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Shifting Cultivation Effects on Brazil Nut (Bertholletia excelsa) Regeneration


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1 SHIFTING CULTIVATION EFFECTS ON BRAZIL NUT ( Bertholletia excelsa ) REGENERATION By JAMIE NICOLE COTTA A THESIS PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE UNIVERSITY OF FLORIDA 2007

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2 Copyright 2007 by Jamie Nicole Cotta

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3 To my parents, who made it all possible.

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4 ACKNOWLEDGMENTS This thesis could not have been complete d without the steadfast support of my ever encouraging advisor, Dr. Karen Kainer, who ge nerously shared her e xpertise and provided counsel throughout the research and writing processe s. I am equally grateful to my committee member, Dr. Lcia Wadt, who welcomed me into her home in Brazil three years in a row, offering not only research advice, but personal and professional guidance th at contributed to the success of my project in Brazil. I also tha nk the members of my supervisory committee: Dr. Emilio Bruna and Dr. Kaoru Kitajima, for their comments and suggestions during the writing of my thesis. I am immensely thankful for Chri stine Staudhammers statis tical counsel; she has a true gift for communicating her st atistical expertise with forestry professionals. I also thank Meghan Brennan for sharing he r statistical knowledge. I sincerely thank my internat ional sponsor, EMBRAPA-Acre ( Empresa Brasileira de Pesquisa Agropecuria) in Rio Branco, Brazil, for gener ously supporting this research. A number of Brazilian colleagues and assistants provided essential insights and offered their talents to this research project, including Paulo Wadt Paulo Rodrigues de Carvalho, Pedro Raimundo R. de Arajo, Aldeci da Silva O liveira, Airton, Freire, Sergio, a nd Rivelino. I would like to sincerely thank the extractivist families, specifi cally Valderi, Maria Alzenira, Duda and Bil Mendes, and Amarilzo da Rocha Bento, for generously sharing their insights, and providing me with a true home away from home. Their j oyful, affectionate child ren will also never be forgotten. I would also like to thank Tim Martin, Christie Klimas, Cara Rockwell, Chris Baraloto, Marisa Tohver, Roberta Veluci, Al exander Cheesman, Skya Rose Murphy, Shoana Humphries, Valerio Gomes, and Marco Lentini fo r their contributions to my final draft. This research would not have been possible without the support of my Department, the School of Natural Resources and Environment, a nd a generous scholarship from the School of

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5 Forest Resources and Conservation. I am also grateful for summer research grants from the University of Florida Graduate School a nd the Tropical Conservation and Development Program. The Institute of Food and Agricultural Sciences, the School of Natural Resources and Environment, and the UF Graduate Student Council also generously provided me with finds to return research results to communities. Most importantly, I thank my family. My pare nts and sister represent the most admirable of role models and, through their guidance and li ving examples, I have learned how to face great challenges and accomplish personal goa ls, while never losing focus of others. Because of them I understand true compassion and generosity of sp irit. My grandparents, aunts, uncles, and cousins have each shared a special part of them selves, which have all contributed to my success and, without their presence in my life I would not have been able to pursue and complete this degree.

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6 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................4 LIST OF TABLES................................................................................................................. ..........8 LIST OF FIGURES................................................................................................................ .........9 ABSTRACT....................................................................................................................... ............10 CHAPTER 1 INTRODUCTION..................................................................................................................12 2 SHIFTING CULTIVATION EFFECTS ON BRAZIL NUT ( Bertholletia excelsa ) REGENERATION.................................................................................................................17 Introduction................................................................................................................... ..........17 Species Description............................................................................................................ ....19 Study Area..................................................................................................................... .........20 Methods........................................................................................................................ ..........21 Plot Installation and Sampling Scheme...........................................................................21 Regeneration Environments by Forest Type...................................................................22 Seedling and Sapling Densities.......................................................................................24 B. excelsa Survival and Growth......................................................................................24 Data Analysis.................................................................................................................. .25 Results........................................................................................................................ .............28 Regeneration Environments by Forest Type...................................................................28 Seedling and Sapling Densities.......................................................................................29 B. excelsa Survival and Growth......................................................................................29 Seedlings..................................................................................................................29 Saplings....................................................................................................................30 Discussion..................................................................................................................... ..........31 Densities Explained.........................................................................................................31 Seed sources and dispersal.......................................................................................31 Seedling survival and growth...................................................................................33 Sapling survival and growth.....................................................................................36 Greater Ecological and Ma nagement Implications.........................................................37 3 CONCLUSION..................................................................................................................... ..39 APPENDIX A TRANSECT CREATION......................................................................................................46 B SCHEMATIC OF PLOT INSTAL LATION AND SAMPLING SCHEME..........................47

