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The Cultivation and Management of Chamaedorea Palms in The THE CULTIVATION AND MANAGEMENT OF CHAMAEDOREA PALMS IN THE UNDERSTORY OF A TROPICAL RAIN FOREST IN MEXICO A THESIS SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAI'I IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE IN BOTANICAL SCIENCES (Botany-Ecology, Evolution, and Conservation Biology) AUGUST 2004 By Parker Clayton Trauemicht Thesis Committee: Tamara Ticktin, Chairperson Donald R. Drake Travis Idol ACKNOWLEDGMENTS As for inspiration, the foundation ofwho I am and my love for science, nature and life - all credit goes to Rindy Trauernicht. The first set ofthanks goes to my family, who has so often provided the means and the encouragement to pursue the many opportunities that have presented themselves over the course ofmy life so far. Many thanks also to my friends here in Hawai'i and elsewhere for their support and healthy amount ofdistraction. I must thank my advisor, Tamara Ticktin, for tracking me down to let me know that she wanted me to be her student. I am truly grateful for her invaluable contribution to my education as both teacher and friend. The other members ofmy committee, Don Drake and Travis Idol, provided helpful criticism and feedback concerning methodology, analyses and the eventual write-up ofthis thesis. Thanks also to Mark Merlin for throwing around ideas. Also, mahalo to Holli and Gerry in the Botany office-the entire department would fall apart without them. Thanks to all ofthe people who helped me in Mexico. First, gracias to the folks at El Proyecto Sierra de Santa Marta in Xalapa, Veracruz. I'm truly grateful for being able to share ideas there with Fernando Ramirez who helped enormously with getting me set up and out into the field. Muchas gracias to the friends that put me up in Xalapa, especially Rosi Rodriguez and Carmen and Candido Hernandez. Also many thanks for the support ofthe director ofthe Los Tuxtlas Biosphere Reserve, Antofio Gonzalez, and all ofthe folks at the reserve office in Catemaco. Thanks to Miguel Martinez for his valuable suggestions for my research and letting me walk out ofhis lab in Morelia with $4,000 worth in camera equipment. 111 I would not have a project had it not been for the logistical support and cooperation ofthe people ofthe ejido, Adolfo Lopez Mateos. They truly made me feel part ofthe community. I must thank those that guided me in the field and helped with data collection, including Don Angel Abrajan, Don Juan Aguilar, Don Casimiro Mena, and Loco Martin. Most ofall I have to thank German Lopez, with whom I probably spent the majority ofmy time in Mexico and who is one ofthe most honest and kind­ hearted people I have ever met. This project would not have been possible without his knowledge ofthe forest and his patience with my questions. Finally, this research would not have been possible without funding through the Beatrice Kraus Fellowship from the Department ofBotany at the University ofHawai'i, a Research Award from U.H. Ecology, Evolution, and Conservation Biology program, and the Environmental Protection Agency's Minority Academic Institution Fellowship. IV ABSTRACT The cultivation ofnontimber forest products in forest understories is being promoted in many tropical regions in order to conserve habitat and provide income for rural communities. This study describes the management ofplantations ofthe palm Chamaedorea hooperiana Hodel in an area ofprimary rain forest in southeastern Mexico and assesses how management affects forest structure and composition, understory light availability and patterns ofseedling establishment. Compared to unmanaged forest, plantation sites were found to have significantly lower density and diversity ofwoody species in smaller size classes (:SID cm dbh). Analyses ofhemispherical photographs revealed that canopy openness and light availability were significantly greater in plantations than in unmanaged forest. A trend towards higher proportions ofshade­ intolerant species ofcanopy tree seedlings in plantations may reflect these altered light conditions. The conservation implications ofthese ecological changes are discussed in the context ofcommunity composition and forest regeneration. v TABLE OF CONTENTS Acknowledgments .iii Abstract. v List ofTables vii List ofFigures viii Chapter I. Introduction 1 Objectives 3 Literature Review 3 Nontimber Forest Products 3 The Ecology ofLight in Tropical Forests 6 Chamaedorea palms 9 Study Site 12 Literature Cited 14 Chapter II. The Management ofChamaedorea Plantations and the Effects on Plant Community Structure and Composition 21 Introduction 21 Chamaedorea Plantation Management. .23 Objectives 25 Methods 26 Analysis 29 Results 32 Discussion 34 Literature Cited 40 Chapter III. The Understory Light Environment of