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- Beta-Diversität (1)
- Habitat Fragmentation (1)
- Human Disturbance (1)
- Kakamega Forest (1)
- Kenya (1)
- Plant Communities (1)
- Vegetation (1)
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.
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.
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.