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Spatial and Matrix Influences on the Biogeography of Insect Taxa in Forest Fragments in Central Uganda

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Spatial and Matrix Influences on the Biogeography of Insect Taxa in Forest Fragments in Central Uganda Spatial and matrix influences on the biogeography of insect taxa in forest fragments in central Uganda Perpetra Akite Dissertation for a cotutelle award of Doctor of Philosophy Degree of Makerere University, Uganda and University of Bergen, Norway Makerere University University of Bergen 2016 Department of Biological Sciences, Makerere University Department of Biology, University of Bergen ii DECLARATION OF ORIGINALITY This is my own work and it has never been submitted for any degree award in any University iii TABLE OF CONTENTS DECLARATION OF ORIGINALITY......................................................................................iii LIST OF CONTENTS...............................................................................................................iv ACKNOWLEDGEMENTS.......................................................................................................vi LIST OF PAPERS....................................................................................................................vii Declaration of authors’ contributions…………………….…...……………...……...viii ABSTRACT...............................................................................................................................x BACKGROUND........................................................................................................................1 Problem statement..........................................................................................................……….2 Objectives........................................................................................................................3 Research questions..........................................................................................................4 INTRODUCTION......................................................................................................................5 Forest fragmentation and Biodiversity............................................................................6 Does the Matrix matter?……………………………………………………………......9 Roles of insects in ecosystem dynamics……………………………………..……… 12 How much is known about insects in Uganda?………………………………………14 STUDY AREAS AND SITES.................................................................................................15 MATERIALS AND METHODS…………………………………………………….………17 Study protocols…………………………………………………………………………….…17 Paper I…………………………………………………..……………………….……17 Paper II…………………………………………………………………………..……19 Paper III……………………………………………………………………………….21 Paper IV…………………………………………………………………………….…23 RESULTS AND DISCUSSION...............................................................................................24 Temporal dynamics of forest biodiversity....................................................................26 iv Species distribution along the vertical continuum........................................................29 Habitat use by insects in a patch-matrix landscape.......................................................30 The need for focused sampling in face of limited inventory………………….………33 Challenges of biodiversity conservation in modified forest landscapes in Uganda…. 34 The social and political realities and their effects on forest protection in Uganda…...36 CONCLUSIONS......................................................................................................................39 RECOMMENDATIONS.........................................................................................................40 REFERENCES..........................................................................................................…….…..42 PLATES SHOWING REPRESENTATIVE SPECIES OF DIFFERENT GROUPS INDIVIDUAL PAPERS I – IV APPENDICES OF LISTS OF SPECIES OF DIFFERENT INSECT TAXA v ACKNOWLEDGEMENTS This study was supported by the Norwegian Research Council (FRIMUF programme) funded the study through the MATRIX project (# 184912) and supported by the University of Bergen–Makerere University Collaboration. I am grateful to my supervisors for guiding me through my doctoral study. You have been inspiring and supportive to my research decisions. Your invaluable expertises in the different aspects of this study are highly regarded. I am indebted to Vigdis Vandvik PI-MATRIX project) for her enthusiastic, encouraging and valuable discussions throughout this work. Your fast feedback on manuscripts, open minded face to face discussions and very constructive criticisms were instrumental in making this study end a reality. Amy Eycott, it’s said better late than never and your coming on board the Matrix project was an honour for me. Thank you for your tireless