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Vegetation Ecology of Forest-Savanna Ecotones in the Comoé National Park (Ivory Coast): Border and Ecotone Detection, Core-Area Analysis, and Ecotone Dynamics

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Vegetation Ecology of Forest-Savanna Ecotones in the Comoé National Park (Ivory Coast): Border and Ecotone Detection, Core-Area Analysis, and Ecotone Dynamics Vegetation ecology of forest-savanna ecotones in the Comoé National Park (Ivory Coast): Border and ecotone detection, core-area analysis, and ecotone dynamics Dissertation zur Erlangung des akademischen Grades Doctor rerum naturalium (Dr. rer. nat.) der Mathematischen-Naturwissenschaftlichen Fakultät der Universität Rostock Institut für Biowissenschaften (IfBI) Abteilung Allgemeine und Spezielle Botanik vorgelegt von Klaus Josef Hennenberg, geb. am 18.12.1970 in Münster (Westf.) aus Rostock Rostock, 17.12.2004 Gutachter: Prof. Dr. Stefan Porembski Prof. Dr. Wilhelm Barthlott Prof. Dr. Florian Jeltsch Tag der Verteidigung: 23.05.2005 Contents Contents ......................................................................................................................................3 List of Figures.............................................................................................................................5 List of Tables ..............................................................................................................................7 List of Abbreviations ..................................................................................................................8 Summary...................................................................................................................................10 Zusammenfassung ....................................................................................................................12 1 Introduction.......................................................................................................................14 1.1 Transect analysis.......................................................................................................17 1.1.1 Transect analysis of univariate data..................................................................17 1.1.2 Transect analysis of multivariate data ..............................................................18 1.2 Biomass and surface fires along forest-savanna ecotones........................................18 1.3 Microclimate along forest-savanna ecotones............................................................19 1.4 Vegetation composition along forest-savanna ecotones...........................................20 1.5 Core-area analysis.....................................................................................................21 1.6 Dynamics of forest-savanna ecotones ......................................................................22 2 Material and Methods.......................................................................................................24 2.1 Study area.................................................................................................................24 2.1.1 Climate..............................................................................................................25 2.1.2 Geology and soils .............................................................................................26 2.1.3 Vegetation.........................................................................................................27 2.1.4 Characteristics of Anogeissus leiocarpus .........................................................29 2.2 Sampling design........................................................................................................29 2.3 Sampling of vegetation .............................................................................................31 2.3.1 Vegetation structure and composition ..............................................................31 2.3.2 Sampling of tree and shrub individuals ............................................................32 2.3.3 Density measurement of tree and shrub individuals.........................................33 2.3.4 Sampling of surface biomass ............................................................................33 2.4 Sampling of abiotic parameters ................................................................................33 2.4.1 Microclimatic parameters .................................................................................33 2.4.2 Fire occurrence, shading, and soil depth ..........................................................34 Contents 2.5 Border-and-ecotone-detection analysis (BEDA) for univariate transect data ..........35 2.6 Statistical analysis.....................................................................................................37 2.6.1 General transect description.............................................................................37 2.6.2 Surface biomass and surface-fire probability ...................................................37 2.6.3 Microclimate.....................................................................................................38 2.6.4 Vegetation composition....................................................................................38 2.6.5 Core-area analysis.............................................................................................39 2.6.6 Size-class distribution of tree species...............................................................40 3 Results...............................................................................................................................42 3.1 General transect description.....................................................................................42 3.2 Surface biomass and surface-fire probability ...........................................................45 3.3 Seasonal variability in microclimatic borders and ecotones.....................................50 3.4 Border and ecotone detection by means of vegetation composition ........................56 3.5 Core-area analysis.....................................................................................................59 3.6 Size-class distribution of tree species with a focus on Anogeissus leiocarpus.........62 4 Discussion.........................................................................................................................69 4.1 Border and ecotone detection ...................................................................................69 4.1.1 Border-and-ecotone detection analysis (BEDA) for univariate transect data...69 4.1.2 SMWDA and MWRA for multivariate transect data .......................................70 4.2 Relation of surface biomass and surface fire............................................................71 4.3 Seasonal variability in microclimatic borders and ecotones.....................................74 4.4 Border and ecotone detection by means of vegetation composition ........................76 4.5 Core-area analysis.....................................................................................................77 4.6 Dynamics of forest-savanna ecotones and the role of Anogeissus leiocarpus .........80 5 Conclusions.......................................................................................................................84 References.................................................................................................................................87 Acknowledgements.................................................................................................................102 Appendix.................................................................................................................................104 4 List of Figures List of Figures Fig. 1.1. The hierarchical continuum model combined with the climax pattern hypothesis expressed as species response curves along a single environmental gradient..................20 Fig. 2.1. Map of the study area in the southwestern part of the Comoé National Park, Ivory Coast, West Africa. .................................................................................................25 Fig. 2.2. Mean monthly and mean annual rainfall for three climate stations in northeastern Ivory Coast and for the climate station at the research camp of the University of Würzburg in the southwestern part of the CNP..........................................26 Fig. 2.3. Idealized model of the vegetation distribution along a topographical gradient of a pediplain.........................................................................................................................28 Fig. 2.4. Distribution of dominant vegetation types in the northern and southern part of the CNP.............................................................................................................................29 Fig. 2.5. Sampling designs along forest-savanna transects being adapted to the study parameters.........................................................................................................................31 Fig. 2.6. Scheme of the border-and-ecotone detection analysis (BEDA).................................35 Fig. 2.7. Demonstration of the method to value the width of an ecotone at a significant border detected by the split-moving-window dissimilarity analysis (SMWDA) using moving-window regression analysis (MWRA) ................................................................39 Fig. 3.1. Values of three structural parameters and soil depth along eight studied transects...43 Fig. 3.2. Sample scores of the first and second DCA axes for the cover of all species along the intensely studied transect ..................................................................................44 Fig. 3.3. Box-plot of dry mass of grasses, herbs, and litter for
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