Institut für Integrierte Naturwissenschaften, Abt. Biologie
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Agricultural intensification is leading to a severe decline in farmland biodiversity worldwide. The resulting landscape simplification through the expansion of monocultures and removal of non-crop habitats has a major impact on arthropod communities in agricultural landscapes. While arable fields are often highly disturbed and ephemeral habitats that are unsuitable for many species, non-crop habitats in agroecosystems can provide important refugia. The creation of non-crop habitats through agri-environmental schemes (AES) in intensive agricultural landscapes, such as the ‘Maifeld’ region in western Germany, is intended to mitigate the negative effects of agricultural intensification, although the effectiveness of these measures for nature conservation is still controversial. Therefore, this work focuses on the taxonomic and functional diversity of beetles (Coleoptera) and spiders (Araneida), being important providers of ecosystem services, between wheat fields and different non-crop habitats, namely grassy field margins adjacent to wheat and oilseed rape fields, small- and large-scale set-aside areas sown with wildflowers, and permanent grassland fallows. Arthropods were collected between 2019 and 2020 using pitfall traps and suction sampling. Land-use type influenced beetle and spider diversity in the study area, with significantly higher values in grassland fallows than wheat fields. Surprisingly, species diversity differed little among all non-crop habitats, but all harboured distinct species assemblages. In particular, large long-term grassland fallows showed the largest within-group variation of beetle and spider assemblages and represented important habitats, especially for habitat specialists and threatened species, likely due to their variable soil moisture and complex habitat structure. In contrast, the homogeneous arthropod assemblages of wheat fields exhibited lower trait richness and were dominated by a few predatory species adapted to such disturbed, man-made habitats. Interestingly, all conservation measures complemented each other in that they contributed in different ways to supporting beetles and spiders in agricultural landscapes. Even small-scale non-crop habitats and existing habitat boundaries in an agricultural matrix appear to be valuable habitats for farmland arthropods by enhancing taxonomic diversity. Field margins and small wildflower-sown patches can link isolated non-crop habitats and contribute to a heterogeneous agricultural landscape. Consequently, a combination of various small- and large-scale greening measures leads to increased compositional and configurational landscape heterogeneity, resulting in improved beetle and spider diversity. Considering the ongoing loss of farmland biodiversity worldwide, agri-environmental schemes should be promoted in the future, as they are particularly important for arthropod conservation in intensive agricultural landscapes such as the Maifeld region.
In this thesis, I present the results of my studies on taxonomy, systematics, and biogeography of Impatiens (Balsaminaceae) in Madagascar and the Comoro islands.
In Chapter 1 I reviewed the literature on taxonomy and classification of Balsaminaceae, on habitat, world distribution, morphology, molecular phylogenetics and infrageneric classification of the genus Impatiens. In Chapters 2-15 (Fischer & Rahelivololona 2002, 2003, 2004, 2007, 2015a, 2015b, 2015c, 2016, Fischer et al. 2004. 2017, 2018a, b submitted, Rahelivololona et al. 2003) I presented the first results of a revision of Balsaminaceae of Madagascar and the Comoro islands including the description of 78 new species. In Chapter 16 (Yuan et al. 2004) we worked on the phylogeny and biogeography of Balsaminaceae inferred from ITS sequences using combined results from molecular phylogenetic and morphological analyses. In Chapter 17 (Rahelivololona et al. 2018) we conducted a phylogeny and assessment of the infrageneric classification of species in the Malagasy Impatiens (Balsaminaceae) with a particular emphasis on taxa collected from Marojejy.
Below I summarise the most important findings of each chapter and provide an outlook for future studies.
How many species of Impatiens occur in Madagascar and the Comoro islands?
To provide additional information on the taxonomic revision of Impatiens in Madagascar and the Comoro islands, the identification of already described species as well as the description of new species was conducted. Based on herbarium specimens from BR, G, K, NEU, P, TAN and on living plants collected during several field trips, 78 new species and 6 nomina nova have been published and another 70 new taxa are already identified. Actually more than 260 species occur in Madagascar and the Comoro islands and all of them are endemic. For each species, a description of the morphology, phenology, ecology and known distribution range was provided. Apart from new taxa, the delimitation of already described species like Impatiens firmula Baker and Impatiens hildebrandtii Baill. could be clarified by studying the types and by observing the variability in the field.
