Institut für Integrierte Naturwissenschaften, Abt. Biologie
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Coat color and pattern are a distinguished feature in mammalian carnivores, shaped by climatic cycles and habitat type. It can be expressed in various ways, such as gradients, polymorphisms, and rare color variants. Although natural selection explains much of the phenotypic variation found in the wild, genetic drift and heterozygote deficiency, as prominent in small and fragmented populations, may also affect phenotypic variability through the fixation of recessive alleles. The aim of this study was to test whether rare color variants in the wild could relate to a deficiency of heterozygotes, resulting from habitat fragmentation and small population size. We present an overview of all rare color variants in the order Carnivora, and compiled demographic and genetic data of the populations where they did and did not occur, to test for significant correlations. We also tested how phylogeny and body weight influenced the presence of color variants with phylogenetic generalized linear mixed models (PGLMMs). We found 40 color-variable species and 59 rare color variants. In 17 variable phenotypic populations for which genetic diversity was available, the average AR was 4.18, HO = 0.59, and HE= 0.66, and FIS= 0.086. We found that variable populations displayed a significant reduction in heterozygosity and allelic richness compared to non-variable populations across species. We also found a significant negative correlation between population size and inbreeding coefficients. Therefore, it is possible that small effective size had phenotypic consequences on the extant populations. The high frequency of the rare color variants (averaging 20%) also implies that genetic drift is locally overruling natural selection in small effective populations. As such, rare color variants could be added to the list of phenotypic consequences of inbreeding in the wild.
The diversity within amphibian communities in cultivated areas in Rwanda and within two selected, taxonomically challenging groups, the genera Ptychadena and Hyperolius, were investigated in this thesis. The amphibian community of an agricultural wetland near Butare in southern Rwanda comprised 15 anuran species. Rarefaction and jackknife analyses corroborated that the complete current species richness of the assemblage had been recorded, and the results of acoustic niche analysis suggested species saturation of the community. Surveys at many other Rwandan localities showed that the species recorded in Butare are widespread in cultivated and pristine wetlands. The species were readily distinguishable using morphological, bioacoustic, and molecular (DNA barcoding) features, but only eight of the 15 species could be assigned unambiguously to nominal species. The remaining represented undescribed or currently unrecognized taxa, including three species of Hyperolius, two Phrynobatrachus species, one Ptychadena species, and one species of Amietia. The diversity of the Ridged Frogs in Rwanda was investigated in two studies (Chapters III and IV). Three species of Ptychadena were recorded in wetlands in the catchment of the Nile. They can be distinguished by morphological characters (morphometrics and qualitative features) as well as by their advertisement calls and genetics. The Rwandan species of the P. mascareniensis group was shown to differ from the topotypic population as well as from other genetic lineages in sub-Saharan Africa and an old available name, P. nilotica, was resurrected from synonymy for this lineage. Two further Ptychadena species were identified among voucher specimens from Rwanda deposited in the collection of the RMCA, P. chrysogaster and P. uzungwensis. Morphologically they can be unambiguously distinguished from each other and the three other Rwandan species. A key based on qualitative morphological characters was developed, which allows unequivocal identification of specimens of all species that have been recorded from Rwanda. DNA was isolated from a Rwandan voucher specimen of P. chrysogaster, and the genetic analysis corroborated the species" distinct status.
A species of Hyperolius collected in the Nyungwe National Park was compared to all other Rwandan species of the genus and to morphologically or genetically similar species from neighbouring countries. Its distinct taxonomic status was justified by morphological, bioacoustic, and molecular evidence and it was described as a new species, H. jackie. A species of the H. nasutus group collected at agricultural sites in Rwanda was described as a new species in the course of a revision of the species of the Hyperolius nasutus group. The group was shown to consist of 15 distinct species which can be distinguished from each other genetically, bioacoustically, and morphologically.
The aerial performance, i.e. parachuting, of the Disc-fingered Reed Frog, Hyperolius discodactylus, was described. It represents a novel observation of a behaviour that has been known from a number of Southeast Asian and Neotropical frog species. Parachuting frogs, including H. discodactylus, exhibit certain morphological characteristics and, while airborne, assume a distinct posture which is best-suited for maneuvering in the air. Another study on the species addressed the validity of the taxon H. alticola which had been considered either a synonym of H. discodactylus or a distinct species. Type material of both taxa was re-examined and the status of H. alticola reassessed using morphological data from historic and new collections, call recordings, and molecular data from animals collected on recent expeditions. A northern and a southern genetic clade were identified, a divide that is weakly supported by diverging morphology of the vouchers from the respective localities. No distinction in advertisement call features could be recovered to support this split and both genetic and morphological differences between the two geographic clades are marginal and not always congruent and more likely reflect population-level variation. Therefore it was concluded that H. alticola is not a valid taxon and should be treated as a synonym of H. discodactylus.