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7 LIST OF REFERENCES............................................................................................................. ..48 BIOGRAPHICAL SKETCH.........................................................................................................55

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8 LIST OF TABLES Table page 2-1 Mean ( sd) soil properti es observed within fallows and mature forest plots in RESEX Chico Mendes and PAE Chico Mendes...............................................................43 2-2 Mixed model results based on B. excelsa seedling and sapling data subsets....................44 2-3 Comparison of B. excelsa seedling and sapling characte ristics in fallow and mature forest......................................................................................................................... .........45

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9 LIST OF FIGURES Figure page 2-1 Location of study sites PAE Chico Me ndes and RESEX Chico Mendes in Acre, Brazil......................................................................................................................... .........41 2-2 Mean ( SE) PPFD at four times of day in four fallows and two mature forest plots.......41 2-3 Bertholletia excelsa seedling and sapling densitie s (means SE) in fallow and mature forest.................................................................................................................. ....42 2-4 Relationship between B. excelsa seedling height and photon flux density (PPFD) measured in 2005...............................................................................................................42 2-5 Relationship between B. excelsa seedling total leaf area and photon flux density (PPFD) measured in 2005..................................................................................................43 B-1 Mature forest and fallow plot design................................................................................47

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10 Abstract of Thesis Presen ted to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Science SHIFTING CULTIVATION EFFECTS ON BRAZIL NUT ( Bertholletia excelsa) REGENERATION By Jamie N. Cotta May 2007 Chair: Karen Kainer Major Department: School of Natural Resources and Environment Brazil nut ( Bertholletia excelsa ), has emerged as the cornerst one of the extractive economy in much of the Amazon, but the debate continues as to whether or not curr ent harvest levels have a detrimental effect on Brazil nut seedling recruitm ent. Regeneration studies to date have been conducted solely within mature forest, but my st udy provides further insight into current Brazil nut regeneration dynamics, with a unique first lo ok at regeneration in swidden fallows within two multiple-use areas in Acre, Brazi l. Recruitment of individuals 10 cm dbh was evaluated within 25 x 25 m and 10 x 10 m subplots in four 9 ha mature forest plot s and six fallows (0.5-1.0 ha each), respectively. Individuals 10 cm dbh were grouped into two size classes: seedlings (individuals < 1.5 m in height) and saplings (i ndividuals between 1.5 m in height and 10 cm dbh). General linear mixed model anal yses revealed higher densities of B. excelsa in fallow than mature forest. Survival and growth of young B. excelsa individuals appear to be enhanced by higher light levels found in fallows Greater light availability is positively related to seedling height and height growth, seedli ng leaf number and leaf area, and seedling survival, and may indirectly enhance saplin g growth and survival. This enha nced survival and growth likely contributes to the observed higher densities of seedlings and sap lings in fallows. Not only can anthropogenic disturbance, in the form of shifting cultivation, play a posit ive role in Brazil nut

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11 regeneration, it could explain curre nt Brazil nut densities and distri butions. Finally, in light of these findings, swidden fallows could potentially be managed for enhanced Brazil nut densities, which may provide an opportunity for greater in come for extractive families while contributing to the sustainability of Brazil nut extraction in the long term.

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12 CHAPTER 1 INTRODUCTION Degraded and secondary forests are widesp read throughout the tropi cs, representing 60% of remaining tropical forests (ITTO 2002). In the Brazilian Amazon alone, over 12% of the forest mosaic is classified as degraded and secondary forest (ITTO 2002). As these forests undergo succession they are capab le of conferring significan t environmental and livelihood benefits. Secondary forests provide people w ith food resources, timber and non-timber forest products, including fuelwood, and qua lity hunting sites (ITTO 2002). They also regulate water regimes, protect soils from er osion, store carbon, and serve as biological corridors and refuges for species (ITTO 2002). In additi on, regenerating secondary forest s are especially important in the life histories of a number of tropical plant species, including some that are of particular socioeconomic and ecological importance, such as Cedrela spp and Swietenia spp (Whitmore 1989). Similarly, two other valued species, Astrocaryum murumuru and Dipteryx micrantha, benefit from seed dispersal to treefall gaps (Cintra and Horna 1997) All four of these species exhibit gap-opportunistic traits, achieving enhanced growth in higher light conditions found in canopy openings or young regenerating forest environments. Brazil nut ( Bertholletia excelsa, H.B.K. ) is another highly valued tropical tree reported to be a gap-opportunistic speci es (Mori and Prance 1990, Myers et al 2000), benefiting from canopy openings during early stages of its life history. Brazil nut has emerged as the cornerstone of the extractive economy in much of the Amazon (Clay 1997), and the seeds, commonly referred to as nuts, are used for a variety of products, including raw and dried nuts, oils, flour, medicines, and personal care products (Ortiz 199 5), making Brazil nut a versatile and valuable non-timber forest product (NTFP). To assess recru itment status as related to long-term harvest sustainability of this valuable species, several regeneration studies have been conducted in recent

