Institut für Ökologie, Evolution und Diversität
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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 intends to provide a detailed 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 lakustrine (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, which are 2 to 10 times higher in fossil than in modern enamel; secondary enrichments in fossil dentin and cement are 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 very 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 variation.
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 granite 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 recrystallization of existing and addition of new apatite during maturation. Perhaps 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 composition, 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.
A Bayesian framework to estimate diversification rates and their variation through time and space
(2011)
Background: Patterns of species diversity are the result of speciation and extinction processes, and molecular phylogenetic data can provide valuable information to derive their variability through time and across clades. Bayesian Markov chain Monte Carlo methods offer a promising framework to incorporate phylogenetic uncertainty when estimating rates of diversification.
Results: We introduce a new approach to estimate diversification rates in a Bayesian framework over a distribution of trees under various constant and variable rate birth-death and pure-birth models, and test it on simulated phylogenies. Furthermore, speciation and extinction rates and their posterior credibility intervals can be estimated while accounting for non-random taxon sampling. The framework is particularly suitable for hypothesis testing using Bayes factors, as we demonstrate analyzing dated phylogenies of Chondrostoma (Cyprinids) and Lupinus (Fabaceae). In addition, we develop a model that extends the rate estimation to a meta-analysis framework in which different data sets are combined in a single analysis to detect general temporal and spatial trends in diversification.
Conclusions: Our approach provides a flexible framework for the estimation of diversification parameters and hypothesis testing while simultaneously accounting for uncertainties in the divergence times and incomplete taxon sampling.
Background: Taxonomy or biological systematics is the basic scientific discipline of biology, postulating hypotheses of identity and relationships, on which all other natural sciences dealing with organisms relies. However, the scientific contributions of taxonomists have been largely neglected when using species names in scientific publications by not citing the authority on which they are based.
Discussion: Consequences of this neglect is reduced recognition of the importance of taxonomy, which in turn results in diminished funding, lower interest from journals in publishing taxonomic research, and a reduced number of young scientists entering the field. This has lead to the so-called taxonomic impediment at a time when biodiversity studies are of critical importance.
Here we emphasize a practical and obvious solution to this dilemma. We propose that whenever a species name is used, the author(s) of the species hypothesis be included and the original literature source cited, including taxonomic revisions and identification literature - nothing more than what is done for every other hypothesis or assumption included in a scientific publication. In addition, we postulate that journals primarily publishing taxonomic studies should be indexed in ISISM.
Summary: The proposal outlined above would make visible the true contribution of taxonomists within the scientific community, and would provide a more accurate assessment for funding agencies impact and importance of taxonomy, and help in the recruitment of young scientists into the field, thus helping to alleviate the taxonomic impediment. In addition, it would also make much of the biological literature more robust by reducing or alleviating taxonomic uncertainty.
Keywords: Taxonomy crisis; taxonomic impediment; impact factor; original species description; citation index; systematics
Savannas are the most important timber and non-timber forest products (NTFPs) providing ecosystems in West Africa. They have been shaped by traditional human land-use (i.e. agriculture, grazing, and harvesting) for thousands of years. In the last decades, land-use has drastically changed due to the rapid population growth and the growing production of cash-crop in West Africa and this process is still continuing. The percentage of land intensively used for agriculture has increased, while the length of fallow periods has decreased. Such changes have enormous ecological, economic, and social consequences. In the context of land-use changes, there is an urgent need to better understand and evaluate the impact of land-use on savannas. Such an understanding provides insights on appropriate management activities that ensure the maintenance of savannas and guarantee the availability of savanna products for subsistence and commercial use of rural West African people.
The major objective of the present thesis was to study the impact of land-use on savanna vegetation and diversity as well as on populations of two important NTFP-providing tree species in a semi-arid area in West Africa. The study area was located in the south-eastern part of Burkina Faso and comprised the protected W National Park and its adjacent communal area.
In the first study (chapter 2), I investigated in cooperation with a colleague from Burkina Faso (Blandine Nacoulma) the impact of land-use on the savanna vegetation. We analyzed which environmental factors determine the occurrence of the vegetation types and investigated the effect of land-use on vegetation structure and the occurrence of life forms and highly valued tree species. Furthermore, we tested whether land-use has an impact on plant diversity pattern and if this impact differed between the vegetation types and layers (woody and herb layer). Vegetation relevés were performed and the vegetation and plant diversity of the protected W National Park were compared with those of its surrounding communal area. Our results reveal five vegetation types occurring in both areas. Elevation and physical soil characteristics and thus soil water availability for plants played the most important role for the occurrence of the vegetation types. The influence of land-use on plant diversity differed between the five vegetation types and the two layers. The impact was highest on the vegetation types with the most favorable soil conditions for cultivation and lowest on rocky habitats with poor soils. While the diversity of the woody layer was increased under human land-use, the diversity of the herb layer was diminished. Overall, as land-use effects were not only negative, our findings suggest that land-use does not automatically lead to a loss of plant species and to a degradation of savanna habitats. We conclude that both protected and communal areas are of great importance for the conservation of savanna vegetation and diversity. Our study highlights furthermore the importance of different management strategies for each vegetation type.
