Phylogenetic Analysis of Plant Community Assemblages in the Kruger National Park, South Africa
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PHYLOGENETIC ANALYSIS OF PLANT COMMUNITY ASSEMBLAGES IN THE KRUGER NATIONAL PARK, SOUTH AFRICA By KOWIYOU YESSOUFOU Thesis submitted in fulfilment of the requirements for the degree PHILOSOPHIAE DOCTOR In BOTANY In the Faculty of Sciences At the University of Johannesburg Supervisor: Prof. Michelle van der Bank Co-supervisor: Prof. Vincent Savolainen January 2012 Declaration Declaration I declare that this thesis has been composed by me and the work contained within, unless otherwise stated, is my own ------------------------------------------- K. Yessoufou (January 2012) Table of contents Table of contents……………………………………………………………………i Abstract…………….………………………………………………………………..v Acknowledgements..…………………………………………………….............vii List of tables………………………………………………………………………..ix List of figures……………………………………………………………………….x Dedication…………………………………………………………………………xiii List of abbreviations……………………………………………………………..xiv Table of contents Table of content Chapter 1- General introduction…………………………………………………..1 1. Savanna ecology…………………………………………………………………1 1.1. Features of tropical savanna………………………………………………….1 1.2. Role of fire in savanna ecology……………………………………………….3 1.3. Importance of herbivory in savanna ecology………………………………..3 2. Community ecology in a savanna context……………………………………..4 2.1. Theories of tree-grass coexistence…………………………………………..4 2.2. Tree-tree coexistence………………………………………………………….5 3. Phylogenetic investigation of community assembly…………………………..6 3.1. The use of DNA barcoding techniques to reconstruct phylogenetic tree…………………………………………………………………………………….6 3.2. Rationale of phylogenetic approach in ecology……………………………..7 3.3. Framework in community phylogenetics…………………………………….8 3.4. Emerging patterns in community phylogenetics…………………………..10 4. Study site and objectives of the study………………………………………...12 4.1. Study site………………………………………………………………………12 4.2. Objectives……………………………………………………………………...18 4.3. Outline of the thesis…………………………………………………………..19 Chapter 2- The phylogeny of trees and shrubs in the Kruger National Park using DNA barcodes…………………….………………………………………...23 1. Introduction……………………………………………………………………..23 2. Materials and methods………………………………………………………..25 i Table of contents 2.1. Sample collection…………………………………………………………….25 2.2. DNA extraction, PCR and sequencing……………………………………..25 2.3. Sequence alignment………………………………………………………….27 2.4. Tree reconstruction…………………………………………………………...28 3. Results…………………………………………………………………………..29 3.1. Tree statistics……………………………………………………………….29 3.2. Phylogeny of trees and shrubs of the KNP……………………………...30 4. Discussion………………………………………………………………………45 4.1. Clade 1: Fabidae…………………………………………………………...48 4.2. Clade 2: Malvidae………………………………………………………….50 4.3. Clade 3: Vitales…………………………………………………………….52 4.4. Clade 4: Santalales………………………………………………………..52 4.5. Clade 5: Lamiidae………………………………………………………….53 4.6. Clade 6: Campanulidae……………………………………………………55 4.7. Clade 7: Ericales…………………………………………………………...55 4.8. Clade 8: Caryophyllales…………………………………………………...56 4.9. Clade 9: Basal Eudicots…………………………………………………..56 4.10. Clade 10: Magnoliidae…………………………………………………….57 4.11. Clade 11: Monocotyledoneae…………………………………………….57 5. Conclusion………………………………………………………………………58 Chapter 3- Testing suitability of evolutionary models using traits of woody plants in the KNP…………………………………………………………………..59 1. Introduction..……………………………………………………………………59 2. Materials and methods….…………………………………………………….61 ii Table of contents 2.1. Plant ecological traits………………………………………………………61 2.2. Data collection and measurements………………………………………63 2.3. Data analysis……………………………………………………………….64 3. Results………………………………………………………………….............65 4. Discussion………………………………………………………………………71 5. Conclusion………………………………………………………………………74 Chapter 4- Characterising diversity and phylogenetic structure of woody plant communities in the Kruger National Park, South Africa………………………..75 1. Introduction……………………………………………………………………..75 2. Materials and methods………………………………………………………...77 2.1. Dataset………………………………………………………………………77 2.2. Data analysis……………………………………………………………….78 3. Results…………………………………………………………………………..82 3.1. Overall diversity and community structure in the KNP.………………..82 3.2. Community phylogenetic structure within and among sites…………...84 3.3. Community trait-based structure within and among sites……………..86 4. Discussion………………………………………………………………………86 5. Conclusion………………………………………………………………………89 Chapter 5- The role of megaherbivores in shaping the structure of subtropical plant communities...