EFFECTS OF LANDUSE CHANGE AND FOREST FRAGMENTATION ON THE BIODIVERSITY AND ECOSYSTEM FUNCTIONING IN THE TROPICAL LOWLANDS OF SRI LANKA ENOKA PRIYADARSHANI KUDAVIDANAGE B.Sc (Zoology), M.Sc (Environmental Sciences) A THESIS SUBMITTED FOR THE DEGREE OF DOCTOR OF PHILOSOPHY DEPARTMENT OF BIOLOGICAL SCIENCES NATIONAL UNIVERSITY OF SINGAPORE 2011 “The forest is a peculiar organism of unlimited kindness and benevolence that makes no demands for its sustenance and extends generously the products of its life activity; it provides protection to all beings, offering shade even to the man who destroys it.” — Gautama Buddha i Dedicated to my parents ii ACKNOWLEDGEMENTS I am grateful to late Prof Navjot Sodhi, my ex-supervisor whose unique combination of intelligence, determination, intimidation and social skills defined my path into the field of conservation biology I am indebted, to my supervisor Prof Edward Webb for persuasion and guidance that enabled me to complete the PhD thesis, and to the constant encouragement provided by Prof Richard Corlett I extend my gratitude to Prof Sarath Kotagama, my supervisor in Sri Lanka, and mentor for the last 15 years, for his guidance provided throughout my academic life I thank all examiners for their most valuable and helpful comments that were adopted in the final version of my thesis Many thanks to all the past and present staff member and colleagues of the Conservation Ecology, Applied Plant Ecology laboratories and the terrestrial ecology group including Daniel Friess; David Bickford; Mary Rose Posa, Nanthinee Jeevanadam, Reuben Clement, Arvin Diesmos and Lee Wei Kit; the technical and administrative staff including Tommy Tan, Reena Devi Samayanadan, Priscilla Lee, members of the Biodiversity group and the Raffles Museum for Biodiversity research of the Department of Biological Sciences for all the encouragement assistance and inspiration extended in one-way or another Special thanks towards Janice Lee and Qie Lan, my fellow dung girls for the comradeship of dung beetles Prof David J Lohman (Lohman Lab, City College of New York) is kindly acknowledged for extending a generous helping hand in many challenges faced during my PhD study including correcting manuscripts, assisting analytical problems, helpful discussions and providing resources I’m grateful to Darren J Mann, Eleanor Slade and the staff of the Oxford University Museum of Natural History for patiently guiding me through the intense process of species identification and for the opportunity to use the scarab beetle collection at the American Museum iii of Natural History Thanks to the ScarabNet research team including Finbarr Horgan for sharing with me their knowledge about dung beetles I acknowledge all my collaborators; Thomas Wanger (Center for Conservation Biology , Stanford University), Neil Collier (Lohman Lab, City College of New York) for analytical work; Priyadarshan Dharmarajan (Ashoka Trust, India), Manori Gunetilleke (National Museums, Sri Lanka) and Deepchandi Lekamge (Sabaragamuwa University) for the taxonomic study I acknowledge the Department of Wildlife Conservation and the Forest Department, Sri Lanka for granting me access to work in the protected area and permission to export specimen for taxonomic work I thank the Vice- Chancellor of the Sabaragamuwa University of Sri Lanka and the Department of Natural Resources for kindly facilitating my graduate studies Gratitude extended to Professors Nimal Gunatilleke, Savithri Gunatilleke and Drs Eben and Uromi Goodale for expertise knowledge, guidance and helpful discussions Ravi Amarasinghe, Chamitha de Alwis, Vimukthi Herath, Krishan and Udeni Ariyasiri, Vishan, Amila Perera, Nuwan Hegodaaarachchi, the Wildlife Conservation Society of Galle and the local assistants who supported the field work are acknowledged The Field research was funded by NAGAO Natural Environmental Foundation, Japan and