Are the groups of Impatiens in Madagascar monophyletic, and what is the systematic position of Trimorphopetalum?
Yuan & al. (2004) conducted a molecular phylogenetic study to examine the morphological and karyological evolution, and the historical biogeography of the Balsaminaceae family by using nucleotide sequence data of internal transcribed spacer regions of nuclear ribosomal DNA. The results support the monophyly of the Malagasy endemic section Trimorphopetalum and show that the cleistogamous Impatiens inaperta should be included in the sect. Trimorphopetalum which is the most derived within Impatiens. Therefore, the section Preimpatiens proposed by Perrier de la Bâthie (1934) is paraphyletic.
Rahelivololona & al. (2018) provided a phylogenetic study focused on three subdivisions (based on macromorphological characters) proposed by Perrier de la Bâthie (1934). The analysis was done using two nuclear AP3/DEF homologues (ImpDEF1 and ImpDEF2) and the plastid atpB-rbcL spacer to reassess or assess the monophyly of the Malagasy Impatiens, of the sections Preimpatiens (Humblotianae and Vulgare groups) and Trimorphopetalum. A focus was on the species of Impatiens from the Marojejy National Park and of the morphologically variable species I. elatostemmoides, I. “hammarbyoides”, I. inaperta and I. manaharensis, using monophyly as the primary criterion.
As results the Malagasy Impatiens are paraphyletic and the section Preimpatiens sensu Perrier de la Bâthie (1934) (= subgen. Impatiens sensu Fischer & Rahelivololona 2002) was not resolved as a monophyletic group. The section Trimorphopetalum sensu Perrier de la Bâthie (1934) (= subgen. Trimorphopetalum sensu Fischer & Rahelivololona 2002), however, was strongly confirmed as a monophyletic lineage (BS: 92; BPP: 1). Neither the Humblotianae group nor the Vulgare group was supported as monophyletic. None of the morphologically variable species appeared to be monophyletic and the sampled species of Impatiens from the Marojejy National Park do also not form a monophyletic group.
What are the biogeographical position and the distribution patterns of Impatiens in Madagascar and the Comoro islands?
Investigation of the geographical affinities and species distribution of section Impatiens (including Humblotianae group and Vulgare group) and section Trimorphopetalum were conducted and the origin and evolution as well as species richness and endemism were discussed.
The isolation, the climate and the complex topography of Madagascar have generated the microhabitats and ecological niches favourable to the diversification of Impatiens species. Impatiens of Madagascar with 260 endemic species is actually the largest genus in Madagascar. Therefore, Madagascar and the Comoro islands are among the most species-rich regions in the world for Impatiens.
Future studies
In Impatiens on Madagascar, there remain numerous unresolved questions that need to be adressed:
• A further study based on a much larger molecular data set and sampling from the entire geographic ranges of Impatiens in Madagascar is needed to retest the monophyly of the different subgenera and sections, as well as a molecular dating of the Malagasy Impatiens.
• The study of pollinators as a key for understanding the radiation and species richness is required: Within Impatiens the different shapes of spur are related to pollinators (bees, birds, butterflies and moths). Therefore pollinator observation of specific species need to be done to understand the radiation of species by adaptation and coevolution with these pollinators. A pollination study with a large number of species within section Trimorphopetalum will help to understand the mechanism of complete disappearance of the spur, the shift of pollinators and the evolution of species richness.
• The destruction of the natural habitats of Impatiens and the subsequent reduction of humidity in logged area constitute a severe threat for the survival of many species. The conservation and reforestation of vulnerable areas such as Ankaratra, Daraina, Mandraka and Col des Tapia near Antsirabe is required.
• In terms of conservation and to mitigate the threat on the genus, a study on the ex-situ-conservation of Malagasy Impatiens species is very important as long as some species are suitable for horticultural purposes (e.g. Impatiens mayae-valeriae, Impatiens emiliae and species with broad red spur).
• Finally, the publication of the revision of Impatiens of Madagascar and the Comoro islands will help other botanists to identify the species and will thus increase our knowledge on the group.