Chamaedorea Plantations and Implications for Patterns ofRegeneration 50 Introduction 50 Objectives 51 Methods 53 Analysis 56 Results 61 Discussion 64 Literature Cited 71 Chapter IV. Conclusions 81 Ecological impacts ofLimpiando 81 Effects ofManagement Variation 84 Potential for Forest Conservation 86 Future Research 87 Literature Cited 90 Appendix A. ANOVA tables for Chapter II 93 Appendix B. Functional Group Classifications 97 Appendix C. ANOVA tables for Chapter III 102 VI LIST OF TABLES 1.1. Species ofChamaedorea palms exploited in the Los Tuxtlas Biosphere Reserve 20 1.2. Prices paid for Chamaedorea leaves at each level ofthe distribution chain in Mexico .20 2.1. Results from the nested ANOVAs for percent vegetative cover. 49 3.1. Canonical structure coefficients and class means for the canonical analysis ofdiscriminance 80 Vll LIST OF FIGURES Figure 1.1. Map ofStudy Site 19 2.1. Diagram of Sampling Design .43 2.2. Plant Species Richness for Woody Species Life History Groups and Palms .43 2.3. Brillouin's Index ofDiversity for Woody Species Life History Groups and Palms .44 2.4. Simpson's Index ofDiversity for Woody Species Life History Groups and Palms 44 2.5. Overall Stem Density ofWoody Species Life History Groups and Palms .45 2.6. Stem Density ofEconomic Palm Species .45 2.7. Stem Density ofAstrocaryum mexicanum and Chamaedorea hooperiana .46 2.8. Stem Density and Relative Abundance of Timber Saplings 46 2.9. Frequency Distribution ofBasal Area Per Size Class 47 2.10. Percent Vegetative Cover for Plantation and Forest Sites 47 2.11. Size Class Distributions ofWoody Species .48 2.12. Size Class Distributions ofPalms and Cycads .48 3.1. Diagram ofSampling Design 74 3.2 Frequency Distribution ofCanopy Openness 74 3.3 Frequency Distribution ofTotal Transmittance 75 Vlll LIST OF FIGURES (continued) Figure 3.4 Seeding Size Class Distributions Based on Stem Height. 75 3.5 Seedling Size Class Distributions Based on Crown Area 76 3.6 Seeding Size Class Distributions Based on Stem Height For Each Plantation Site 76 3.7 Seedling Size Class Distributions Based on Crown Area for Each Plantation Site 77 3.8. Functional Group Proportions for Seedlings 77 3.9. Functional Group Proportions for Seedlings Excluding Block 1 78 3.10. Functional Group Proportions for Adult Trees 78 3.11. Histogram ofCanonical Scores from Multivariate Analysis ofDiscriminance 79 4.1 Photograph ofChamaedorea Plantation in Primary Forest. 92 4.2 Photograph ofChamaedorea Plantation in Secondary Forest. 92 IX CHAPTER I Introduction Tropical ecosystems are currently facing unprecedented rates ofhabitat degradation and species loss (Wilson 1999). This biological crisis is largely due to pressure from a growing human population coupled with the implementation of destructive land uses such as cattle ranching, unsustainable timber extraction and industrialized agriculture (Anderson 1990a). In response to the widespread depletion of biological resources in tropical forests, many nations and conservation organizations have promoted the establishment ofnational parks and protected areas. Alcorn (1995) notes, however, that despite the presence ofprotected areas offorests, most ofthe biodiversity in the tropics exists, and will continue to exist, in areas occupied by humans. Land management in tropical regions must therefore cope with the seemingly paradoxical problem ofmaintaining biodiversity and supporting a burgeoning human population. Although nearly all forest reserves in the tropics are surrounded by a matrix of forested and deforested land under human management, the interactions between areas of managed forest and adjacent protected areas are not well understood. Still, integrating this surrounding matrix with the conservation ofprotected areas has become the basis for the widespread establishment ofBiosphere Reserves in both temperate and tropical regions. Biosphere Reserves designate various zones ofprotection, consisting of a central zone ofstrict preservation surrounded by so-called "buffer zones," in which a variety ofland uses are practiced by local communities. The theory behind the biosphere reserve model is to employ sustainable land practices in the buffer zone that integrate habitat conservation with economic development. In practice, however, management in 1 buffer zones ranges from potentially sustainable practices such as the extraction of nontimber forest products and small-scale shifting agriculture to large-scale forest conversion for cattle ranching and agriculture. Despite the relative ubiquity ofthe Biosphere Reserve design, its overall effectiveness at conserving forested areas is still debatable (e.g., Duffy et al. 2001; Hayes et al. 2002). This study examines some ofthe ecological consequences ofthe management of forest resources in an area
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