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13 years (Viana et al 1998, Zuidema and Boot 2002, Peres et al. 2003, Serrano 2005), yielding differing reports on the impact of nut harvest on Brazil nut population st ructures. However, studies to date have been conduc ted solely within mature forest sites. The modern Amazonian landscape consists of a mosaic of mature forest, regenerating s econdary forest, swidden fallow, and pasture; therefore, in light of the increasing extent of seconda ry forests in the region, the role of these sites in critical economic and conservation species recruitment and regeneration processes should not be underestimated. In Acre, Brazil, local extractivists report rela tively high levels of Brazil nut recruitment in regenerating swidden fallows, which may be due to specific biotic and abiotic characteristics associated with these sites, in cluding differences in seed disp erser activity (Clay 1997, personal obs.), light availability (Kainer et al. 1998, Myers et al. 2000, Zuidema 2003), and nutrient availability (Kainer et al 1998), as well as increased resistan ce to pathogens in higher light environments (Augspurger 1984b). The aim of my study is to assess to what extent swidden fallows constitute favorable regeneration environments for B. excelsa Although various studies have assessed the performance of tropical s eedlings in natural gaps (Popma and Bongers 1991, Cintra and Horna 1997, Myers et al. 2000), typically formed by treef alls, conclusions regarding recruitment, survival and growth in these canopy openings should not be a pplied to all cases of canopy disturbance. For the purpose of this study I highlight some characteristics unique to secondary fallow forests, in comparison with treef all gaps, as well as othe r regenerating forest types, which may each contribute considerably different influences on B. excelsa regeneration. Fallows forests differ from treefall gaps in many ways. To begin with, the most common treefall gap size is < 20 m2 (Brokaw 1982), while swidden plots av erage 2 hectares (equivalent to 20,000 m2) (Fujisaka et al. 1998). Abiotic conditions, includi ng light and temperature levels,

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14 vary within gaps (Denslow et al. 1990). As a result, one might observe a larger range of light and temperature levels in larger gap areas, such as swidden plots. Also, herbivore activity may differ as a result of this difference in gap size (Janzen 1990), as some vertebrates prefer secondary forest environments (Pea-Claros an d De Boo 2002), but othe rs avoid crossing large clearings (da Silva et al. 1996). What may differentiate fallows most from othe r secondary forests is the influence of land use history on succession and regeneration (Uhl et al. 1988, Guariguata and Ostertag 2001, Mesquita et al. 2001). In general, swidden fallows devel op after 2 to 4 years of cultivation, and are left to regenerate for a period of 5 to 15 years (Montagnini and Me ndelsohn 1997). In fields which have been subjected to little burning, Cecropia spp. are often the dominant pioneers to establish in young fallows, however, wher e burning has been more frequent, Vismia spp often dominate (Borges and Stouffer 1999). The dominance of these early pioneers directly influences subsequent vegetation succession (Nascimento et al. 2006). For example, in Cecropia dominated areas heavy litterfa ll and increased shading can inhibit seedling establishment (Mesquita 1995, cited in Nascimento et al. 2006). As succession proceeds, differences in forest structure, stem density, and species composition can be observed. Fujisaka et al. (1998) found species richness to be higher in 3 5 year old fa llows than in 1 2 year old fallows, and higher in fallows than in pasture sites. Also, vines we re more abundant in mature forest and fallows of 3 5 years than in cultivated sites and pasture (Fujisaka et al. 1998). Furthermore, grass and weed dominance was reduced in fallows, compared to pasture and agroforestry sites, resulting in less competitive suppression (Ferguson et al. 2003). More shrub speci es were found in cropped and fallow sites than in mature forest and pasture (Fujisaka et al. 1998), and herbaceous plants were most common in cropped fields and pastur e. Finally, as a resu lt of burning and crop

