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Tree Growth and Tree Regeneration in Two East African Rain Forests as related to the Abiotic Environment after Human Disturbance Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften (Dr. rer. nat.) Fakultät Naturwissenschaften Universität Hohenheim Institut für Botanik (210) vorgelegt von Robert Gliniars aus Wallasey (England) 2010 Dekan: Prof. Dr. H. Breer 1. Prüfer (Betreuer): Prof. Dr. M. Küppers 2. Prüfer (Mitberichter): Prof. Dr. A. Fangmeier 3. Prüfer: Prof. Dr. J. Steidle Eingereicht am: 06.September 2010 Mündliche Prüfung am: 17.Dezember 2010 Die vorliegende Arbeit wurde am 02.Dezember 2010 von der Fakultät Naturwissenschaften der Universität Hohenheim als „Dissertation zur Erlangung des Doktorgrades der Naturwissenschaften“ angenommen. Diese Dissertation ist auch in digitaler Form unter folgender Adresse verfügbar: www://opus.ub.uni-hohenheim.de Table of Contents Table of content s i Acronyms v Summary 1 Zusammenfassung 3 1 Introduction 5 1.1 Characteristics of tropical rain forests 6 1.2 Tree succession and disturbance 7 1.2.1 Seedling regeneration 9 1.2.2 Stem growth 12 1.3 Aim of this study 14 2 Study Sites 17 2.1 Kakamega Forest in Kenya 18 2.1.1 Geography 18 2.1.2 Geology and Soils 20 2.1.3 Climate 20 2.1.4 Vegetation 22 2.1.5 Human impact 22 2.2 Budongo Forest of Uganda 23 2.2.1 Geography 23 2.2.2 Geology and Soils 24 2.2.3 Climate 25 2.2.4 Vegetation 26 2.2.5 Human Impact 26 2.3 Main differences between the two forests 27 3 Material and Methods 29 3.1 Study plots 29 3.1.1 Disturbance classification 32 3.1.2 Recording of forest stand structure and tree diversity 33 3.1.2.1 Hemispherical photography 34 3.1.2.2 Canopy parameters and light measurements 35 3.1.2.3 Species identification and collected material 35 3.1.3 Additional data on the abiotic environment 36 3.2 Stem growth measurements 38 3.2.1 Dendrometer bands 39 3.2.2 Selected species 40 3.2.3 Tree stem growth measurement 41 3.2.4 Stem growth rates 42 3.2.5 Tree phenology surveys 43 3.2.6 Tree age calculation 43 3.3 Seedling surveys 44 3.4 Data handling and statistics 44 i Table of Contents 3.4.1 General Statistics 45 3.4.2 Estimates of the actual species richness 45 3.4.3 α-diversity 46 3.4.4 β-diversity 46 4 Results: Kakamega Forest 47 4.1 Tree diversity 47 4.1.1 α-diversity 47 4.1.1.1 α-diversity as related to environmental factors 48 4.1.2 Effect of disturbance on β-diversity 49 4.2 Tree dynamics 50 4.2.1 Stem growth of all study trees 50 4.2.2 Stem growth of selected tree species 51 4.2.3 Temporal constancy of stem growth 52 4.2.4 Stem growth as related to phenological dynamics 53 4.2.5 Tree turnover 55 4.2.5.1 Tree age 55 4.2.5.2 Self-thinning and tree mortality 56 4.2.5.3 Balance of tree turnover 56 4.2.6 Stem growth as related to structural tree parameters 58 4.2.6.1 Between species stem growth by DBH categories 58 4.2.6.2 Between species stem growth by height categories 61 4.3 Forest structure as related to human disturbance 64 4.3.1 Effect of plot size 66 4.3.2 Canopy parameters 66 4.4 Effects of climatic factors on tree dynamics 68 4.4.1 Effect of incident precipitation and canopy throughfall on stem growth 68 4.4.2 Effect of temperature on stem growth 71 4.4.3 Effect of climatic factors on tree phenology 74 4.4.4 Effect of soil parameters on tree growth 74 4.4.4.1 Plot scale 74 4.4.4.2 Tree individual scale 76 4.5 Tree seedling establishment 77 4.5.1 Tree seedling α-diversity 77 4.5.1.1 Tree seedling diversity as related to abiotic factors 78 4.5.2 Dynamics of seedlings in space and time 79 4.5.2.1 Seedling recruitment 79 4.5.2.2 Dynamics of seedlings as related to canopy throughfall 82 4.5.2.3 Seedling dynamics and incident precipitation 83 4.5.2.4 