Joe Grove memorial award, UK, while the taxonomic study was funded by the Oxford University Museum of Natural History, London The World Bank sponsored IRQUE project and the National University of Singapore Research Scholarship supported the PhD studies Finally, I thank everybody including my family who stood by me extending their constant support, patience and understanding through the graduate studies I owe all that I am to my parents, who made it their lifetimes’ goal to help me fulfill my dreams I remember with heartfelt gratitude and warmth, my father who passed away one month before I entered the graduate iv school of his choice and my mother who stood by me all my life before she passed away seven months ago She shared my love for nature and all living things, supported my chosen career and enjoyed every one of my field expeditions, participating with great enthusiasm This thesis is a product of her faith in me v Contents ACKNOWLEDGEMENTS iii THESIS SUMMARY LIST OF FIGURES LIST OF TABLES APPENDICES CHAPTER 1: GENERAL INTRODUCTION CHAPTER 2: STUDY SITE 13 2.1 Sri Lanka and the Lowland Wet Zone: a biodiversity hotspot 13 2.2 From game protection to biodiversity conservation: the history and status of biodiversity conservation in the wet zone of Sri Lanka 15 2.3 Study sites: Sinharaja, Kanneliya and Kottawa 20 2.3.1 Sinharaja Man and Biosphere Reserve 20 2.3.2 Kanneliya conservation forest 23 2.3.3 Kottawa Kombala conservation forest 24 CHAPTER 3: AMPHIBIAN AND BUTTERFLY DIVERSITY ACROSS A LOWLAND TROPICAL LAND-USE GRADIENT IN SRI LANKA 28 3.1 Introduction 28 3.2 Materials and methods 29 3.2.1 Study area 29 vi 3.2.2 Sampling diversity and abundance 30 3.2.3 Environmental variables 31 3.2.4 Analysis of species richness and abundance data 32 3.2.5 Analysis of environmental variables 33 3.3 Results 34 3.3.1 Butterflies 34 3.3.2 Amphibians 35 3.3.3 Environmental determinants 37 3.4 Discussion 38 3.4.1 Butterfly response to land-use change 38 3.4.2 Amphibian response to land-use change 40 3.4.3 Environmental predictors of butterfly and amphibian diversity patterns 42 3.5 Conclusions 43 4.1 Introduction 51 4.2 Dung beetles (Insecta: Coleoptera: Polyphaga: Scarabaeoidea: Scarabaeidae: Scarabaeinae and Aphodiinae) 51 4.2 Materials and methods 53 4.2.1 Study site 53 4.2.2 Beetle sampling and identification 54 4.3 Results and discussion 55 4.3.1 Species distribution overview 55 4.3.2 Species distribution in contrast to previous records 56 vii 4.3.4 Possible natural and anthropogenic environmental determinants of observed distribution patterns of dung beetle species 58 4.3.5 Limitations and future work 62 CHAPTER 5: EFFECTS OF LAND-USE CHANGE ON THE COMMUNITIES OF DUNG BEETLES IN LOWLAND TROPICS OF SRI LANKA 67 5.1: Introduction 67 5.4 Materials and methods 70 5.4.3 Beetle sampling 71 5.5 Data analysis 72 5.8 Conclusion 84 CHAPTER DUNG BEETLE COMMUNITIES IN LOWLAND FOREST FRAGMENTS 90 6.1 Introduction 90 6.2 Study site 94 6.3 Materials and methods 94 6.3.1 Dung beetle sampling 94 6.3.2 Environmental variables and fragment characteristics 95 6.3.3 Fragment characteristics 95 6.4 Data analysis 96 6.4.1 Summary statistics and community parameters 96 6.4.2 Comparison of species richness, abundance across the fragments 96 6.5 Results 99 6.5.1 Community comparison 100 viii 6.5.2 Relating species richness, abundance, fragment characteristics and environmental variables 101 6.6 Discussion 104 6.7 Conclusion 111 CHAPTER HABITAT CHANGE IMPAIRS ECOLOGICAL SERVICES PROVIDED BY SCARABAEINAE DUNG BEETLES IN THE TROPICAL LOWLANDS OF SRI LANKA 122 7.1 Introduction 122 7.2 Materials and methods 123 7.2.1 Study area 123 7.2.2 Sampling design for dung removal and dung beetle trapping 123 7.2.3 Dung removal experiments 124 7.2.4 Beetle sampling 125 7.2.5 Statistical analysis 125 7.3 Results 126 7.4 Discussion 127 7.4.1 Functional responses of dung beetles 127 7.4.2 Economic