This study was conducted in Nyungwe National Park (NNP); a biodiversity hotspot Mountain rainforest of high conservation importance in Central Africa, but with little knowledge of its insect communities including butterflies, good indicators of climate change, and forest ecosystem health. The study aimed at availing baseline data on butterfly species diversity and distribution in NNP, for future use in monitoring climate change-driven shifts and the effects of forest fragmentation on the biodiversity of Nyungwe. Butterflies were collected seasonally using fruit-baited traps and hand nets along elevational transects spanning from 1700 m up to 2950 m of altitude. Two hundred forty-two species including 28 endemics to the Albertine Rift and 18 potential local climate change indicators were documented. Species richness and abundance declined with increasing elevation and higher seasonal occurrence was observed during the dry season. This was the first study on the spatial and temporal distribution of butterflies in NNP and further studies could be conducted to add more species and allow a depth understanding of the ecology of Nyungwe butterflies.
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.
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.
Eine Ursache des Insektenrückgangs ist die Abnahme der für Arthropoden wichtigen Lebensräume. Der kleinstrukturierte Obstanbau als Dauerkultur mit vielfältigen Strukturen (Bäume, Grünland, offener Boden) kann als Refugium wirken. Gerade Dauerkulturen, welche über viele Jahre bestehen bleiben, können einen großen ökologischen Wert für Insekten und Spinnentiere darstellen, wenn die negativen Einflüsse der Bewirtschaftung durch gezielte Maßnahmen minimiert werden. Im Gegensatz zu den gut untersuchten Streuobstwiesen liegen bisher nur wenige oder ungenaue Daten über die Arthropodenfauna in Erwerbsobstanlagen vor. Ziele dieser Arbeit sind eine genauere Erfassung der Arthropodenfauna in Erwerbsobstanlagen, Benennung der Einflussfaktoren und Erarbeitung einer indikatorgestützten Bewertungsmethode mit Entwicklung von Maßnahmen zur Förderung der Biodiversität. Es wurde eine intensive Erfassung der Insekten und Spinnentiere auf Basis von Individuenzahlen, Verteilung der Tiere auf Großgruppen, Artenzahlen der Käfer (inkl. Rote Liste) und Wanzen, deren Diversität und der sie beeinflussenden Faktoren auf vier Erwerbsobstflächen und einer Streuobstwiese als Referenzfläche in der Gemeinde Zornheim (Rheinland-Pfalz) untersucht. Mittels des erarbeiteten Bewertungsbogens wurden die Auswirkungen der fünf Faktorenkomplexe: Sonderstrukturen, Anlagenstruktur, Beschattung, Mahdregime und Einsatz von Pflanzenschutzmitteln auf die Arthropodenfauna erhoben und die auf der Fläche zu erwartende Biodiversität ermittelt. Die visuelle Darstellung der Einflussfaktoren mit Netzdiagrammen zeigt biodiversitätsbeeinträchtigende Faktoren und ermöglicht so Empfehlungen zur Aufwertung der Biodiversität. Die Bewertungsmethode kann nach Verifizierung im Rahmen von erfolgsorientierten Agrarumwelt- und Klimamaßnahmen (AUKM), zur Anerkennung von Kompensationsmaßnahmen (z.B. PIK) und zur Ermittlung des ökologischen Wertes (Ökosystemdienstleistung) von Obstanlagen genutzt werden. In modifizierter Form könnte es ebenfalls im Bereich der Flächenbewertung in Landschafts- und Bauleitplanung eingesetzt werden.
The stands surveyed are among the last closed canopy forests in Rwanda. Their exploration began in the early twentieth century and is still ongoing. Previous studies were mainly concerned with plant sociological issues and presented references to environmental factors in anecdotal form, at best using indirect ordination methods. The present study undertakes a classification of the vegetation with numerical methods and establishes quantitative relationships of the species’ distributional structure to environmental parameters using spatially explicit procedures. For this purpose, 94 samples were taken in 100 m² hexagonal plots. Of these, 70 samples are from Nyungwe, 14 are from Gishwati, and 10 are from Cyamudongo. Given the homogeneity of the terrain and vegetation, all vegetation types encountered, all types of stands, and all vegetation strata were included. The beta diversity is expressed by an average Bray-Curtis dissimilarity of 0.92, and in JOST’S (2007) numbers equivalents, 37.90 equally likely samples would be needed to represent the diversity encountered. Within the survey, 1198 species in 127 families were collected. Among the specimens are 6 local endemics and 40 Albertine Rift endemics. Resulting from UPGMA and FCM-NC, 20 to 40 plant communities were established depending on the level of resolution. It can be inferred by means of a Mantel correlogram that the mean zone of influence of a single vegetation stand, as sampled by a 100 m² plot in Nyungwe Forest, ranges between 0.016 and 3.42 km. Of the communities compiled using FCM-NC and UPGMA, 50% consist of individual samples. Beyond undersampling, natural small-scale discontinuities are reflected by this result. Partial db-RDA resulted in an explained variation of 9.60% and 14.41% for environmental and soil factors, respectively. Utilising variation partitioning analyses based on CCA and tb-RDA, between 21.70% and 37.80% of the variation in vegetation data could be explained. The spatially structured fraction of these parameters accounts for between 30.50% and 49.80% of the explained variation (100%). The purely environmental parameters account for a share of 10.30% to 16.30%, whereby the lower limit originates from the unimodal approach and has lost its statistical significance. The soil variables, also after partial analysis, account for a share of 19.00% to 35.70%. While the residual impact of the climatic parameters is hardly significant, the effect of the soil properties is prevalent. In general, the spatially structured fraction of the parameters is predominant here. While on the broad-scale climatic factors, the altitude a.s.l. and the geology are determining factors, some soil parameters and matrix components also show their impacts here. In the mid-range of the scale, it is the forest matrix, the soil types, and the geology that determine species distribution. While in the fine range of the scale, some unrecorded parameters seem to have an effect, there are also neutral processes that determine species composition.
