Institut für Integrierte Naturwissenschaften, Abt. Biologie
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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.
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.
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.
The implementation of physiological indicators reflecting the response of organisms to changes in their environment is assumed to provide potential benefits for ecological studies. By analysing the physiological condition of organisms in freshwater ecological studies rather than their ultimate effects, physiological indicators can contribute to a faster assessment of effects than using traditional ecological indicators, such as the evaluation of the benthic community structure or the determination of the reproductive success of organisms. This can increase the effectiveness of environmental health assessment and experimental ecology. In this respect the thesis focuses on physiological measures characterizing the energetic condition and energy consumption (the concentration of energy storage compounds, the adenylate energy charge, the energy consumption in vivo), as well as individual growth (RNA:DNA ratio) of organisms. Although these sub-individual indicators are commonly applied in marine ecology and more recently in ecotoxicology, they have been rarely applied in freshwater ecology to date. With respect to an increased use of physiological indicators in freshwater ecological studies, the objectives of the present thesis are twofold. First, it highlights the potential of assessing the individual fitness by means of physiological indicators in freshwater ecological studies. For that reason, Chapter 2 provides the basic assumptions as well as the theoretical and methodological fundamentals necessary for the application of physiological indicators within freshwater ecology and, furthermore, points out their applicability by several case studies. As second objective, the thesis addresses selected ecophysiological aspects of native and non-native freshwater amphipods, which are considered suitable candidates for the determination of physiological indicators in ecological studies due to their function as keystone species within aquatic habitats. The studies presented in Chapters 3−5 of the thesis provide information on (i) species- and sex-specific seasonal variations within the energetic condition of natural Gammarus populations (G. fossarum, G. pulex), (ii) differences in metabolic activity and behaviour between different amphipod species (G fossarum, G. roeselii and D. villosus), as well as (iii) the direct effects of ambient ammonia on the physiology and behaviour of D. villosus. The fundamental conclusions drawn from the conducted field and laboratory studies, as well as their relevance and general implications for the application of physiological indicators in freshwater ecological research are discussed in Chapter 6.
Taxonomy and Systematics of Spiny-Backed Treefrogs, Genus Osteocephalus (Amphibia: Anura: Hylidae)
(2015)
The pan-Amazonian treefrog genus Ostecephalus is poorly understood both on a taxonomic and phylogenetic level. The status of several frogs already or not yet referred to the genus is unclear and the relationships among the genus and with respect to related genera is not understood. In this work O. cabrerai (Cochran and Goin, 1970) from Colombia and Peru is redescribed and O. festae (Peracca, 1904) from the foothills of the Andes in Ecuador is revalidated. Hyla inframaculata Boulenger, 1882, from the lower Amazon in Brazil, is reallocated to Osteocephalus and O. elkejungingerae (Henle, 1981) from the Andean foothills in Peru is shown to be a synonym of Hyla mimetica (Melin, 1941), the valid name being O. mimeticus. Hyla vilarsi Melin, 1941 is considered a valid species in the genus Osteocephalus and revalidated from the synonymies of several other frogs. Three new species, O. castaneicola from northern Bolivia and southern Peru, O. duellmani from a sub-Andean mountain range in southern Ecuador, and O. camufatus from central Amazonian Brazil, are described. A phylogenetic analysis based on up to nine mitochondrial genes and one nuclear one reveals the paraphyly of the genus as previously understood with respect to the genus Tepuihyla. A new taxonomy is proposed, securing the monophyly of Osteocephalus and Tepuihyla by rearranging and redefining the content of both genera. A new genus, Dryaderces, is erected for the sister group of Osteocephalus. The colouration of newly metamorphosed individuals is proposed as a morphological synapomorphy for Osteocephalus. Five monophyletic species groups within Osteocephalus are recognized, three species of Osteocephalus (O. germani, O.rnphasmatus, O. vilmae) and three species of Tepuihyla (T. celsae, T. galani, T. talbergae) synonymized and three species (Hyla helenae to Osteocephalus, O.rnexophthalmus to Tepuihyla and O. pearsoni to Dryaderces gen. n.) reallocated. Furthermore, nine putative new species are flagged (an increase to 138% of the current diversity), an indication that species numbers are largely underestimated, with most hidden diversity centred on widespread and polymorphic nominal species. The evolutionary origin of breeding strategies within Osteocephalus is discussed in the light of this new phylogeny and a novel type of amplexus (gular amplexus) is described.