In the following two studies (chapter 3 and 4), the impact of land-use - and in particular of harvesting - on populations of Adansonia digitata L., the baobab tree, and Anogeissus leiocarpa (DC.) Guill. & Perr. was examined. These two tree species were chosen as they provide several NTFPs for the local population and as they show different levels of human protection and opposed life histories. Thus, they may react differently to land-use. Stands of the protected W National Park were compared with those of its surrounding communal area (in fallows, croplands, and villages). I applied dendrometric methods to study the population structures and combined it with rates and patterns of NTFP-harvesting (debarking and chopping/pruning). Furthermore, the impact of land-use and harvesting on the fruit production of A. digitata and on the sprouting ability of A. leiocarpa were studied. The inverse J-shaped size class distribution curve indicates that the stands of A. digitata were in a healthy state in the park, while the low number of smaller size classes in fallows, croplands, and villages may give evidence of an ageing population. However, a high number of seedlings were recorded in villages. The stands of A. leiocarpa were also in healthy states in the park and likewise in fallows. In contrast, the absence of saplings gives evidence of a declining population in croplands. Both species were strongly harvested by local people and harvesting was tree size-specific. Pruning in interaction with tree-size had a significant impact on fruit production of A. digitata. While smaller trees were more vulnerable to pruning, bigger trees benefited from slight-pruning. A. leiocarpa had a great ability to respond to chopping by sprouting. The sprouting ability increased even with higher chopping intensity. Results suggest that despite the intense harvesting and the land-use impact, populations of both species are still well preserved. While A. digitata can withstand the harvesting and land-use pressure by its longevity, extremely low adult mortality rates, and particularly due to positive human influences, A. leiocarpa is able to withstand the use pressure by its fast growing, high recruitment, and high sprouting ability. I conclude that a none protected tree species (A. leiocarpa) might not necessarily be at higher risk to the harvesting and land-use impact than a protected tree species (A. digitata) as the adverse impact of harvesting and land-use can be compensated by its specific life history.
Important additional information to such ecological findings can be provided by local people. Learning from traditional knowledge and management systems of local people will help to produce culturally and ecologically reasonable conservation and management strategies. Thus, I investigated local uses and management strategies of A. digitata and A. leiocarpa in the last two studies (chapter 5 and 6). Quantitative ethnobotanical surveys among the Gulimanceba people were conducted in the communal area in order to document uses of the different plant parts, harvesting modes, perceptions about the population status, and conservation status of both species. Hereby, differences in knowledge between gender, generations, and people from different villages were tested. Interviews reveal that both species are harvested for multipurpose and emphasize the high importance of both species for local people. Especially the leaves and fruits of A. digitata add valuable minerals and vitamins to the otherwise micronutrient-“poor” staple crops of the Gulimanceba people. In comparison with other studies in West Africa, it has turned out that people in this area could benefit even more from A. leiocarpa, e.g. for dyeing of clothes, for treatment of malaria and skin problems. Local knowledge did not differ between genders and generations, while it slightly differed between people from different villages. The lack of age differences suggests that the traditional knowledge about these two species is passed on from one generation to another. Differences between people from different villages might be explained by influences from the neighboring countries Niger and Benin. Current local harvesting modes and management strategies of both species resulted in sustainable use. However, ongoing land-use intensifications require adapted harvesting and management techniques to guarantee the persistence of these economically important species. These results provide, in combination with the ecological findings (chapter 3 and 4), appropriate management recommendations for A. digitata and A. leiocarpa that are reliable under currently practiced management strategies.
This article reports on the second Young Environmental Scientists Meeting that was hosted from 28 February to 2 March 2011 by the Institute for Environmental Research at RWTH Aachen University, Germany. This extraordinary meeting was again initiated and organized by the Student Advisory Council under the umbrella of Society of Environmental Toxicology and Chemistry Europe. A movie about the meeting and the abstracts of poster and platform presentations are freely available as supplemental material of this article.