implications for conservation 129 7.5 Conclusions 131 CHAPTER 8: GENERAL DISCUSSION – RESEARCH FINDINGS AND THEIR APPLICABILITY 137 8.1 Ecological trends 137 8.2 Conservation implications 143 BIBLIOGRAPHY 150 ix tardigradus in Masmullah proposed forest reserve, Sri Lanka Mammalia 69, 201210 Nichols, E., Gardner, T A., Peres, C A., and Spector S (2009) Co-declining mammals and dung beetles: an impending ecological cascade Oikos 118, 481-487 Nichols, E., Gardner, T.A., Peres, C.A., and Spector, S (2009) Co-declining mammals and dung beetles: an impending ecological cascade Oikos 118, 481-487 Nichols, E., Larsen, T., Spector, S., Davis, A.L., Escobar, F., Favila, M., and Vuline, K (2007) Global dung beetle response to tropical forest modification and fragmentation: A quantitative literature review and meta-analysis Biol Conserv 137, 1-19 Nichols, E., Spector, S., Louzada, J., Larsen, T., Amequita, S., and Favila, M.E (2008) Ecological functions and ecosystem services provided by Scarabaeinae dung beetles Biol Conserv 141, 1461-1474 Nichols, E.S., and Gardner, T.A (2011) Dung beetles as a candidate study taxon in applied biodiversity conservation research In Ecology and Evolution of Dung Beetles, L.W Simmons, and T.J Ridsdill-Smith, eds (John Wiley and Sons), pp 267-290 Novacek, M.J., and Cleland, E.E (2001) The current biodiversity extinction event: Scenarios for mitigation and recovery Proceedings of the National Academy of Sciences of the United States of America 98, 5466-5470 Nummelin, M., and Hanski, I (1989) Dung Beetles of the Kibale Forest, Uganda Comparison between Virgin and Managed Forests Journal of Tropical Ecology 5, 349-352 172 Nyeko, P (2009) Dung Beetle Assemblages and Seasonality in Primary Forest and Forest Fragments on Agricultural Landscapes in Budongo, Uganda Biotropica 41, 476-484 Osberg, D.C., Doube, B.M., and Hanrahan, S.A (1994) Habitat specificity in African dung beetles: The effect of soil type on the survival of dung beetle immatures (Coleoptera Scarabaeidae) Tropical Zoology 7, 1-10 Peeris, C.V.S (1975) The ecology of the endemic tree species of Sri Lanka in relation to their conservation PhD thesis (University of Aberdeen, UK) Pemadasa, M.A., and Gunatilleke, C.V.S (1981) Pattern in a rain-forest in Sri-Lanka J Ecol 69, 117-124 Peres, C.A (2001) Synergistic effects of subsistence hunting and habitat fragmentation on Amazonian forest vertebrates Conservation Biology 15, 1490-1505 Pethiyagoda, R and Manamendra-Arachchi, K (1998) Evaluating Sri Lanka’s amphibian diversity Occasional Papers Wildlife Heritage Trust 2, 1–12 Pethiyagoda, R (2005) Exploring Sri Lanka’s biodiversity Raffles Bulletin of Zoology (Suppl.) 12, 1-4 Philips, T.K., Pretorius, E., and Scholtz, C.H (2004) A phylogenetic analysis of dung beetles (Scarabaeinae: Scarabaeidae): unrolling an evolutionary history Invertebrate Systematics 18, 53-88 Pineda, E., and Halffter, G (2005) Cloud forest fragmentation and frog diversity in a montane landscape in Mexico m3m Monografias Tercer Milenio 4, 165-176 Pollard, E and Yates, T.J (1993) Monitoring butterflies for ecology and conservation Chapman and Hall London Posa, M R C and Sodhi, N S (2006) Effects of anthropogenic land use on forest birds and butterflies in Subic Bay, Philippines Biol Cons 129, 256-270 173 Pulliam, H.R (1988) Sources, sinks, and population regulation American Naturalist 132, 652-669 Qie, L., T M Lee, N S Sodhi, and S L H Lim Dung beetle assemblages on tropical land-bridge islands: small island effect and vulnerable species J Biogeogr 38: 792-804 Quintero, I., and Roslin, T (2005) Rapid recovery of dung beetle communities following habitat fragmentation in central Amazonia Ecology 86, 3303-3311 R Development Core Team (2009) R: A language and environment for statistical computing R Foundation for Statistical Computing, Vienna, Austria ISBN 3900051-07-0, URL http://www.R-project.org R