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Regeneration and Growth of Several Canopy Tree Species in the Maya Forest of Quintana Roo, Mexico: the Role of Competition and Microhabitat Conditions

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Regeneration and Growth of Several Canopy Tree Species in the Maya Forest of Quintana Roo, Mexico: the Role of Competition and Microhabitat Conditions AN ABSTRACT OF THE DISSERTATION OF NaDene S. Sorensen for the degree of Doctor of Philosophy in Forest Science presented on July 26, 2006. Title: Regeneration and Growth of Several Canopy Tree Species in the Maya Forest of Quintana Roo, Mexico: The Role of Competition and Microhabitat Conditions. Abstract approved: David E. Hibbs We evaluated the regeneration behavior and early growth rates of 10 non-pioneer canopy tree species in medium-height, semi-evergreen dry tropical forest in Quintana Roo, Mexico. These species provide timber and non-timber forest products for local communities and include evergreen and deciduous species with varied dispersal mechanisms. The species were Coccoloba spicata, Cordia dodecandra, Dendropanax arboreus, Guettarda combsii, Lysiloma latisiliquum, Manilkara zapota, Metopium brownei, Piscidia piscipula, Platymiscium yucatanum, and Sabal yapa. An evaluation of seedling abundances in understory and open conditions, understory seedling spatial aggregation, and adult size frequency distributions revealed divergent regeneration behaviors. Among the 10 species, we detected three processes limiting regeneration: seed availability, resource conditions, and negative density dependence. Based on observed regeneration behaviors, species were assigned to five regeneration behavior groups, reflecting, in part, inferred shade tolerance. Highly shade intolerant species would have been favored by past slash and burn agriculture. We evaluated growth responses of individuals regenerating in 0.5 ha artificial clearings to a number of biotic and abiotic factors using neighborhood measures and model selection based on Akaike’s Information Criterion (AIC). Neighbors reduced both height and diameter growth rates for most, but not all, target species within 4-5 years of opening creation. Species responses to soil factors were highly individualistic. For some species, competitive response depended on edaphic factors. Individualistic responses of species to these biotic and abiotic factors suggest that diversity may be maintained, in part, by very early niche partitioning. Neither inherent growth rate nor competitive response alone corresponded with species groupings derived from shade tolerance characteristics. However, shade tolerance was associated with an early size hierarchy among the species and with their final crown class positions. The silvicultural implications of our research results, other studies in Quintana Roo, and the historic disturbance regime are discussed. Slash and burn agriculture is identified as a central and more important component of the historic disturbance regime than hurricanes. Based on the wide range of observed regeneration behaviors, recommended silviculture techniques to maintain current species composition include variably-sized patch cuts, site preparation, control of competing vegetation, and seed tree and structural retention. Regeneracion y Crecimiento de Varias Especies Arbóreas del Estrato Superior en la Selva Maya de Quintana Roo, México: El Rol de la Competencia y las Condiciones del Microhabitat Hemos evaluado el comportamiento de la regeneración y el crecimiento inicial de 10 especies arbóreas no pioneras del estrato superior en la selva mediana subperennifolia en Quintana Roo, México. Estas especies proveen madera y productos no maderables a las comunidades locales e incluyen especies siempre verdes y deciduas con diversos mecanismos de dispersión. Las especies fueron: Coccoloba spicata, Cordia dodecandra, Dendropanax arboreus, Guettarda combsii, Lysiloma latisiliquum, Manilkara zapota, Metopium brownei, Piscidia piscipula, Platymiscium yucatanum, y Sabal yapa. La evaluación de la abundancia de la regeneración en el sotobosque y en claros, la agregación espacial de la regeneración en el sotobosque, y la distribución de adultos por frecuencia de tamaños, reveló comportamientos divergentes de la regeneración. Entre las 10 especies hemos detectado tres procesos que limitan la regeneración: la disponibilidad de las semillas, la condición de los recursos y una dependencia negativa a la densidad. Basado en el comportamiento de la regeneración observado, las especies fueron asignadas en cinco grupos, los cuales reflejan, en parte, su tolerancia a la sombra. Hemos evaluado las respuestas en crecimiento de individuos regenerando en claros de 0.5 ha a un determinado número de factores bióticos y abióticos utilizando las mediciones vecinales y la selección de modelos basados en el Criterio de Información de Akaike (AIC por sus siglas en inglés). La vegetación vecina redujo las