The intensive foraging of insectivorous birds and bats is well known to reduce the density of arboreal herbivorous arthropods but quantification of collateral leaf damage remains limited for temperate forest canopies. We conducted exclusion experiments with nets in the crowns of young and mature oaks, Quercus robur, in south and central Germany to investigate the extent to which aerial vertebrates reduce herbivory through predation. We repeatedly estimated leaf damage throughout the vegetation period. Exclusion of birds and bats led to a distinct increase in arthropod herbivory, emphasizing the prominent role of vertebrate predators in controlling arthropods. Leaf damage (e.g., number of holes) differed strongly between sites and was 59% higher in south Germany, where species richness of vertebrate predators and relative oak density were lower compared with our other study site in central Germany. The effects of bird and bat exclusion on herbivory were 19% greater on young than on mature trees in south Germany. Our results support previous studies that have demonstrated clear effects of insectivorous vertebrates on leaf damage through the control of herbivorous arthropods. Moreover, our comparative approach on quantification of leaf damage highlights the importance of local attributes such as tree age, forest composition and species richness of vertebrate predators for control of arthropod herbivory.
Recovery of an ecosystem following disturbance can be severely hampered or even shift altogether when a point disturbance exceeds a certain spatial threshold. Such scale-dependent dynamics may be caused by preemptive competition, but may also result from diminished self-facilitation due to weakened ecosystem engineering. Moreover, disturbance can facilitate colonization by engineering species that alter abiotic conditions in ways that exacerbate stress on the original species. Consequently, establishment of such counteracting engineers might reduce the spatial threshold for the disturbance, by effectively slowing recovery and increasing the risk for ecosystem shifts to alternative states. We tested these predictions in an intertidal mudflat characterized by a two-state mosaic of hummocks (humps exposed during low tide) dominated by the sediment-stabilizing seagrass Zostera noltii) and hollows (low-tide waterlogged depressions dominated by the bioturbating lugworm Arenicola marina). In contrast to expectations, seagrass recolonized both natural and experimental clearings via lateral expansion and seemed unaffected by both clearing size and lugworm addition. Near the end of the growth season, however, an additional disturbance (most likely waterfowl grazing and/or strong hydrodynamics) selectively impacted recolonizing seagrass in the largest (1 m2) clearings (regardless of lugworm addition), and in those medium (0.25 m2) clearings where lugworms had been added nearly five months earlier. Further analyses showed that the risk for the disturbance increased with hollow size, with a threshold of 0.24 m2. Hollows of that size were caused by seagrass removal alone in the largest clearings, and by a weaker seagrass removal effect exacerbated by lugworm bioturbation in the medium clearings. Consequently, a sufficiently large disturbance increased the vulnerability of recolonizing seagrass to additional disturbance by weakening seagrass engineering effects (sediment stabilization). Meanwhile, the counteracting ecosystem engineering (lugworm bioturbation) reduced that threshold size. Therefore, scale-dependent interactions between habitat-mediated facilitation, competition and disturbance seem to maintain the spatial two-state mosaic in this ecosystem.
Background: Fragmented distribution ranges of species with little active dispersal capacity raise the question about their place of origin and the processes and timing of either range fragmentation or dispersal. The peculiar distribution of the land snail Tudorella sulcata s. str. in Southern France, Sardinia and Algeria is such a challenging case. Methodology: Statistical phylogeographic analyses with mitochondrial COI and nuclear hsp70 haplotypes were used to answer the questions of the species' origin, sequence and timing of dispersal. The origin of the species was on Sardinia. Starting from there, a first expansion to Algeria and then to France took place. Abiotic and zoochorous dispersal could be excluded by considering the species' life style, leaving only anthropogenic translocation as parsimonious explanation. The geographic expansion could be dated to approximately 8,000 years before present with a 95% confidence interval of 10,000 to 3,000 years before present. Conclusions: This period coincides with the Neolithic expansion in the Western Mediterranean, suggesting a role of these settlers as vectors. Our findings thus propose that non-domesticated animals and plants may give hints on the direction and timing of early human expansion routes.
The influence of dispersal limitation on species ranges remains controversial. Considering the dramatic impacts of the last glaciation in Europe, species might not have tracked climate changes through time and, as a consequence, their present-day ranges might be in disequilibrium with current climate. For 1016 European plant species, we assessed the relative importance of current climate and limited postglacial migration in determining species ranges using regression modelling and explanatory variables representing climate, and a novel species-specific hind-casting-based measure of accessibility to postglacial colonization. Climate was important for all species, while postglacial colonization also constrained the ranges of more than 50 per cent of the species. On average, climate explained five times more variation in species ranges than accessibility, but accessibility was the strongest determinant for one-sixth of the species. Accessibility was particularly important for species with limited long-distance dispersal ability, with southern glacial ranges, seed plants compared with ferns, and small-range species in southern Europe. In addition, accessibility explained one-third of the variation in species' disequilibrium with climate as measured by the realized/potential range size ratio computed with niche modelling. In conclusion, we show that although climate is the dominant broad-scale determinant of European plant species ranges, constrained dispersal plays an important supplementary role.