Cultural eutrophication due to excessive inputs of nutrients seriously threatens aquatic ecosystems worldwide and is one of the major anthropogenic stressors on aquatic biota in European rivers. In streams and shallow rivers, its effects include excessive periphyton growth, which causes biological clogging and thereby oxygen depletion in the hyporheic zone. The result is a serious degradation of habitat quality for benthic invertebrates as well as for the eggs and larvae of gravel-spawning fish. Unlike in standing waters, efficient tools for controlling eutrophication in rivers are lacking. However, top-down control of the food-web by manipulating fish stocks, similar to the biomanipulation successfully applied in lakes, offers a promising approach to mitigating the effects of eutrophication in shallow rivers, especially those in which major reductions in nutrient inputs are not feasible. The overall aim of this thesis was to assess the potential for top-down control by two large cypriniform fish, the common nase (Chondrostoma nasus), the only obligate herbivorous fish species in European rivers, and the omnivorous European chub (Squalius cephalus), to mitigate the effects of eutrophication in medium-sized rivers. I therefore conducted field experiments on different spatial and temporal scales in the hyporhithral zone of a eutrophic gravel-bed river. Generally, the results of those experiments revealed the crucial role of fish-mediated top-down effects in river food webs. In a 4-year reach-scale experiment, the key contribution of my thesis, the enhancement of fish densities significantly increased both oxygen availability and water exchange in the upper layer of the hyporheic zone, even though the top-down effects of the fish on periphyton biomass were relatively small. These findings were supported by those of a 4-week mesocosm experiment, which also provided insights into the mechanisms underlying the mitigation of eutrophication effects by nase and chub. The top-down effects of both fish species reduced hyporheic oxygen depletion, suggesting a reduction of biological clogging. The positive effects of herbivorous nase on hyporheic oxygen availability could be attributed to benthic grazing, whereas the reduction of hyporheic oxygen depletion in the presence of omnivorous chub was best explained by the enhanced bioturbation induced by the fish’s benthic foraging. Overall, the results of my thesis demonstrate that biomanipulation achieved by enhancing herbivorous and omnivorous fish stocks can mitigate the effects of eutrophication in medium-sized European rivers. The results may be the first step towards the establishment of biomanipulation as a supportive management tool for eutrophication control in running waters and therefore as a strategy to preserve aquatic biodiversity.
Aim of this study was the assessment of the conservation status of vascular plants in East African rain forests with the background of establishing an ex-situ culture of local endangered plants at the Botanic Garden of the Maseno University (Kenya).
For a sustainable implementation it was first necessary to learn more about the general species inventory, especially concerning species composition and abundance under human impact, and to assess the conservation priority of each plant species. Representative for East African rain forests, Kakamega Forest (Kenya) and Budongo Forest (Uganda) were selected to serve as model forests.
Beside the general floristic investigations including all vascular plants, a special focus was laid on vascular epiphytes and their vulnerability to forest disturbance. To assess the conservation priority of the plants, a rating system was developed based on seven threat criteria. By carrying out first plant collections, the exsitu culture in Maseno Botanic Garden was already initiated.
Ecological assessment approaches based on benthic invertebrates in Euphrates tributaries in Turkey
(2019)
Sustainable water management requires methods for assessing ecological stream quality. Many years of limnological research are needed to provide a basis for developing such methods. However, research of this kind is still lacking in Turkey. Therefore, the aim of this doctoral thesis was to provide basic research in the field of aquatic ecology and to present methods for the assessment of ecological stream quality based on benthic invertebrates. For this purpose, I selected 17 tributaries of the Euphrates with a similar typology/water order and varying levels of pollution or not affected by pollution at all. The characterisation of the natural mountain streams was the first important step in the analysis of ecological quality. Based on community indices, I found that the five selected streams had a very good ecological status. I also compared the different biological indications, collected on two occasions ¬– once in spring (May) and once in autumn (September) – to determine the optimal sampling time. The macroinvertebrate composition differed considerably between the two seasons, with the number of taxa and Shannon index being significantly higher in autumn than in spring. In the final step, I examined the basal resources of the macroinvertebrates in the reference streams with an isotope analysis. I found that FPOM and biofilm were the most relevant basal resources of benthic invertebrates. Subsequently, based on the similarity of their community structures, I divided the 17 streams into three quality classes, supported by four community indices (EPT [%], EPTCBO [%], number of individuals, evenness). In this process, 23 taxa were identified as indicators for the three quality classes. In the next step, I presented two new or adapted indices for the assessment of quality class. Firstly, I adapted the Hindu Kush-Himalaya biotic index to the catchment area of the Euphrates and created a new, ecoregion-specific score list (Euph-Scores) for 93 taxa. The weighted ASPT values, which were renamed the Euphrates Biotic Score (EUPHbios) in this study, showed sharper differentiations of quality classes compared to the other considered ASPT values. Thus, this modified index has proved to be very effective and easy to implement in practical applications. As a second biological index, I suggested the proportion of habitat specialists. To calculate this index, the habitat preferences of the 20 most common benthic invertebrates were identified using the new habitat score. The proportion of habitat specialists differed significantly among the three quality classes with higher values in natural streams than in polluted streams. The methods and results presented in this doctoral thesis can be used in a multi-metric index for a Turkish assessment programme.