efforts, endless reading of manuscripts, mapping and all other help you extended to me. I am especially grateful to Paul Waring and Hugh Rowell for all the expert field trainings, Ian Kitching and Alessandro Giusti from the Natural History Museum London, Torben Larsen (R.I.P) for identifying difficult butterfly specimens, research associates Therese Kronstad and Jenny Reiniö and our local assistants, colleagues at Ecology and Environmental Research Group (University of Bergen) and the Department of Biological Sciences, Makerere University for a friendly and inspiring work atmosphere, Cathy Jenks; my welfare hero in Bergen, Josephine Esaete for all the ‘big sister’ counsel and fantastic coordination, Ulf Bjelke for financial help following gruesome road accident and for sharing my passion for insects and photography, Janet McCrae, Joseph Chipperfield, Mikail Erdogan and Jan Skogvang. Above all, I am grateful to God for holding everything together to the very end. This thesis is dedicated to a dear friend, Petter Hansen (the cruel hand of death did not let you see the end of this journey), and to all who appreciate nature. vi LIST OF PAPERS The thesis is based on the following four papers which will be referred to by their Roman numerals hereafter. Paper I: Akite, P., Telford, R.J., Waring, P., Akol, A.M. & Vandvik, V. (2015) Temporal patterns in Saturnidae (silk moth) and Sphingidae (hawk moth) assemblages in protected forests of Central Uganda. Ecology and Evolution 5 (8): 1746–1757. Paper II: Akite, P., Akol, M.A., Kronstad, T., Vandvik, V & Telford, R.J. Vertical distribution of fruit-feeding butterflies in three protected forests in Central Uganda. Manuscript under revision. Paper III: Akite, P., Akol, M.A., Eycott, A.E., Vandvik, V. & Telford, R.J. The use of matrix habitats by forest insect species. Manuscript. Paper IV: Akite, P. & Rowell, C.H.F. (2013) Oshwea dubiosa rediscovered in Uganda. Journal of Orthoptera Research 22 (1): 45–49. vii Declaration of authors’ contributions Paper I: P. Akite, R.J. Telford, P.Waring, A.M. Akol & V. Vandvik. Temporal patterns in Saturnidae (silk moth) and Sphingidae (hawk moth) assemblages in protected forests of Central Uganda. Akite, P.: Compiling historical data, survey design, field work, laboratory work, data processing, statistical analyses, writing Telford, R.J.: Project design, analytical design, data processing, statistical analyses, editing Waring, P.: Survey design, field work, editing Akol, A.M.: Analytical design, statistical analyses, editing Vandvik, V.: Project design, analytical design, statistical analyses, editing Paper II: Akite, P., Akol, M.A., Kronstad, T., Vandvik, V & Telford, R.J. Vertical distribution of fruit-feeding butterflies in three protected forests in Central Uganda. Akite, P.: Survey design, field work, laboratory work, data processing, statistical analyses, writing Akol, M.A.: Analytical design, editing Kronstad, T.: Field work, laboratory work, data processing, editing Vandvik, V.: Project design, analytical design, statistical analyses, editing Telford, R.J.: Project design, analytical design, data processing, statistical analyses, editing viii Paper III: Akite, P., Akol, M.A., Eycott, A.E., Vandvik, V. & Telford, R.J. The use of matrix habitats by forest insect species. Akite, P.: Survey design, field work, laboratory work, data processing, statistical analyses, writing Akol, M.A.: Editing Eycott, A.E.: Data processing, statistical analyses, editing Vandvik, V.: Project design, analytical design, statistical analyses, editing Telford, R.J.: Project design, analytical design, statistical analyses, editing Paper IV: Perpetra Akite & C.H.F. Rowell (2013). Oshwea dubiosa rediscovered in Uganda. Akite, P.: Field work, data processing, editing Rowell, C.H.F.: Field work, laboratory work, data processing, writing ix ABSTRACT How best to manage forest patches, mitigate the consequences of forest fragmentation, and enable landscape permeability are key questions facing conservation scientists and managers. In Uganda, most protected forests have undergone considerable changes from a range of human activities in recent decades. As such once continuous forests now exist as ‘islands’ (patches) in a matrix of non-forest habitats. In fragmented forest landscapes, the capacity of matrix habitats to support forest species varies. However, very little is known of how faunal communities and in particular insect assemblages utilize resources within the patch-matrix landscape. This study explored spatial and temporal patterns of diversity and distribution of butterflies, moths and grasshoppers in the patch-matrix landscapes of three protected forests: Mabira (largely stable and recovering from past encroachment, albeit with minor illegal logging, but disturbed at the edges), Zika and Mpanga (relatively undisturbed and unchanged internally, but with substantially altered matrix),
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