The protected areas of Rwanda are facing various challenges resulting from the anthropogenic activities of the surrounding communities especially in the adjacent area to Cyamudongo isolated rain forest, which results in climate change, soil degradation, and loss of biodiversity. Therefore, this study aims to broaden current knowledge on the impact of sustainable Agroforestry (AF) on the Carbon (C) stock and Biodiversity conservation on the surroundings of Cyamudongo isolated rain forest and Ruhande Arboretum.
To understand this, the permanent sample plots (PSPs) were established mainly in the designed four transects of four km long originating on the boundary of the Cyamudongo isolated rain forest following the slope gradient ranging from 1286 to 2015 m asl. A total number of 73 PSPs were established in the Cyamudongo study area while 3 PSPs were established in the Ruhande AF plot. The Arc Map GIS 10.4 was used to design and map the sampling areas while GPS was used for localization of collected items. Statistical significance was analyzed through the R-software especially for wood and soil variables while for biodiversity indicator species, MVSP Software 3.0 was used to determine the Shannon Diversity indices and similarities among species.
In this study, I have obtained comprehensive results demonstrating that in all study areas, the various AF tree species contribute differently to C stock and C sequestration and the amount of C stored and removed from the atmosphere depends on different factors such as tree species, plantation density, growth stage, or the age of establishment, applied management practices, wood specific density (WSD), wood C concentration, and climatic conditions. The estimated quantity of sequestrated C for 2 years and 34 years AF species were 13.11 t C ha -1 yr-1 (equivalent to 48 t CO2 ha -1 yr-1) and 6.85 t ha-1 yr-1 (equivalent to 25.1 t CO2 ha -1 yr-1) in Cyamudongo and Ruhande respectively. The estimated quantity of C stored by the Ruhande AF plot is 232.94 t ha-1. In Cyamudongo, the overall C stored by the AF systems was 823 t ha-1 by both young tree species established by the Cyamudongo Project (35.84 t ha-1) and C stored by existed AF species before the existence of the Project (787.12 t ha-1). In all study areas, the Grevillea robusta was found to contribute more to overall stored C compared to other species under this study.
The tests revealed differences in terms of nutrient contents (C, N, C: N ratio, K, Na, Ca, and Mg) for various AF tree species of Cyamudongo and Ruhande study areas. The differences in terms of correlation for various variables of AF tree species in different study areas varied with tree species, age, stage of growth, and tree shape. By comparing the correlation coefficients for various tree variables for young and mature AF tree species, the results showed a high correlation variability for young species than mature or old species recorded in different environmental conditions of Cyamudongo and Ruhande study areas.
The recorded soil pH mean value across in Cyamudongo study area is 4.2, which is very strongly acidic. The tests revealed that the soil pH, C, C: N ratio, OM, NH4+, NO3-+NO2-, PO43-, and CEC were significantly (P < 0.05) different in various soil depths whereas the N was not statistically significant. The pH, N, C: N ratio, CEC, NH4+, PO43-, and Al3+ showed a significant difference across land uses whereas the C and NO3-+NO2- did not show any statistical difference. All tested chemical elements showed a statistical difference as far as altitude ranges are concerned. The only NH4+, PO43-, and CEC showed significant differences with time whereas all other remaining chemical elements did not show any statistical significance. The bulk density of soil was statistically different across land uses and altitude ranges. The soil pH was very strongly correlated with CEC, Mg, and Ca in cropland (CL) whereas it was strongly correlated in both AF and natural forest (NF) except for Mg, which was moderately correlated in AF. Furthermore, its correlation with K was strong in CL, moderate in AF while it was weak in NF. Finally, the pH correlation with Na was weak in both AF and CL whereas it was negligible in NF. The overall estimated soil C stock of the study area was 16848 t ha -1.
The sustainable AF practices changed significantly the frequency of reptiles, amphibians, and flowering plants while there was no statistical change observed on ferns with time. In terms of species richness, 16 flowering plants, 14 ferns, 5 amphibians, and 3 reptiles were recorded and monitored. These findings add to a growing body of literature on the impact of AF on the C stock, soil improvement, and Biodiversity. It is recommended that further researches should be undertaken for the contribution of other AF tree species to the C stock found in the agricultural landscape around all protected areas of Rwanda and the impact on them on the soil and biodiversity.
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.
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.