Because silver nanoparticles (Ag NPs) are broadly applied in consumer products, their leaching will result in the continuous release of Ag NPs into the natural aquatic environment. Therefore, bacterial biofilms, as the prominent life form of microorganisms in the aquatic environment, are most likely confronted with Ag NPs as a pollutant stressor. Notwithstanding the significant ecological relevance of bacterial biofilms in aquatic systems, and though Ag NPs are expected to accumulate within these biofilms in the environment, the knowledge on the environmental and ecological impact of Ag NPs, is still lagging behind the industrial growth of nanotechnology. Consequently, aim of this thesis was to perform effect assessment of Ag NP exposure on bacterial biofilms with ambient Ag NPs concentrations and under environmentally relevant conditions. Therefore, a comprehensive set of methods was applied in this work to study if and how Ag NPs of two different sizes (30 and 70 nm) affect bacterial biofilms i.e. both monospecies biofilms and freshwater biofilms in environmentally relevant concentrations (600 - 2400 µg l-1). Within the first part of this work, a newly developed assay to test the mechanical stability of
monospecies biofilms of the freshwater model bacterium Aquabacterium citratiphilum was validated. In the first study, to investigate the impact of Ag NPs on the mechanical stability of bacterial biofilms, sublethal effects on the mechanical stability of the biofilms were observed with negative implications for biostabilization. Furthermore, as it is still challenging to monitor the ecotoxicity of Ag NPs in natural freshwater environments, a mesocosm study was performed in this work to provide the possibility for the detailed investigation of effects of Ag NPs on freshwater biofilms under realistic environmental conditions. By applying several approaches to analyze biofilms as a whole in response to Ag NP treatment, insights into the resilience of bacterial freshwater biofilms were obtained. However, as revealed by t-RFLP fingerprinting combined with phylogenetic studies based on the 16S gene, a shift in the bacterial community composition, where Ag NP-sensitive bacteria were replaced by more Ag NP-tolerant species with enhanced adaptability towards Ag NP stress was determined. This shift within the bacterial community may be associated with potential detrimental effects on the functioning of these biofilms with respect to nutrient loads, transformation and/or degradation of pollutants, and biostabilization. Overall, bringing together the key findings of this thesis, 4 general effect mechanisms of Ag NP treatment have been identified, which can be extrapolated to natural freshwater biofilms i.e. (i) the identification of Comamonadaceae as Ag NP-tolerant, (ii) a particular resilient behaviour of the biofilms, (iii) the two applied size fractions of Ag NPs exhibited similar effects independent of their sizes and their synthesis method, and (iv) bacterial biofilms show a high uptake capacity for Ag NPs, which indicates cumulative enrichment.
Foliicolous lichens are one of the most abundant epiphytes in tropical rainforests and one of the few groups of organisms that characterize these forests. Tropical rainforests are increasingly affected by anthropogenic disturbance resulting in forest destruction and degradation. However, not much is known on the effects of anthropogenic disturbance on the diversity of foliicolous lichens. Understanding such effects is crucial for the development of appropriate measures for the conservation of such organisms. In this study, foliicolous lichens diversity was investigated in three tropical rainforests in East Africa. Godere Forest in Southwest Ethiopia is a transitional rainforest with a mixture of Afromontane and Guineo-Congolian species. The forest is secondary and has been affected by shifting cultivation, semi-forest coffee management and commercial coffee plantation. Budongo Forest in West Uganda is a Guineo-Congolian rainforest consisting of primary and secondary forests. Kakamega Forest in western Kenya is a transitional rainforest with a mixture of Guineo-Congolian and Afromontane species. The forest is a mosaic of near-primary forest, secondary forests of different seral stages, grasslands, plantations, and natural glades.
