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Edge Effects in a Forest Mosaic: Implications for Oak Regeneration in the Highlands of Chiapas, Mexico

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Edge Effects in a Forest Mosaic: Implications for Oak Regeneration in the Highlands of Chiapas, Mexico Edge effects in a forest mosaic: implications for oak regeneration in the Highlands of Chiapas, Mexico. by Fabiola López-Barrera A thesis submitted to the University of Edinburgh for the degree of Doctor of Philosophy 2003 Declaration I hereby declare that this thesis is composed of work carried out by myself unless otherwise acknowledged and that this thesis is of my own composition. The research was carried out in the period of March 2000 to September 2002. This thesis has not in whole or in part been previously submitted for any other degree or professional qualification. Fabiola López-Barrera i ii Acknowledgements I would like to thank to my supervisors Adrian Newton, Colin Legg, Robert Manson and Mario González-Espinosa for their guidance on the research design, logistics of fieldwork, and the production of this dissertation. I especially appreciate the time that Adrian Newton and Robert Manson took to visit to the field site. I also wish to thank Duncan Golicher, Pedro Quintana-Ascencio and Colin Legg for providing assistance in the statistical analysis. Thanks to Neptalí Ramírez-Marcial, Susana Ochoa-Gaona, Bruce Haines and Mike Steele who offered relevant information and valuable comments during the research. I am extremely grateful to the many people who helped at different stages with the fieldwork, including: Alfonso Luna-Gómez, Javier Bautista-Bolóm, Juan C. Bautista-Bolóm, Pedro Girón-Hernández, Maricela García-Bautista, Felipe Barragan- Torres, Henry E. Castañeda-Ocaña, Enrique Ocaña and José A. Montero-Solano. Miguel Martínez-Icó deserves a special mention for his invaluable assistance and enthusiasm on several field trips. I also wish to thank to Mario Guillén who provided me with the equipment that was necessary for the fieldwork. I am especially indebted to Consuelo Lorenzo-Monterrubio who kindly allowed me to use her live traps for small mammals and gave me access to the rodent collections. My thanks also to Oscar Retana who offered professional support in rodent manipulation and identification. I’m most grateful to Yolanda Barrera-López who helped to capture data and to tag thousands of acorns. Many thanks to the owners of the study plots in Rancho Merced Bazom who provided access to the sites: Nicolas Martínez, Pedro Pérez-Martínez, Manuel Díaz- López, Pedro Martínez-Alvarez, Miguel Pérez-Martínez and Nicolas Pérez-Ton. I am grateful to the staff of SEMARNAT (Region Altos) who transferred the seedlings from the nursery to the field sites. I am also grateful to Diego M. Díaz-Bonifaz for his assistance with map production, Hermilo Cruz-García, Mario Zuñiga-Trejo and Maria E. Martínez-Pérez for iii maintaining an efficient library service at ECOSUR. My thanks to other staff in ECOSUR including Mario Ishiki-Ishihara, Abraham Mena-Farrera, Carla Gasca- Suárez who were always very helpful. Thanks to Efraín Ascencio-Cedillo for helping me with the pictures. Particular thanks to Graham French, Cécile Bâcles, Simoneta Negrete-Yankelevich, Barbra Harvie and Tanya Ogilvy for stimulating conversations and advice during the writing up stage. Recognition is extended to the administrative and technical staff in the Institute of Atmospheric and Environmental Science at the University of Edinburgh, including Graham Walker and Connie Fox. El Colegio de la Frontera Sur (ECOSUR) provided nursery and part of the fieldwork facilities. I also visited the Instituto de Ecología A.C. Xalapa, Mexico and the World Conservation Monitoring Centre, Cambridge, over the course of the research. The British Council provided a scholarship for the first year of this research (Chevening programme Mex-2900177) and the “Consejo Nacional de Ciencia y Tecnología” (CONACYT) then funded the following three years (scholarship 131197). CONACYT also covered part of the research expenses. Financial support to pay the rent for the study plots was provided by the European Commission under the INCO- DC programme (framework 4) as part of the SUCRE project (ERBIC-18 CT 97- 0146) and the BIOCORES project (PL ICA4-2000-10029). iv Dedication To my parents Yolanda Barrera López and Roberto López Alexandre La culminación de esta tésis fue gracias al constante apoyo y estímulo de mi familia: Yolanda, Roberto, Lorena y Roberto. Mis amigos: Olga, Efraín, José Antonio, Gabriel y Tarin y a la tolerancia de mi compañero de siempre, de vida… José Antonio. v vi ABSTRACT Forest edges created by scattered-patch clear-cutting have become a common landscape feature in tropical montane forests, including pine-oak and evergreen cloud forests. Forest-edge-pasture gradients were characterised with respect to changes in vegetation structure and ecological