Distribution of tree seedlings as related to soil parameters 85 4.5.2.5 Survival of tree seedlings from initial census 85 5 Results: Budongo Forest 87 5.1 Tree diversity 87 5.1.1 α-diversity 87 5.1.1.1 α-diversity as related to environmental factors 88 5.1.2 β-diversity 88 ii Table of Contents 5.2 Tree dynamics 89 5.2.1 Stem growth of all study trees 89 5.2.2 Stem growth of selected tree species 90 5.2.3 Temporal constancy in stem growth 91 5.2.4 Stem growth as related to phenological dynamics 92 5.2.5 Tree turnover 94 5.2.5.1 Tree age 94 5.2.5.2 Self-thinning and tree mortality 94 5.2.5.3 Balance of tree turnover 95 5.2.6 Stem growth as related to structural parameters 96 5.2.6.1 Between species growth by DBH categories 96 5.2.6.2 Between species growth by height categories 99 5.3 Forest structure as related to human disturbance 100 5.3.1 Canopy parameters 101 5.4 Effect of climatic factors on tree dynamics 103 5.4.1 Effect of incident precipitation on stem growth 103 5.4.2 Effect of temperature on stem growth 104 5.4.3 Effect of climatic factors on tree phenology 108 5.5 Tree seedling establishment 109 5.5.1 Tree seedling α-diversity 109 5.5.1.1 Seedling diversity as related to abiotic factors 109 5.5.2 Dynamics of seedlings in space and time 111 5.5.2.1 Seedling recruitment 111 5.5.2.2 Dynamics of seedlings as related to climatic factors 112 5.5.2.3 Distribution of seedlings as related to the abiotic environment 115 5.5.2.4 Survival of tree seedlings from initial census 115 6 Forest Comparison 117 6.1 Tree diversity and forest structure 117 6.2 Stem growth 118 6.2.1 Tree growth as related to abiotic factors 118 6.2.2 Species growth comparison 119 6.3 Seedling density as related to abiotic factors 120 7 Discussion 121 7.1 Influence of human disturbance on the two forests 121 7.1.1 Tree diversity and growth on forest scale 121 7.1.2 Different successional stages 124 7.2 Tree succession in disturbed forest sites 130 7.2.1 The pioneer-climax continuum 130 7.2.2 Role of the temporal variability of climatic factors 133 7.2.3 Role of the spatial variability of soil parameters 136 7.3 Conclusion 139 8 Acknowledgements 141 9 References 143 iii Table of Contents List of Figures 159 List of Tables 165 10 Appendix 171 10.1 Kakamega Forest 171 10.2 Budongo Forest 178 Erklärungen 187 Curriculum Vitae 191 iv Acronyms Acronyms % circ: Relative growth abs.: Absolute growth ACE: Species estimator acetate: Ammonium acetate extracted acid: Nitric acid extracted a.s.l.: Above sea level B: Biso (part of Budongo Forest) BA: Basal Area BCFS: Budongo Conservation Field Station BCI: Barro Colorado Island Forest BDO: Biodiversity Observatory BF: Budongo Forest BIOTA: Biodiversity Monitoring Transect Analysis BU: Buyangu Hill (part of Kakamega Forest) C: Carbon Ca: Calcium CA: Campforest (part of Kakamega Forest) CDI: Commercial Disturbance Index CE: Crown exposure CEC: Cation Exchange Capacity Chao 1: Species estimator Chao 2: Species estimator circ.: Circumferential Cmol: Coulomb per mole CO: Colobus (part of Kakamega Forest) DBH: Diameter at Breast Height DBH max: Maximum DBH DBH min: Minimum DBH EA: East African herbarium in Nairboi, Kenya EC: Electrical Conductivity e.g.: exempli gratia EM: Ectomycorrhiza FD: Forest Department, Kenya Fisher’s alpha: Diversity Index FTEA: Flora of Tropical East Africa ha: Hectare Height max: Maximum height Height min: Minimum height v Acronyms HOH: Herbarium of the University of Hohenheim IDH: Intermediate disturbance hypothesis IM: Intermediate indiv.: Individuals IPav: Monthly Incident Precipitation IP2: Summed two-monthly Incident Precipitation IP3: Summed three-monthly Incident Precipitation IK: Ikhuywa (part of Kakamega Forest) IS: Isecheno (part of Kakamega Forest) IT: