Miocene (late Orleanian to early Astaracian), late Middle Miocene (late Astaracian), and early Late Miocene (Vallesian) ages.
Conclusions/Significance: This study documents complex faunal mixing of classical Dinotheriensand fauna, covering at least six million years, during a time of low subsidence in the Mainz Basin and shifts back the origination of the Rhine River by some five million years. Our results have severe implications for biostratigraphy and palaeobiology of the Middle to Late Miocene. They suggest that turnover events may be obliterated and challenge the proposed ‘supersaturated’ biodiversity, caused by Middle Miocene superstites, of Vallesian ecosystems in Central Europe.
Sponges play a key role in Antarctic marine benthic community structure and dynamics and are often a dominant component of many Southern Ocean benthic communities. Understanding the drivers of sponge distribution in Antarctica enables us to understand many of general benthic biodiversity patterns in the region. The sponges of the Antarctic and neighbouring oceanographic regions were assessed for species richness and biogeographic patterns using over 8,800 distribution records. Species-rich regions include the Antarctic Peninsula, South Shetland Islands, South Georgia, Eastern Weddell Sea, Kerguelen Plateau, Falkland Islands and north New Zealand. Sampling intensity varied greatly within the study area, with sampling hotspots found at the Antarctic Peninsula, South Georgia, north New Zealand and Tierra del Fuego, with limited sampling in the Bellingshausen and Amundsen seas in the Southern Ocean. In contrast to previous studies we found that eurybathy and circumpolar distributions are important but not dominant characteristics in Antarctic sponges. Overall Antarctic sponge species endemism is ~43%, with a higher level for the class Hexactinellida (68%). Endemism levels are lower than previous estimates, but still indicate the importance of the Polar Front in isolating the Southern Ocean fauna. Nineteen distinct sponge distribution patterns were found, ranging from regional endemics to cosmopolitan species. A single, distinct Antarctic demosponge fauna is found to encompass all areas within the Polar Front, and the sub-Antarctic regions of the Kerguelen Plateau and Macquarie Island. Biogeographical analyses indicate stronger faunal links between Antarctica and South America, with little evidence of links between Antarctica and South Africa, Southern Australia or New Zealand. We conclude that the biogeographic and species distribution patterns observed are largely driven by the Antarctic Circumpolar Current and the timing of past continent connectivity.
Non-indigenous species that become invasive are one of the main drivers of biodiversity loss worldwide. In various freshwater systems in Europe, populations of native amphipods and fish are progressively displaced by highly adaptive non-indigenous species that can perform explosive range extensions. A total of 40 Ponto-Caspian round gobies Neogobius melanostomus from the Rhine River near Düsseldorf, North Rhine-Westphalia, Germany, were examined for metazoan parasites and feeding ecology. Three metazoan parasite species were found: two Nematoda and one Acanthocephala. The two Nematoda, Raphidascaris acus and Paracuaria adunca, had a low prevalence of 2.5%. The Acanthocephala, Pomphorhynchus tereticollis, was the predominant parasite species, reaching a level of 90.0% prevalence in the larval stage, correlated with fish size. In addition, four invasive amphipod species, Corophium curvispinum (435 specimens), Dikerogammarus villosus (5,454), Echinogammarus trichiatus (2,695) and Orchestia cavimana (1,448) were trapped at the sampling site. Only D. villosus was infected with P. tereticollis at a prevalence of 0.04%. The invasive goby N. melanostomus mainly preys on these non-indigenous amphipods, and may have replaced native amphipods in the transmission of P. tereticollis into the vertebrate paratenic host. This study gives insight into a potential parasite-host system that consists mainly of invasive species, such as the Ponto-Caspian fish and amphipods in the Rhine. We discuss prospective distribution and migration pathways of non-indigenous vertebrate (round goby) and invertebrates (amphipods) under special consideration of parasite dispersal.
We study how species richness of arthropods relates to theories concerning net primary productivity, ambient energy, water-energy dynamics and spatial environmental heterogeneity. We use two datasets of arthropod richness with similar spatial extents (Scandinavia to Mediterranean), but contrasting spatial grain (local habitat and country). Samples of ground-dwelling spiders, beetles, bugs and ants were collected from 32 paired habitats at 16 locations across Europe. Species richness of these taxonomic groups was also determined for 25 European countries based on the Fauna Europaea database. We tested effects of net primary productivity (NPP), annual mean temperature (T), annual rainfall (R) and potential evapotranspiration of the coldest month (PETmin) on species richness and turnover. Spatial environmental heterogeneity within countries was considered by including the ranges of NPP, T, R and PETmin. At the local habitat grain, relationships between species richness and environmental variables differed strongly between taxa and trophic groups. However, species turnover across locations was strongly correlated with differences in T. At the country grain, species richness was significantly correlated with environmental variables from all four theories. In particular, species richness within countries increased strongly with spatial heterogeneity in T. The importance of spatial heterogeneity in T for both species turnover across locations and for species richness within countries suggests that the temperature niche is an important determinant of arthropod diversity. We suggest that, unless climatic heterogeneity is constant across sampling units, coarse-grained studies should always account for environmental heterogeneity as a predictor of arthropod species richness, just as studies with variable area of sampling units routinely consider area.
The aim of this study was to assess the invasion risk of freshwater habitats and determine the environmental variables that are most favorable for the establishment of alien amphipods, isopods, gastropods, and bivalves. A total of 981 sites located in streams and rivers in Germany. Therefore we analyzed presence-absence data of alien and indigenous amphipods, isopods, gastropods, and bivalves from 981 sites located in small to large rivers in Germany with regard to eight environmental variables: chloride, ammonium, nitrate, oxygen, orthophosphate, distance to the next navigable waterway, and maximum and minimum temperature. Degraded sites close to navigable waters were exposed to an increased invasion risk by all major groups of alien species. Moreover, invaded sites by all four groups of alien species were similar, whereas the sites where indigenous members of the four groups occurred were more variable. Increased temperature and chloride concentration as well as decreased oxygen concentration were identified as major factors for the invasibility of a site. Species-specific analyses showed that chloride was among the three most predictive environmental variables determining species assemblage in all four taxonomic groups. Also distance to the next navigable waterways was similarly important. Additionally, the minimum temperature was among the most important variables for amphipods, isopods, and bivalves. The bias in the occurrence patterns of alien species toward similarly degraded habitats suggests that the members of all four major groups of freshwater alien species are a non-random, more tolerant set of species. Their common tolerance to salinity, high temperature, and oxygen depletion may reflect that most alien species were spread in ballast water tanks, where strong selective pressures, particularly temperature fluctuations, oxygen depletion, and increased salinity may create a bottleneck for successful invasion. Knowledge on the major factors that influence the invasion risk of a habitat is needed to develop strategies to limit the spread of invasive species.
We describe the frog species Diasporus citrinobapheussp. n. from the Cordillera Central of western Panama. The new species differs from all other species in its genus in coloration, disk cover and disk pad shape, skin texture, advertisement call, and size. It is most similar to Diasporus tigrillo, from which it differs in dorsal skin texture, relative tibia length, number of vomerine teeth, ventral coloration, dorsal markings, and relative tympanum size, and to Diasporus gularis, from which it can be distinguished by the lack of membranes between the toes, adult size, posterior thigh coloration, and position of the choanae. We provide data on morpho- logy, vocalization, and distribution of the new species, as well as brief information on its natural history.