The ongoing loss of species is a global threat to biodiversity, affecting ecosystems worldwide. This also concerns arthropods such as insects and spiders, which are especially endangered in agricultural ecosystems. Here, one of the main causing factors is management intensification. In areas with a high proportion of traditionally managed grassland, extensive hay meadows that are cut only once per year can still hold high levels of biodiversity, but are threatened by conversion into highly productive silage grassland. The Westerwald mountain range, western Germany, is such a region. In this thesis, I compare the local diversity of bees, beetles, hoverflies, leafhoppers, and spiders of five grassland management regimes along a gradient of land-use intensity. These comprise naturally occurring grassland fallows, three types of traditionally managed hay meadows, and intensively used silage grassland. By using three different sampling methods, I recorded ground-dwelling, flower-visiting, and vegetation-dwelling species. The results show that in most cases species richness and diversity are highest on fallows, whereas variation among different managed grassland types is very low. Also, for most sampled taxa, fallows harbour the most distinct species assemblages, while that of other management regimes are largely overlapping. Management has the largest effect on species composition, whereas environmental parameters are of minor importance. Long-term grassland fallows seem to be highly valuable for arthropod conservation, even in a landscape with a low overall land-use intensity, providing structural heterogeneity. In conclusion, such fallows should be subsidized agri-environmental schemes, to preserve insect and spider diversity.
In the present study the flora and vegetation of Kakamega Forest, an East African rainforest in Western Kenya, was investigated. Kakamega Forest is highly degraded and fragmented and is an ideal model to study the anthropogenic influence on the forest inventory. The main focus was to analyse the influence of human impact on the vascular plant species composition. During five field phases in the years 2001 to 2004 a total of 19 study sites scattered over the whole forest including all fragments were investigated regarding forest structure, species composition and plant communities. The different forest sites were analysed by three different methods, phytosociological relevés, line-transect and with the variable-area transect method. The forest survey revealed about 400 taxa of vascular plant species, among them 112 trees, 62 shrubs, 58 climbers and 114 herbs. Several species are restricted to this forest in Kenya, but only one endemic species, the herb Commelina albiflora, could be discovered. About 15 species were recorded as new for Kenya and probably at least one species is new to science. Kakamega Forest is a unique mixture of Guineo-Congolian and Afromontane floral elements. About one half of the vascular plant species has its origin in the lowland forests of the Congo basin and one third originates from Afromontane habitats. The present study represents the first description of plant communities of Kakamega Forest. An analysis of different forest sites and plantations resulted in 17 different vegetation units. For the mature forest sites eleven plant communities were described. The young succession stage consists of two plant communities. Since the disturbance history and the age of the different plant communities could be estimated, their chronology was also described. An exception are the study sites within the plantations and afforested sites. The four defined vegetation units were not described as plant communities, because they are highly affected by man and do not belong to the natural succession of Kakamega Forest. Nevertheless, the regeneration potential of such forests was investigated. Due to the different succession stages the changing species composition along a disturbance gradient could be analysed. Most of Kakamega Forest consists of middle-aged secondary forest often surrounded by very young secondary forest. A true primary rainforest could not be found due the massive influence by over-exploitation. In all parts of the forest the anthropogenic influence could be observed. The forest develops towards a climax stage, but a 2 Abstract comparison with former surveys shows that the regeneration is much slower than expected. Human impact has to be avoided to allow the forest to develop into a primary-like rainforest. But several climax tree species might be missing anyway, because after the broad logging activities in the past there are not enough seed trees remaining. Species richness was highest in disturbed forest sites. A mixture of pioneer, climax and bushland species could be recorded there. Therefore, a high species richness is not a suitable indicator for forest quality. The proportion of climax species typical for Kakamega Forest would be a better measure. Compared to the main forest block the forest fragments do not lack in diversity as expected due to fragmentation processes. Instead, the only near primary forest could be recorded in Kisere, a northern fragment. The high amount of climax species and the more or less undisturbed forest structure is a result of the strict protection by the Kenya Wildlife Service and due to low logging activities. Differences in species composition between the studied forest sites are either a result of the different logging history or management regime rather than due to different edaphic or climatic conditions.