tasas de crecimiento en altura y diámetro para la mayoría de las especies de interés para los 4 a 5 años después de la creación de los claros. Las respuestas de las especias a los factores edáficos fueron altamente individualistas. Para algunas especies, las respuestas a la competencia variaron con los factores edáficos. Las respuestas individualistas de las especies a los factores bióticos y abióticos sugieren que la diversidad arbórea podría ser mantenida, en parte, por la temprana división de los nichos. Ni la tasa de crecimiento inherente ni las respuestas a la competencia por sí solas concordaron con el agrupamiento de las especies derivado de sus características de tolerancia a la sombra. Sin embargo, la tolerancia a la sombra estuvo asociada a una jerarquización temprana de las especies por tamaño y la posición final de la copa en el dosel. Discutimos las implicancias silvícolas de los resultados de este y otros estudios y el régimen histórico de los disturbios en Quintana Roo. Identificamos la agricultura de roza, tumba y quema como componente central del régimen histórico de disturbios, más importante que los huracanes. Basado en el amplio rango de comportamiento de la regneración observado y suponiendo el objetivo de mantener la diversidad de especies, recomendamos técnicas silvícolas que incluyen la apertura de claros de tamaño variable, la preparación de los sitios, el control de la vegetación competitiva, la retención de árboles semilleros y estructurales. ©Copyright by NaDene S. Sorensen July 26, 2006 All Rights Reserved Regeneration and Growth of Several Canopy Tree Species in the Maya Forest of Quintana Roo, Mexico: The Role of Competition and Microhabitat Conditions by NaDene S. Sorensen A DISSERTATION submitted to Oregon State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy Presented July 26, 2006 Commencement June 2007 Doctor of Philosophy dissertation of NaDene S. Sorensen presented on July 26, 2006. APPROVED: Major Professor, representing Forest Science Head of the Department of Forest Science Dean of the Graduate School I understand that my thesis will become part of the permanent collection of Oregon State University libraries. My signature below authorizes release of my thesis to any reader upon request. NaDene S. Sorensen, Author ACKNOWLEDGEMENTS First and foremost I offer my sincere gratitude to Dave Hibbs. More than being just a major professor, Dave has become a friend and someone to whom I can happily say “mi casa es su casa”. I appreciate his commitment to me as a graduate student, his visits to the field sites in Mexico, and his patience and advice throughout the writing process. I also want to thank my committee members: Barbara Bond, who has been on my committee since the beginning and whose support helped me through many difficult moments along the path to this degree; Manuela Huso, who patiently helped with the statistical analyses, challenged my thinking on numerous fronts, and still found a way to make me feel good about my research; Klaus Puettman who kept me looking for ways to broaden my knowledge of forest ecology, and Ken Johnson, my graduate representative. In addition to my committee I want to thank fellow graduate students and friends who provided much needed moral support over several years, both in and out of Corvallis. Special thanks to Luisa Camara, Sarah Karr, Julia Jones, Melissa Morris, Tara Nierenberg, Jeff Shatford, and Suzanne Simard. Thanks also to Jessica Halofsky, Allison Cross and Carlos Sierra for interesting scientific, and non-scientific, conversation that made many months in the computer lab actually enjoyable. Julia Jones provided many insightful comments on Chapter 2 and Jessica Halofsky helped clarify the text. Thanks to Cristina Eisenberg for her friendship and editing of Chapter 3. In Mexico my research was facilitated by Patricia Negreros-Castillo and Laura Snook who set up the research sites and provided me with an entry into the communities where this research was conducted. I especially thank Patricia for her friendship, her enthusiasm and her collegiality. Funding and collaboration: Initial funding for setting up the treatment plots used in my studies was provided to Drs. Laura Snook and Patricia Negreros-Castillo by the Worldwide Fund for Nature (WWF). Additional funding provided to NaDene Sorensen included: Lucille Berger, Walt A. Gruetter and Forest Science Fellowships (Oregon State University), Sports Lottery Scholarship (Oregon State System of Higher Education), International Trade and Development Graduate Fellowship (Oregon State System of Higher Education). Invaluable collaboration was provided by the Instituto Tecnológico No. 16 (ITA No. 16), the Organización de Ejidos Productores Forestales de la Zona Maya (OEPFZM), and the ejidos of Limones/Cafetal, Naranjal Poniente, X-Pichil and the private ranch of the late Don Antonio Uh and his family. Numerous hours were spent in the herbarium
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