Development Core Team (2011) R: A language and environment for statistical computing R Foundation for Statistical Computing, Vienna, Austria ISBN 3900051-07-0, URL http://www.R-project.org Rahbek, C 1995 The elevation gradient of species richness: a uniform pattern? Ecography 18: 200-205 Raheem, D C., Naggs, F., Preece, R C., Mapatuna, Y., Kariyawasan, L and Eggleton P (2008) Structure and conservation of Sri Lankan land-snail assemblages in fragmented lowland rainforest and village home gardens J Appl Ecol 45, 10191028 Raheem, D.C., Naggs, F., Chimonides, P.D.J., Preece, R.C., and Eggleton, P (2009) Fragmentation and pre-existing species turnover determine land-snail assemblages of tropical rain forest Journal of Biogeography 36, 1923-1938 Ratcliffe, B.C and M.L Jameson 2000 Key to the families and subfamilies of Scarabaeoidea of the New World URL: http://www.museum.unl.edu/research/entomology/Guide/Scarabaeoidea/Scarabae 174 oideaK.htm In, B.C Ratcliffe and M.L Jameson (eds.), Generic Guide to New World Scarab Beetles (URL: http://wwwmuseum.unl.edu/research/entomology/Guide/index4.htm) Reid, J., and De Souza, W.C (2005) Infrastructure and conservation policy in Brazil Conservation Biology 19, 740-746 Ricketts, T (2001) Aligning conservation goals: are patterns of species richness and endemism concordant at regional scales? Animal Biodiversity and Conservation 24, 91-99 Ridsdill-Smith, T.J., and Edwards, P.B (2011) Biological control: ecosystem functions provided by dung beetles In Ecology and Evolution of Dung Beetles, L.W Simmons, and T.J Ridsdill-Smith, eds (John Wiley & Sons), pp 245-266 Rosenzweig, M.L (2003.) Reconciliation ecology and the future of species diversity Oryx 37, 194-205 Rosenzweig, M L 1995 Species Diversity in Space and Time Cambridge University Press, Cambridge, UK Roslin, T., and Viljanen, H (2011) Dung beetle populations: structure and consequences In Ecology and Evolution of Dung Beetles, L.W Simmons, and T.J RidsdillSmith, eds (John Wiley and Sons), pp 220-243 Rutledge, D 2003 Landscape indices as measures of the effects of fragmentation: can pattern reflect process? Doc science internal series 98 Department of Conservation, PO Box 10-420 Wellington, New Zealand Samways, M.J (1993) Insects in biodiversity conservation: some perspectives and directives Biodiversity and Conservation 2, 258-282 Saunders, D A and Hobb, R J., eds (1991) Nature conservation 2: The role of corridors Surrey Beatty and Sons, Chipping Norton, New South Wales 175 Saunders, D A., Hobbs, R J., and Margules, C R (1991) Biological consequences of ecosystem fragmentation: a review Conservation Biology, 5, 18–32 ScarabNet Global Taxon Database Version 1.5 [04-09-2011] www.scarabnet.org [Accessed 4th September 2011] Scheffler, P (2005) Dung beetle (Coleoptera: Scarabaeidae) diversity and community structure across three disturbance regimes in eastern Amazonia Journal of Tropical Ecology 21, 9-19 Schulze, C H., Linsenmaier, K E and Fiedler, K (2001) Understorey versus canopy: patterns of vertical stratification and diversity among Lepidoptera in a Bornean rain forest Plant Ecol 153, 133–152 Schulze, C H., Waltert, M., Kessler, P J A., Pitopang, R., Shahabuddin, Veddeler, D., Mühlenberg, M., Gradstein, S R., Leuschner, C., Steffan-Dewenter, I and Tscharntke, T (2004) Biodiversity indicator groups of tropical land-use systems: comparing plants birds and insects Ecol Appl 14, 1321–1333 Scott, D.M., Brown, D., Mahood, S., Denton, B., Silburn, A., and Rakotondraparany, F (2006) The impacts of forest clearance on lizard, small mammal and bird communities in the arid spiny forest, southern Madagascar Biological Conservation 127, 72-87 Sekercioglu, C.H (2002) Effects of forestry practices on vegetation structure and bird community of Kibale National Park, Uganda Biological Conservation 107, 229240 Sekercioglu, C H., P R Ehrlich, G C Daily, D Aygen, D Goehring, and R