A taxonomic revision of the genus Pteris in tropical Africa revealed 26 species. An identification key to the species is provided. Morphological characters were used to prepare a cladistic analysis of the relevant taxa. Each species was evaluated concerning the IUCN red list status. Only Pteris mkomaziensis was considered as Near Threatened, and all other species only as Least Concern. An inventory of the ferns of Kakamega Forest / Kenya and Budongo Forest / Uganda revealed 85 species in Kakamega and 66 species in Budongo. Life form spectra were analysed and the ferns were studied for their value for bioindication.
The first group that was revised within my study is Ochralea Clark, 1865 (Hazmi & Wagner 2010a). I have checked the type specimen of most species that were originally described in Ochralea and there is no doubt that this genus is clearly distinct from Monolepta. Weise (1924) has synonymised Galeruca nigripes (Olivier, 1808) with O. nigricornis Clark, 1865 and the valid name of the species is O. nigripes (Olivier, 1808). Out of ten species originally described in this genus, only this species remain valid and O. pectoralis is a new synonym of O. nigripes. Additionally, Monolepta wangkliana Mohamedsaid, 2000 is very closely related to O. nigripes and need to be transferred to Ochralea. The second genus where the revision is still published is Arcastes Baly, 1865 (Hazmi & Wagner 2010b). I have checked the genitalic characters of A. biplagiata, and most of the type species of other Arcastes. Arcastes biplagiata possesses a peculiar shape of the median lobe and asymmetrically arranged endophallic structures. These peculiar characters are very useful to delimit this genus from the others. Therefore, only three valid species remain in Arcastes, while two new synonyms are found and fourrnother species need to be transferred to other genera. While checking the genitalic characteristics of type species of Arcastes sanguinea, thernmedian lobe as well as the spermatheca of this species possesses strong differences to A. biplagiata. The species was redescribed and transferred in a monotypic new genus Rubrarcastes Hazmi & Wagner, 2010c. The fourth genus that was already revised is Neolepta Jacoby, 1884. It was originally described on base of only two species by that time, N. biplagiata and N. fulvipennis. Jacoby has not designated a type species of the genus, and Maulik (1936) did it later, with the designation of N. biplagiata. Jacoby in his original description has only commented that Neolepta is very close and similar to Monolepta Chevrolat, 1837 and Candezea Chapuis, 1879. Subsequent authors have described further eight species, and transferred one species from Luperodes to it, summing up the total number of eleven described species in Neolepta. I have checked the genitalic characters of the type, N. biplagiata and have found out that the median lobe is not incised apically and stronger sclerotised ventral carinae with an apical hook close to the apex occur. Out of all described species, only two are closely related to the genero-type, N. sumatrensis (Jacoby, 1884) new combination and N. quadriplagiata Jacoby, 1886 that will remain in this group after the revision. All other species need to be transferred to other genera, including the newly described Paraneolepta and Orthoneolepta. The last distinct paper of this thesis presented the results on Monolepta Chevrolat, 1837. The massive number of Monolepta from the entire Oriental Region, with about 260 described species names is a more long-life project and not practicable within a PhD-study. Thus I have focused on the species of Monolepta known from the Sundaland area in this work. A comprehensive revision including the study of the primary types of the described species, has never been done for Monolepta from this sub-region, while new species have also been described in the last decade (e. g. Mohamedsaid 1993, 1997, 1998, 1999, 2000a,b, 2001, 2002, 2005).
On base of the most current species lists of Mohamedsaid (2001, 2004, 2005) and Kimoto (1990), the number of valid species described from this region is about 72. After my revision, only thirteen valid species can remain in Monolepta in the sense of the generotype M. bioculata (Wagner 2007), while seven species have been found as new synonyms, three have been already transferred to other genera and further 49 species need to be transferred to other genera.
Nandi forests (South and North Nandi forests) are situated in the Rift Valley Province of Kenya very close to Kakamega forest. From previous documents it has been seen that Kakamega and Nandi forests were connected to each other forming one big "U" shaped forest block till the beginnings of 1900s. Due to human pressures, currently there are three different forests form the previous one block forest. Although they were one forest, information on Nandi forests is very scanty when it is compared to that of Kakamega forest. The species composition and diversity as well as plant communities and population structure of Nandi forests have not been studied. Information is not available about the similarity status of South and North Nandi forests. Furthermore the natural regeneration potential (seedling bank) of these forests is not well studied and documented. Hence this study aims to fill these gaps.