function in the Highlands of Chiapas, Mexico. In this study two edge types were recognised by assessing vegetation structure and floristic composition along a gradient from adjacent clearings into the forest interior. The influence of edge type (either “hard” or “soft”) was investigated with respect to acorn production, predation, dispersal and germination, seedling establishment and abundance of small mammals during two consecutive years. There was no significant difference between the forest interior and the edge in plant density and basal area. Floristic richness and composition did not vary with depth-of- edge influence. Edge type influenced the relationship between probability of acorn removal by small mammals and distance from the edge in both mast fruiting and non-mast fruiting years. Movements of metal-tagged acorns supported the hypothesis that soft edges are permeable to acorn predators and/or dispersers, and that rodents move acorns up to 15 m into the grasslands in soft edges. In hard edges, acorn dispersal and removal was greater within the forest. Peromyscus spp. were the main acorn predators and/or dispersers of acorns. The intensity and rate of acorn removal varied with the year, demonstrating that in mast-fruiting years some seeds may escape predation by animals, according to the predator satiation hypothesis. Grasslands presented fewer insect damaged acorns and higher seedling emergence than the edge and the forest, emphasising the importance of acorn dispersal. Seedling survival and growth were higher for 12 m into grassland from the soft edges, suggesting that the probability of oak expansion into the grassland increases when the grasslands have been abandoned and have a scattered cover of shrubs. The results indicate that edge structure alters edge function (edge permeability), which determines the probabilities of tree invasion into old-fields. These findings have implications for the conservation of montane forest in Mexico where scattered- vii patch clear-cutting is widespread. Recommendations are made for different restoration practices to accelerate tree colonisation into the grasslands across hard and soft edges. viii Contents Contents Declaration……………………..……………………..……………………. i Acknowledgements……………………..……………………..…………… iii Dedication……………………..……………………..…………………….. v Abstract……………………..……………………..……………………….. vii Contents……………………..……………………..………………………. ix CHAPTER 1: General introduction and context of research INTRODUCTION…………………………………………………………. 1 STUDY AREA………………………………………………………….….. 5 Huistan: a forest mosaic in the Highlands of Chiapas, Mexico………... 5 Present threats to forest conservation and regeneration………………. 11 CONTEXT OF RESEARCH……………………………………………… 14 Basic concepts: Edges…………………………………………………….. 14 Edge classifications……………………………………………………….. 15 Edge effects………………………………………………………………... 16 Edge effects on species interactions……………………………………... 17 THESIS OUTLINE………………………………………………………... 19 General objective and hypothesis………………………………………... 19 Project structure…………………………………………………………. 19 CHAPTER 2: Edge effects on plant diversity and vegetation structure in a forest mosaic in the Highlands of Chiapas, Mexico ABSTRACT………………………………………………………………... 23 INTRODUCTION……………………………………………………….… 23 METHODS……………………………………………………………….… 26 Study area……………………………………………………………….… 26 Study sites……………………………………………………………….… 29 Study design………………………………………………………………. 31 Statistical analysis………………………………………………………… 33 RESULTS…………………………………………………………………... 34 Vegetation structure and composition…………………………………... 34 Edge effects………………………………………………………………... 38 Edge sharpness……………………………………………………………. 42 DISCUSSION………………………………………………………………. 45 Edge effects on plant diversity and forest structure……………….…… 45 Edge effects on forest composition………………………….…………… 47 Edge development………………………………………………………… 49 Edge sharpness and its implications…………………………………….. 50 CHAPTER 3: Permeability of forest edges: consequences for oak dispersal ABSTRACT………………………………………………………………... 53 INTRODUCTION…………………………………………………………. 53 METHODS…………………………………………………………………. 58 Experimental plots……………………………………………………….. 58 Acorn crops……………………………………………………………….. 59 Small mammal trapping…………………………………………………. 59 Observation of acorn predators and/or dispersers……………………... 60 ix Contents Post-dispersal removal trials…………………………………………….. 60 Species……………………………………………………………………... 60 Microhabitat characteristics…………………………………………….. 62 Acorn dispersal…………………………………………………………… 62 Statistical analysis………………………………………………………… 64 RESULTS…………………………………………………………………... 66 Acorn production…………………………………………………………. 66 Small mammals…………………………………………………………… 66 Other acorn consumers…………………………………………………... 69 Acorn removal……………………………………………………………. 70 Probability of detecting acorn piles……………………………………... 71 Acorn removal, microhabitat and small mammal abundance………… 74 Fate of metal tagged acorns……………………………………………… 75 DISCUSSION……………………………………………………………….
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