F Sandi (2002) Disappearance of insectivorous birds from tropical forest fragments Proceedings of the National Academy of Sciences of the United States of America 99, 263-267 176 Shahabuddin, N., Hidayat, P., Manuwoto, S., Noerdjito, W.A., Tscharntke, T., and Schulze, C.H (2010) Diversity and body size of dung beetles attracted to different dung types along a tropical land-use gradient in Sulawesi, Indonesia Journal of Tropical Ecology 26, 53-65 Shahabuddin, N., Schulze, C., and Tschnarntke, T (2005) Changes of dung beetle communities from rainforests towards agroforestry systems and annual cultures in Sulawesi (Indonesia) Biodiversity and Conservation 14, 863-877 Shimura, J (editor) 2003 Global Taxonomy Initiative in Asia – Research report from the National Institute for Environmental Studies, Japan 309 pp Singhakumara, B.M.P (1995) Ecological Assessment of the Kanneliya-DediyagalaNakiyadeniya (KDN) Forest Complex (Department of Forestry and Environmental Science, University of Sri Jayewardenepura, Gangodawila, Nugegoda) Slade, E M., Mann, D J., and Lewis O T (2010) Biodiversity and ecosystem function of tropical forest dung beetles under contrasting logging regimes Biological Conservation 144,166-174 Slade, E.M., Mann, D.J., Villanueva, J.F., and Lewis, O.T (2007) Experimental evidence for the effects of dung beetle functional group richness and composition on ecosystem function in a tropical forest Journal of Animal Ecology 76, 1094-1104 Smith, T.B., Mila, B., Grether, G.F., Slabbekoorn, H., Sepil, I., Buermann, W., Saatchi, S., and Pollinger, J.P (2008) Evolutionary consequences of human disturbance in a rainforest bird species from Central Africa Molecular Ecology 17, 58-71 Soberón, J M., Llorente, J B and Onate, L (2000) The use of specimen-label databases for conservation purposes: an example using Mexican Papilionid and Pierid butterflies Biodivers Conserv 9, 1441-1466 177 Sodhi, N S., Koh, L P., Clements, R., Wanger, T C., Hill, J K., Hamer, K C., Clough Y., Tscharntke, T., Posa, M R C., and Lee, T M (2010) Conserving Southeast Asian forest biodiversity in human-modified landscapes Biol Cons 143, 23752384 Sodhi, N S., and Brook, B W (2006) Southeast Asian biodiversity in crisis Cambridge University Press Cambridge Sodhi, N S., Brook, B W., and Bradshaw, C J A (2007) Tropical conservation biology Wiley-Blackwell, Boston, MA Sodhi, N S., Lee, T M., Koh, L P., and Brook B W (2009) A meta-analysis of the impact of anthropogenic forest disturbance on Southeast Asia's biotas Biotropica 41, 103-109 Sodhi, N., Liow, L.H., and Bazzaz, F.A (2004) Avian Extinctions from Tropical and Subtropical Forests Annu Rev Ecol Syst 35, 323-345 Sodhi, N.S., and Ehrlich, P R (Editors) (2010) Conservation Biology for All Oxford University Press, Oxford, UK 344 pp Sodhi, N.S., Koh, L.P., Clements, R., Wanger, T.C., Hill, J.K., Hamer, K.C., Clough, Y., Tscharntke, T., Posa, M.R.C., and Lee, T.M (2007) Conserving Southeast Asian forest biodiversity in human-modified landscapes Biol Conserv 143, 2375-2384 Spector, S (2002) Biogeographic crossroads as priority areas for biodiversity conservation Conserv Biol 16, 1480-1487 Spector, S (2006) Scarabaeine Dung Beetles (Coleoptera: Scarabaeidae: Scarabaeinae): An Invertebrate Focal Taxon for Biodiversity Research and Conservation Coleopterists Bulletin 5, 71-83 178 Spector, S., and Forsyth, A.B (1998) Indicator taxa in the vanishing tropics Pages 181– 210 in: Mace, G.M., and A Balmford, and J.R Ginsberg (editors) Conservation in a Changing World Cambridge University Press, London Spector, S., and Ayzama, S (2003) Rapid turnover and edge effects in dung beetle assemblages (Scarabaeidae) at a Bolivian Neotropical forest-savanna ecotone Biotropica 35, 394-404 Spitzer, K., Jaros, J., Havelka, J and Leps, J (1997) Effect of small-scale disturbance onbutterfly communities of an Indochinese montane rainforest Biol Cons 80, 9– 