Functional Analysis of Secondary Tropical Dry Forests in a Region of the Colombian Caribbean
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FUNCTIONAL ANALYSIS OF SECONDARY TROPICAL DRY FORESTS IN A REGION OF THE COLOMBIAN CARIBBEAN Carolina Castellanos Castro This thesis has been submitted in partial fulfilment of the requirements of the degree of Doctor of Philosophy Bournemouth University October 2013 This copy of the thesis has been supplied on condition that anyone who consults it is understood to recognize that its copyright rests with its author and due acknowledgement must always be made of the use of any material contained in, or derived from, this thesis. 2 FUNCTIONAL ANALYSIS OF SECONDARY TROPICAL DRY FORESTS IN A REGION OF THE COLOMBIAN CARIBBEAN Carolina Castellanos Castro ABSTRACT Secondary tropical forests are increasingly recognized for their role conserving biodiversity in agricultural landscapes and this role is especially important for seasonally dry tropical forests (SDTF), one of the most threatened tropical forested ecosystems. The conservation value of secondary forest is increased by its capacity to maintain ecosystem properties and provide services to humans; which has been hypothesized to have positive links to the species and functional diversity of ecosystems. However very little information is available on the occurrence of this relationship in secondary forests. This dissertation makes an important contribution to the ecological knowledge of secondary SDTF and describes changes in plant species and functional diversity by using a stratified design considering different successional stages along an environmental gradient in a region of the Caribbean coast of Colombia and a multi-trait approach to study functional diversity at three scales: species, communities and landscape. The analysis of the variation in functional traits of SDTF trees at the species level allowed me to support the hypothesis of coordination between leaves and stem traits. Three dimensions of correlated variation were identified: the first related to leaf and stem economy, the second to leaf thickness and organization and the third to the trade-offs between leaf size, stem density and bark thickness. Secondary forests showed high species turnover during succession and increasing diversity from early to late forests. Species composition similarity was higher among late successional forest than early and intermediate stage forests, showing that environmental characteristics are influencing successional trajectories. Frequency distributions of species in the three successional stages showed evidence of functional trait similarity among species and underlined the importance of changes in species abundances determining functional composition during succession. A shift in abundance from individuals with traits that favour survival after disturbance to individuals with denser stems and a more conservative resource use profile was observed from early to late stages of succession. Functional composition was also strongly influenced by environmental variables, especially leaf traits, and a shift of traits from acquisitive to conservative type species was observed with increasing nutrient content and flooding, proxies of water availability. 3 Assessment of ecosystem services was conducted using two independent approaches: semi- structured interviews and field data. A high richness of useful woody plants was recorded and the provision of services was related to a high variability in functional traits among species. The provision of the ecosystem services determined by the presence and abundance of useful species showed significant differences between stages. The relationships observed between ecosystem sevices and functional and species diversity indices were not consistent. In contrast species richess showed significant negative relationships at the plot level with most of the ecosystem services assessed, showing a trade-off among the conservation of species richness and the maintenance of service provision. Overall, this research provides novel insights into the dynamic relationships between biodiversity, ecosystem function and ecosystem services in this globally important, but under-researched forest type. 4 ACKNOWLEDGMENTS This thesis integrates many of the aspects that have shaped my research trajectory, especially my growing interest in the study of plant functional ecology and my years of experience working with seasonally dry forests in Mexico and Colombia. The incredible dynamics of these forests inspired me to study in depth the functional characteristics of their plant community and its relationship with the changing management and successional conditions where they are found. However, aware of the conservation status of these forests I also wanted to make a significant contribution to the information needed to assure their sustainable management. I believe this thesis allowed me to achieve both of these aims. I’m very grateful to Adrian Newton for giving me the opportunity to conduct this research, for his help shaping my ideas and unconditional support during these three years. My fieldwork was possible thanks to all the wonderful people that accompanied me during these months. I thank Luis Motta, Tania Riveros, Gregorio Olivares and Lino Olivares for their enthusiastic support during the longs days of hard work. I especially acknowledge the patience and generosity of Lino while sharing with me his great knowledge of the plants in the region. I’m very grateful to all the owners and administrators of the farms I visited for their hospitality, especially Rafael González, Felix and Wilfran and his family. I was also very fortunate to be hosted during my field visits by Sandra Cabarcas and doña Anita, who made me feel at home and showed great interest in my venture. The amazing cooking of Nancy Olivares, Beatriz Zuñiga, Yudis Sarmiento and Enrique Jimenez definitely gave me all the strength required to survive the long hours of field work. Thanks to the people of Los Límites, Pendales, Hibácharo and Cerrito who received me in their houses and kindly participated in the interviews. Karina and Gina, thanks for your assistance organizing my fieldwork and for your hospitality in Barranquilla. The University of Atlantico allowed me to use its facilities for the analysis of plant samples, I’m very grateful to them for this. The personnel of the Colombian National Herbarium, especially Carlos Parra and Olando Rivera, were very helpful during my visit and assisted me with the identification of plant vouchers. My research was also possible thanks to the financial support of the Administrative Department of Science, Technology and Innovation of Colombia (COLCIENCIAS) for a PhD scholarship grant, and of Bournemouth University and Santander for the funds that allowed me to conduct the field campaigns. At Bournemouth, I’m grateful to Anita Diaz for the discussions that improved my research and to Louise Pearson for her help with the 5 administrative tasks. I also thank all my fellow postgraduate researchers, especially Natalia, Ivis and Ari, with whom I shared all the vicissitudes of doing a PhD. Finally, thanks to my mum, Cristian and Migue, I receive all the strength and support to embark in new adventures from you, espero seguir llevándolos conmigo a todas partes. Sergio thanks for all your patience, for cheering up my days and for always reminding me of all the good things about life. 6 LIST OF CONTENTS LIST OF TABLES ............................................................................................................... 10 LIST OF FIGURES ............................................................................................................. 15 CHAPTER 1. INTRODUCTION ....................................................................................... 18 1.1 SEASONALLY DRY TROPICAL FORESTS ........................................................... 18 1.2 FUNCTIONAL DIVERSITY AND COMMUNITY ASSEMBLY ............................ 19 1.3 SHADE AND DROUGHT TOLERANCE IN TROPICAL TREES .......................... 20 1.4 BIODIVERSITY AND ECOSYSTEM SERVICES ................................................... 22 1.5 THESIS OVERVIEW .................................................................................................. 23 1.6 STUDY AREA ............................................................................................................ 25 1.7 REFERENCES ............................................................................................................ 26 CHAPTER 2. TRAIT VARIATION AND PLANT FUNCTIONAL TYPES IN 113 WOODY SPECIES OF A SEASONALLY DRY TROPICAL FOREST ....................... 34 2.1 ABSTRACT ................................................................................................................. 34 2.2 INTRODUCTION ....................................................................................................... 34 2.3 METHODS .................................................................................................................. 37 2.3.1 Study area .............................................................................................................. 37 2.3.2 Study species ......................................................................................................... 38 2.3.3 Plant trait selection and measure ........................................................................... 38 2.3.4 Data analysis ......................................................................................................... 41 2.4 RESULTS ...................................................................................................................