In this study totally 76 quadrates (49 from South Nandi and 27 from North Nandi) were used to collect data. In the South Nandi forests 27 of the quadrates were laid in the better side of the forest (at Kobujoi) and the remaining 22 were in the heavily disturbed part of this forest (Bonjoge). The quadrates were arranged on transects that have one to one and half km which were parallel to the slope. The distance between the quadrates was 100 meter and transects are 500 m apart. The size of the main quadrate was 400 m2 (20 X 20 m) which also had five small plots (3 X 3 m) distributed on the four corners and in the center. Each woody plants (climbers, shrubs and trees) having more than one meter and greater than two centimeter diameter at breast height (dbh) were measured and recorded. Seedlings and herbaceous plants were sampled in the smaller plots. Individual plants were identified at species level and when it was not possible to identify in the field voucher specimen were prepared and latter identified at the East African Herbarium, National Museum of Kenya, and Nairobi. Clustering and ordination were performed using PC-ORD and CANOCO ecological softwares, respectively. For both clustering and ordination abundance data of the species was used. Shannon diversity index and evenness were computed using PC-ORD while similarity indices, Fisher alpha, rarefaction, species richness estimation (nonparametric species richness estimators) were conducted using EstimateS. Indicator species analysis was undertaken using PC-ORD. Basal area and height class distribution at forests level or site level (Bonjoge and Kobujoi) and diameter (dbh) class distribution for selected trees species were performed to evaluate population structure.
Furthermore importance value (IV) of woody plant species was calculated. SPSS version 16 was used to undertake both parametric (when data assume normal distribution) and nonparametric (when data are not assuming normal distribution) comparison of means, correlation and regression analysis.
In this study totally 321 vascular plant species comprising 92 families and 243 genera were identified in Nandi forests (both South and North Nandi forests). In South Nandi forest 253 plant species form 82 families and 201 genera were recorded while in North Nandi 181 species comprising 67 families and 155 genera were recorded. Jackknife second order estimators gave the highest species richness estimate for both South and North Nandi forests i.e. 284 and 209, respectively. In the case of highly disturbed and less disturbed parts of South Nandi forest 138 and 172 vascular plant species were recorded, respectively. Asteraceae, Rubiaceae and Euphorbiaceae are the top three species rich families of Nandi forests. In terms of different diversity measures (i.e. alpha and beta diversity, Fisher alpha, Shannon diversity and evenness indices) South Nandi is more diverse than North Nandi forest. Sörensen and Jaccard (classic) as well as their respective abundance based similarities showed that there is a low species similarity between South and Nandi forests. The cluster analysis resulted in three different plant communities and this result is supported by the ordination result.
South and North Nandi forest has inverted "J" height class distribution showing that larger proportion of woody plant individuals are found in the lower height classes. Similar pattern is observed when the diameters of all woody plants were considered together. However, different diameter class distributions (seven types) were identified when selected tree species were analyzed separately. It has been observed that the basal area of South Nandi forest is significantly lower than that of North Nandi forest (Mann-Whitney U =358, p < 0.001). Similarly Bonjoge has significantly lower basal area (t-value=3.77, p<0.01) than that of Kobujoi. Number of woody plat seedlings in South Nandi forest is significantly higher than that of North Nandi (Mann-Whitney U = 362.5, p<0.001). In the same way Bonjoge has significantly smaller number of ssedlings than Kobujoi (t-value 4.24, p<0.001). Most of species in both forests are able to resprout from stumps after physical damage; hence this helps the regeneration of the forests in addition to seedling banks. This study enables to fill some of the information gaps about Nandi forests especially of floristic composition, population structure, natural regeneration and human impacts on this ecosystem.
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.
More than 10,000 organic chemicals such as pharmaceuticals, ingredients of personal care products and biocides are ubiquitously used in every day life. After their application, many of these chemicals enter the domestic sewer. Research has shown that conventional biological wastewater treatment in municipal wastewater treatment plants (WWTPs) is an insufficient barrier for the release of most of these anthropogenic chemicals into the receiving waters.
This bears unforeseen risks for aquatic wildlife and drinking water resources. Especially for recently introduced and/or detected compounds (so called emerging micropollutants), there is a growing need to investigate the occurrence and fate in WWTPs. In order to get a comprehensive picture on the behavior in municipal wastewater treatment, the following groups of emerging organic micropollutants, spanning a broad range of applications and physico-chemical properties, were selected as target compounds: pharmaceuticals (beta blockers, psycho-active drugs), UV-filters, vulcanization accelerators (benzothiazoles), biocides (anti-dandruffs, preservatives, disinfectants) and pesticides (phenylurea and triazine herbicides).