15 Steffan-Dewenter, I., Kessler, M., Barkmann, J., Bos, M.M., Buchori, D., Erasmi, S., Faust, H., Gerold, G., Glenk, K., Gradstein, S.R., et al (2007) Tradeoffs between income, biodiversity, and ecosystem functioning during tropical rainforest conversion and agroforestry intensification Proceedings of the National Academy of Sciences of the United States of America 104, 4973-4978 Stouffer, P.C., and Bierregaard, R.O (1995) Use of amazonian forest fragments by understory insectivorous birds Ecology 76, 2429-2445 Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L., and Waller, R.W (2004) Status and trends of amphibian declines and extinctions worldwide Science 306, 1783-1786 Sturtz, S., Ligges, U and Gelman, A (2005) R2WinBUGS: A Package for Running WinBUGS from R J Stat Softw 12, 1-16 Sundriyal, R.C., Rai, S.C., Sharma, E., and Rai, Y.K (1994) Hill agroforestry systems in South Sikkim, India Agroforestry Systems 26, 215-235 179 Tabachnik, B.G and Fidell, L.S (1989) Using Multivariate Statistics Second ed Harper and Row NY Ter Braak, C J F (1986) Canonical Correspondence Analysis: a new eigenvector technique for multivariate direct gradient analysis Ecology 67, 1167-1179 Terborgh, J., Lopez, L., Nunez, P., Rao, M., Shahabuddin, G., Orihuela, G., Riveros, M., Ascanio, R., Adler, G.H., Lambert, T.D., et al (2001) Ecological meltdown in predator-free forest fragments Science 294, 1923-1926 Tilman, D (1999) The ecological consequences of changes in biodiversity: a search for general principles Ecology 805, 1455-? Tregidgo, D.J., Qie, L., Barlow, J., Sodhi, N.S., and Lim, S.L.H Vertical Stratification Responses of an Arboreal Dung Beetle Species to Tropical Forest Fragmentation in Malaysia Biotropica 42, 521-525 Tscharntke, T (2010) Ecosystem services and agroecosystems in a landscape context In Conservation Biology for All, E.P.R Sodhi N.S., ed (Oxford, UK, Oxford University Press), pp 55-56 Tscharntke, T., Clough, Y., Bhagwat, S.A., Buchori, D., Faust, H., Hertel, D., Holscher, D., Juhrbandt, J., Kessler, M., Perfecto, I., et al (2010) Multifunctional shadetree management in tropical agroforestry landscapes - a review Journal of Applied Ecology 48, 619-629 Tscharntke, T., Klein, A.M., Kruess, A., Steffan-Dewenter, I., and Thies, C (2005) Landscape perspectives on agricultural intensification and biodiversity ecosystem service management Ecology Letters 8, 857-874 Turner, I.M., and Corlett, R.T (1996) The conservation value of small, isolated fragments of lowland tropical rain forest Trends Ecol Evol 11, 330-333 180 Turner, I M (1996) Species loss in fragments of tropical rain forest: A review of the evidence Journal of Applied Ecology 33, 200-209 Uduporuwa, R.S.P., Singhakumara, B.M.P., and Ashton, P.M.S (1997) Florsitics of soil seed banks in relation to light and topographic position within a Sri Lankan rain forest In Forests of the humid tropics of south and southeast Asia: A regional seminar commemorating 40 years of UNESCO in Sri Lanka, R.N.D Fonseka, ed (NARESA) UNESCAP (2003) Integrating environmental considerations into economic policy making processes Available from http://www.unescap.org/drpad/publication/integra/volume3/srilanka/3sr04a04.htm (accessed October 2010) Vasconcellos-Neto, J., (1991) Interactions between ithomiine butterflies and Solanaceae feeding and reproductive strategies In Plant–Animal Interactions:Evolutionary Ecology in Tropical and Temperate Regions Price, P.W., Lewinsohn, T.M., Fernandes, G.W and Benson, W.M (Eds.).Wiley-Interscience, New York Vulinec, K., Lambert, J.E., and Mellow, D (2006) Primate and Dung Beetle Communities in Secondary Growth Rainforests: Implications for Conservation of Seed Dispersal Systems International Journal of Primatology 34, 297-309 Vulinec, K (2002) Dung beetle communities and seed dispersal in primary forest and disturbed land in Amazonia Biotropica 34, 297-309 Wanger, T C., Iskandar, D