Human population pressure increased with the population growth around the NNP and Cyamudongo with disturbance impacts on the forests isolating populations into fragments and today, Cyamudongo natural forest is located a way at a distance of at least 8.5 km horizontal distance to Nyungwe main block with a surface area estimated at 300 ha. Under Cyamudongo project implementation, there was a need to understand how the flora diversity responded to human imposed challenges and to forest restoration initiatives. Three physiognomic landscapes forest were identified and considered for three phases of vegetation survey in Cyamudongo and related to the closest area of Nyungwe main block. In this study, 15 transects were laid in each physiognomic forest landscape and 10 and 5 plots were set respectively in Cyamudongo and Gasumo study area. In total, 315 phytosociological relevés were performed and the Braun-Blanquet methods used for three times vegetation surveys. Species life-forms and chorophyte were evaluated and tree species dbh and height have been measured. Data were subjected to different statistical analyses using different softwares such as PAST, R 3.5.2, and SPSS. The mapping was done using Arc GIS and the Multi-Spectral Remote Sensing used to find NDVI for the vegetation classification.
NDVI trends showed that there has been fluctuations in vegetation classifications of the studied area. In this study, 494 vascular plant species from 106 families were harbored in the study area and distributed differently among forest landscapes and study phases. Although, 43.54% were common to Cyamudongo and Gasumo landscapes while 48.54 % of species diversity were hold only by Cyamudongo and 7.92% confined to Gasumo and 12 in total were found new records for Rwanda while several others suspected require detailed research for identification showing how the flora diversity of Cyamudongo is of special interest and extremely important for discoveries.
The finding of the study on diversity indices, the PCA, CA and the Cluster analysis, all statistical analyses (MANOVA, ANOVA) and life form spectra unanimously showed that the anthropogenic disturbance shaped the vegetation cover, the floristic composition, the species diversity, the forest landscapes community structure, the life form spectrum and the phytoclimate of Cyamudongo and Gasumo forest landscapes. Although, the vegetation analysis couldn’t clearly identify communities and sub-communities at the initial and final vegetation surveys and cluster groups were heterogeneous as well as overlapping and species associations not clearly defined due to the high level of similarities in species composition among forest landscapes and vegetation surveys. The species diversity was found high in secondary forest and Gasumo landscape forest and low in the primary forest and the buffer zone of Cyamudongo and the disturbance with gaps openings was found to be associated to the species diversity with a seasonal variation. The patterns of dbh for the buffer zone and of the size classes of all landscapes with an inverted ‘J’ indicated a healthy regeneration in the forest landscapes and tree species explained a good regeneration and recruitment capacity. Different shapes in the pattern of dbh with respectively an inverted ‘J’, ‘J ‘and ‘U’ for the buffer zone, primary and together the secondary and Gasumo forest landscapes indicated differences in the landscapes health and degree of regeneration and recruitment capacity.
Findings from differents measuements showed at which extend human activities have shaped the flora diversity and structure of forest landcapes studied. For instance, disturbances due human activities were daily oberved and trees were logged by neighboring communities such as Batwa populations at Cyamudongo and local populations at Gasumo. Some species were evenly observed targeted for their barks such as Ocotea usambarensis, Parinari excelsa for medicines and many others for their wood quality, fire wood collection and for agricultural purposes.
In the period of Cyamudongo project implementation, important achievements included the increase of forest biomass and therefore the photosynthetic capacity and the evapotranspiration potential that influence the rainfall regime; the regulation of weather conditions and then species diversity; supporting local communities and limiting human activities; raising awareness on conservation and protection of biodiversity and improving of living conditions of neighboring populations by providing paid employment and so to restore to the Cyamudongo forest ecosystem functions. Moreover, Cyamudongo forest remains vulnerable as surrounded by local communities with a high population pressure relying on forest resources for its survival. Cyamudongo harbors a high level of endemism and is a small hotspot for biodiversity conservation. It is therefore recommended to strengthen conservation and protection measures and continue the support of local communities.