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15 harvesting, nutrient availability may be significan tly altered in young fallows (Uhl and Jordan 1984). After a few years of succe ssion, large regenera ting clearings may harbor increased numbers of vertebrates due to gr eater food availability for certain species than that of mature forest (Janzen 1990). Composition of seed dispersers may also differ between gap types, as a result of hunting activities th at often coincide with crop cultivation (Janzen 1990). In large man-made clearings, new recruitment is dependent upon seed dispersal (Ferguson et al 2003), if cropping activiti es are carried out such that adu lts of desired species are removed or avoided, and no seeds are present in the seed bank. In this case, near by patches of mature forest are required for forest tree s to colonize abandoned plots (Uhl et al. 1982). On the other hand, recruitment may result from both dispersal and adult seed rain in a treefall gap. Although swidden fallows are often surr ounded by other fallows or secondary forests of varying ages (Ferguson et al. 2003), recruitment processes may not be drastically compromised relative to those in treefall gaps, since matu re forest is often left intact along at least a portion of the swidden edge (personal obs.). All of the aforem entioned differences associated with forest gap type are relevant to any analys is of recruitment and regenerati on processes of species such as B. excelsa in disturbed areas. My study offers a new look at the seedling ec ology of a valuable NTFP, through a focused study on the post-germination phase of B. excelsa regeneration within two multiple-use areas in the southeastern portion of Acre, Brazil. I predicted that densities of B. excelsa individuals 10 cm diameter at breast hei ght would be greater in fallow than mature forest for two main reasons: 1) local extracti vists report relatively high agouti se ed dispersal in fallows (Clay 1997), and 2) survival and growth of young B. excelsa individuals is reported to be enhanced in the

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16 presence of higher light av ailability in forest gaps (Mori and Prance 1990, Myers et al. 2000). In addition to evaluating seeding and sapling densities, light levels soil nutrient content, leaf herbivory level, and number of nearby seed s ources were evaluated in both fallow and mature forest to determine their respective influences on B. excelsa survival and growth. Chapter 2 is presented as a single, independent chapter rea dy for submission to a peer-reviewed journal. Relevant conclusions and management imp lications are presented in Chapter 3.

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17 CHAPTER 2 SHIFTING CULTIVATION EFFECTS ON BRAZIL NUT ( Bertholletia excelsa ) REGENERATION Introduction Brazil nut ( Bertholletia excelsa, H.B.K.), has emerged as the cornerstone of the extractive economy in much of the Amazon (Clay 1997), which has given rise to the debate over whether or not current harvest levels have a detrimental effect on Brazil nut seedling recruitment. Some scholars report that current ha rvest levels have minimal to no detrimental effect on seedling recruitment in selected Br azil nut populations (Viana et al 1998, Zuidema and Boot 2002), while others report that regeneration is rare or nonexistent in ove r-exploited populations (Peres et al. 2003, Serrano 2005). All previous Brazil nut rege neration studies have been conducted solely within mature forest, however, species regenera tion is not limited to th is forest type. The modern Amazonian landscape is a mosaic of mature forest, regenerating secondary forest, swidden fallow, and pasture; which comprise a continuum of microhabitat suitability for B. excelsa recruitment and regeneration. An evalua tion of Brazil nut regeneration ecology is needed across the entire mosaic of forest types in orde r to provide an accurate picture of current B. excelsa population structures. Bertholletia excelsa may regenerate more successfully in secondary than mature forest, based on reports that it is a gap-opportuni stic species (Mori and Prance 1990, Myers et al 2000). Bertholletia excelsa is one of a number of shade-tolerant species which experience enhanced seedling survival and growth in canopy openings, characterized by higher light levels (Brokaw 1985, Whitmore 1989, Molofsky and Fisher 1993, Cint ra and Horna 1997). According to Myers et al (2000), B. excelsa seedlings not only benefit from fore st gaps, they require a minimum gap size of > 95 m2 or > 10.4% global site factor in order to reach sapling size. One large-scale disturbance yet unexplored for B. excelsa recruitment is that of recent anthropogenic secondary