T., Motzke, I., Brook, B W., Sodhi, N S., Clough, Y and Tscharntke, T (2010) Land-use change affects community composition and traits of tropical amphibians and reptiles in Sulawesi (Indonesia) Cons Biol 24, 795802 181 Wanger, T.C., Saro, A., Iskandar, D.T., Brook, B.W., Sodhi, N.S., Clough, Y., and Tscharntke, T (2009) Conservation value of cacao agroforestry for amphibians and reptiles in South-East Asia: combining correlative models with follow-up field experiments Journal of Applied Ecology 46, 823-832 Webb, E L and Kabir, Md E (2009) Home gardening for tropical biodiversity conservation Cons Biol 23, 1641-1644 Weerakoon D K and Goonatilake W L D P T S de A (2006) Taxonomic status of the mammals of Sri Lanka ed Bambaradeniya, C in The Fauna of Sri Lanka: Status of Taxonomy, Research and Conservation p 216 Wheeler, Q.D., Raven, P.H and Wilson E.O (2004) Taxonomy: impediment or expedient? Science 303, 285 Whitmore, T.C (1997) Tropical forest disturbance, disappearance, and species loss In Tropical forest Remnants: ecology, management, and conservation of fragmented communities, W.F Laurance, and R.O Bierregaard, eds (Chicago, Illinois, University of Chicago Press), pp 3-12 Wijesinghe, L.C.A de S., Gunatilleke, I.A.U.N., Jayawardana, S.D.G., Kotagama, S.W., and Gunatilleke, C.V.S (1993) Biological conservation in Sri Lanka: a national status report IUCN, Sri Lanka Country Office, Colombo Wijesinghe, L.C.D.A (2003) Forestry in Sri Lanka: voyage through time Tropical Agriculture Research and Extension 6, 14-21 Wijesinghe, M.R., and Brooke, M.D (2004) What causes the vulnerability of endemic animals? A case study from Sri Lanka Journal of Zoology 263, 135-140 Wijesinghe, M R., and Brooke, M de L (2005) The distribution of small mammals along a disturbance gradient in Sinharaja, Sri Lanka Journal of Tropical Ecology 21: 291-296 182 Wijesooriya, W A D A., and Gunetilleke, C V S (2003) Buffer zone of the Sinharaja Biosphere Reserve in Sri Lanka and its management strategies J Natn Sci Foundation 31, 57-71 Wikramanayake, E.D., Dinerstein, E., Robinson, J.G., Karanth, U., Rabinowitz, A., Olson, D., Mathew, T., Hedao, P., Conner, M., Hemley, G (1998) An EcologyBased Method for Defining Priorities for Large Mammal Conservation: The Tiger as Case Study .Conservation Biology 12, 865-878 Wilcove, D., McLennan, C.H., and Dobson, A.P (1986) Habitat fragmentation in the temperate zone In Conservation Biology: The science of scarcity and diversity, M.E Soule, ed (Sunderland, Massachusetts, Sinauer Associates), pp 237-256 Willott, S J Lim, D C., Compton, S G and Sutton, S L (2000) Effects of selective logging on the butterflies of a Bornean rainforest Cons Biol 14, 1055–1065 Winfree, R., Dushoff, J., Crone, E.E., Schultz, C.B., Budny, R.V., Williams, N.M., and Kremen, C (2005) Testing simple indices of habitat proximity American Naturalist 165, 707-717 Wright, S.J (2005) Tropical forests in a changing environment Trends Ecol Evol 20, 553-560 Yamamoto, W (2000) Participatory forest conservation in Southwest lowland rainforests of Sri Lanka Journal of Forest Research 5, 195-199 Zuur, A.F., Ieno, E.N., and Elphick, C.S (2010) A protocol for data exploration to avoid common statistical problems Methods in Ecology and Evolution 1, 3-14 183 APPENDICES 185 186 ... community composition and ecosystem functions The present study examines the effects of land use change and forest fragmentation on biodiversity and ecosystem functioning in Sri Lanka using community... present study examines the effects of land use change and forest fragmentation 21 on biodiversity and ecosystem functioning in the tropical lowlands of Sri Lanka More 22 specifically, the study provides... deforestation, intervention by the scientific community, shifting public opinion towards forest 10 conservation, and obligations to international conventions The concepts of biodiversity 11 conservation