………………………………………………………………..90 1. Introduction……………………………………………………………………..90 2. Materials and methods………………………………………………………..94 2.1. Study site: exclosures……………………………………………………..94 iii Table of contents 2.2. Traits of anti-herbivores defences………………………………………..96 2.3. Community sampling in the exclosures………………………………….96 2.4. Statistical analyses………………………………………………………...97 3. Results…………………………………………………………………………..98 4. Discussion……………………………………………………………………..103 4.1. Exclusion of megaherbivores and plant diversity………………………..104 4.2. Exclusion of megaherbivores and phylogenetic diversity………………105 5. Conclusion…………………………………………………………………….106 Chapter 6- General conclusion…..…………………………………………….108 1. Major findings, discussion and contribution to literature..………………..108 1.1. Phylogenetic information database for the KNP………………………108 1.2. Ornstein-Uhlenbeck is more suitable for phylogenetic comparative analysis of plant traits in the KNP………………………………………109 1.3. Plant community assemblages in the KNP are not neutral………….109 1.4. Megaherbivores leave distinct signature on plant community structure…………………………………………………………………...109 2. Future challenges…………………………………………………………….111 Chapter 7- References…………………………………………………………..114 Supplementary Information…………………………………………………...139 Appendix- Submitted paper…………………………………………………….176 iv Abstract Abstract What underlies species distribution and species coexistence has long been of key interest in community ecology. Several methods and theories have been used to address this question. However, it still remains a controversial debate. The recent development of plant DNA barcodes with possibility of merging phylogeny with ecology brings high expectation in uncovering the processes underlying community assemblages. Previous works that used molecular approach in community ecology focused mainly on rainforests. Using a phylogenetic approach, this study brings novel understandings about savanna ecology, especially regarding how megaherbivores impact plant community composition. The Kruger National Park (KNP) is one of the world’s largest reserves, but less studied from a phylogenetic perspective. A DNA database of 445 DNA sequences (plant DNA barcodes, rbcLa + matK) was generated for the woody plants of the KNP. This database proves reliable in reconstructing the phylogeny of Angiosperms of the park. Based on this phylogeny, the present study characterised plant community composition, and investigated how megaherbivores influence this composition. Results indicate that plant communities in the KNP are not neutral, i.e. they are more clustered than expected under various null models. This suggests that ecological forces, most likely habitat filtering may be playing key role in dictating community structure in the KNP. The KNP is well-known for its richness in megaherbivores. The contribution of these animals to the current shape of plant community structures was therefore further investigated. Where megaherbivores have been excluded, plant diversity decreases, but shifts in plant community structure are contingent upon the initial community composition, suggesting that herbivory might be important filter that drives the clustering pattern observed. v Abstract These results also have important implications for management of African woodlands, particularly given the continental decline in megaherbivores. As large herbivores are lost from these ecosystems, one can predict a subsequent reduction in plant diversity, whilst the impact on plant community structure will depend upon the initial composition. Critically, I also show that the loss of phylogenetic diversity (a surrogate for functional diversity) will depend on the relative shifts in phylogenetic community structure, information that has never been considered before in management strategy. Key words: Community phylogenetics, functional diversity, species coexistence, under/overdispersion, evolutionary models, megaherbivores, conservation, extinction, Kruger National Park, South Africa. vi Acknowledgements Acknowledgements This project was financially supported by the University of Johannesburg (UJ, South Africa), the National Research Funds (NRF, South Africa), the Government of Canada through Genome Canada and the Ontario Genomics Institute (2008-OGI-ICI-03), and the Royal Society (UK). The massive contribution of the Canadian Centre for DNA Barcoding (CCDB) through assistance in DNA sequencing is also acknowledged. I am grateful to my supervisors Prof. Michelle van der Bank (UJ, South Africa) and Prof. Vincent Savolainen (Imperial College London, UK) for the wonderful roles they played not only during the run of the project, but also for their guidance that leads to the production of this dissertation. Dr T. Jonathan Davies (McGill University, Canada) also provided invaluable assistance. The University of Johannesburg through the Department of Botany provided excellent work and social environment and must receive here my gratitude. I would like to thank researchers from UJ, especially Dr Motsi Cynthia Moleboheng (former PhD student), who took me