Shade-Intolerant ITCZ: Inter Tropical Convergence Zone Jackknife 1: species estimator Jackknife 2: species estimator K: Potassium KA: Kaimosi (part of Kakamega Forest) KI: Kisere (part of Kakamega Forest) km2 : Square kilometer KF: Kakamega Forest KP11: Kaniyo-Pabidi 11th compartment (part of Budongo Forest) KWS: Kenyan Wildlife Service LAI: Leaf Area Index LDI: Local Disturbance Index MA: Malava (part of Kakamega Forest) MANOVA: Analysis of variance MFNP: Murchison Falls National Park Mg: Magnesium MH: Maximum height MHU: Herbarium of the Makerere University, Uganda Mn: Manganese mNN: Meter über Normalnull N: Nitrogen N3: Nyakafunjo 3rd compartment (part of Budongo Forest) N4: Nyakafunjo 4th compartment (part of Budongo Forest) N15: Nyakafunjo 15th compartment (part of Budongo Forest) n.d.: Not determined NFA: National Forestry Authority, Uganda NESS: Normalized Expected Species Shared NMDS: Non-metrical dimensional scaling No.: Number ODI: Overall Disturbance Index vi Acronyms p: Probability P: Phosphorus PG: Physiological Group pH: Power of hydrogen precip.: Precipitation RD: Number of rainy days RDM: Repeated diameter measurements rel.: Relative SAI: Salazar I (part of Kakamega Forest) SAII: Salazar II (part of Kakamega Forest) SD: Standard Deviation Shannon H’: Shannon-Wiener-H’ Diversity Index Simpson: Simpson Diversity Index ST: Shade-Tolerant T: Temperature TFav: Sum of monthly throughfall TF2: Sum of two-monthly throughfall TF3: Sum of three-monthly throughfall Ti: Titanium UWA: Ugandan Wildlife Authority VAM: Vesicular-arbuscular mycorrhiza W21: Waibira 21st compartment (part of Budongo Forest) W22: Waibira 22nd compartment (part of Budongo Forest) W36: Waibira 36th compartment (part of Budongo Forest) water: Water extracted YA: Yala (part of Kakamega Forest) yr: Year vii Summary Summary This study deals with the stem growth and seedling regeneration of different native tree species in two East African rainforests influenced by human disturbance in Kenya (Kakamega Forest) and Uganda (Budongo Forest), also considering spatially and temporally variable environmental influences.
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    CHANGING FOREST-WOODLAND-SAVANNA MOSAICS IN UGANDA: WITH IMPLICATIONS FOR CONSERVATION Grace Nangendo Promoters: Prof. Dr. F.J.J.M. Bongers Personal Professorship at Forest Ecology and Forest Management Group, Wageningen University, The Netherlands Prof. Dr. Ir. A. De Gier Professor, Geo-information for Forestry / Department of Natural Resources International Institute for Geo-information Science and Earth Observation (ITC), Enschede, The Netherlands Co-promoter: Dr. H. ter Steege Chair Plant Systematics (Ag.), Nationaal Herbarium Nederland - Utrecht Branch Utrecht University, The Netherlands Examining Committee: Dr. J. F. Duivenvoorden, University of Amsterdam, The Netherlands Prof. Dr. M. J. A. Werger, Utrecht University, The Netherlands Dr. J. R. W. Aluma, National Agricultural Research Organization (NARO), Uganda Prof. Dr. M. S. M. Sosef, Wageningen University, The Netherlands CHANGING FOREST-WOODLAND-SAVANNA MOSAICS IN UGANDA: WITH IMPLICATIONS FOR CONSERVATION Grace Nangendo Thesis To fulfil the requirements for the degree of Doctor on the authority of the Rector Magnificus of Wageningen University, Prof. Dr. Ir. L. Speelman, to be publicly defended on Wednesday, June 1, 2005 at 15:00 hrs in the auditorium at ITC, Enschede, The Netherlands. ISBN: 90-8504-200-3 ITC Dissertation Number: 123 International Institute for Geo-information Science & Earth Observation, Enschede, The Netherlands © 2005 Grace Nangendo CONTENTS Abstract .......................................................................................................................