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18 forests, which, in addition to being widespre ad throughout Amerindian history (Denevan 1992, Heckenberger et al. 2003), are becoming increasingly common in the Amazonian landscape (ITTO 2002). In Acre, Brazil, landholders annua lly clear 0.5 2 ha patches of older secondary or mature forest for swidden agriculture. When agricultu ral sites are abandoned, uni que secondary forests succeed in fallow sites, which may constitute par ticularly favorable regeneration environments for B. excelsa Local extractivists describe fallows as having greater B. excelsa recruitment than mature forest, associated with a relatively highe r incidence of agouti seed dispersal to these disturbed sites. In addition, seedling survival ma y also be enhanced in fallow, due to decreased vulnerability to pathogens in higher light environments (Augspurger 1984b). Subsequent establishment and growth of B. excelsa may be greater in fallows because of high light levels (Kainer et al. 1998, Myers et al. 2000, Zuidema 2003) and greater nutrient availabi lity (Kainer et al 1998). An evaluation of the potential of fa llow sites to provide favorable B. excelsa regeneration environments will complement previous assessments of tropical seedling performance in other disturbed sites, specifically natural gaps (Popma and Bongers 1991, Cintra and Horna 1997, Myers et al. 2000). Although both types of di sturbance increase forest light availability, fallows represent ecologically unique microhabitats which may differentially affect seedling recruitment, survival and growth. As a result of the shifting cultivation process, which often includes the application of fire, fallow forests may be charac terized by significantly di fferent nutrient stocks (Uhl and Jordan 1984), plant assemblages (Fujisaka et al. 1998, Ferguson et al. 2003), and vertebrate activity (Schupp 1988, Cintra a nd Horna 1997, Pea-Claros and de Boo 2002).

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19 This study directly tests the hypothesis, based upon aforementioned scientific evidence and extractivist observations, that B. excelsa seedling and sapling densities are greater in fallows than mature forest, by assessing the post-germination phase of B. excelsa recruitment. To understand some of the mechanisms behind observed recruitmen t, effects of light, nutrient availability and proximity of seed sources are also assessed. In addition, one-year survivor ship and growth rates of young individuals, seedling leaf production, an d seedling herbivory, which further shape B. excelsa recruitment, are compared between fallow and mature forest. Species Description Bertholletia excelsa is a monospecific member of the Lecythidaceae family, found in unflooded (terra firme) forests across the Am azon basin and the Guianas (Mori and Prance 1990). This canopy emergent can reach up to 50 m in height (Mori and Prance 1990), and some individuals have been recorded at almost 1000 years of age (Vieira et al. 2005). In the two landholdings surveyed in my study (Filipinas an d Cachoeira), densities of Brazil nut trees 10 cm diameter at breast hei ght (dbh) were 1.4 individuals ha-1 (Wadt et al 2005) and 2.5 individuals ha-1 (Serrano 2005), respectively, which are representative of densities reported by Peres et al. (2003) elsewhere in the study region. Although some authors report that B. excelsa occurs in groves of 50 100 individuals, with ea ch grove separated from another by distances of up to 1 km (Mori and Prance 1990), Wadt et al (2005) reported no existe nce of groves in my study area. Mammal communities play a critical role in th e regeneration of many large-seeded tropical tree species (Sork 1987, Asquith et al. 1997, Snchez-Cordero and Martinez-Gallardo 1998, Forget et al. 2000, Jansen 2003), via seed dispersal a nd seed predation. Brazil nut is no exception, as the regeneration of B. excelsa is highly dependent upon the presence of agoutis (Dasyprocta spp.) the primary dispersers of Brazil nut seeds (Huber 1910, Prance and Mori

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20 1978). Agoutis have been observed burying seeds at clearing edges (Peres and Baider 1997) and in young secondary forest (Forget et al. 2000), and extractivists ha ve often observed agoutis carrying seeds to fallows (Clay 1997, personal obs.). Agoutis may prefer fallow sites because of the protection provided against predators by re generating vegetation (Cintra and Horna 1997). These reports suggest a possibili ty of comparable or greater seed dispersal to fallows than mature forest. Recruitment of B. excelsa in both mature forest and fallows may also depend upon density and proximity of reproductiv e adults, as the proportion of s eeds that arrive at a certain point in the forest should dec line with distance from parent tree (Janzen 1970). Agoutis have been observed dispersing B. excelsa seeds as far as 100 m from seed sources, with the majority of seeds being dispersed within 25 m (Peres a nd Baider 1997). Normal seed shadows can be extended by secondary dispersal, when seeds are un earthed by seed predat ors and reburied even further from the adult tree (Mori and Prance 1990, Peres et al. 1997). Study Area Fieldwork was carried out in th e southeastern portion of Acre Brazil, between 10 and 11 south of the equator in Acre, Brazil. The re gion has undulating topograph y, vegetation classified as humid, moist tropical forest (Holdridge 1978), and a pronounced three month dry-season from June to August. Average annual rainfall is from 1600-2000 mm (IMAC 1991). Soils of the region are classified under the Brazilian cla ssification system as Argissolos (ZEE 2000), which correspond roughly to U.S. Soil Taxonomy System Ultisols. Recruitment of B. excelsa was evaluated within RESEX Chico Mendes and th e Chico Mendes Agro-Extractive Settlement Project (PAE Chico Mendes), multiple use areas of 976,570 ha and 24,898 ha, respectively, separated by 30 kilometers (Fig. 2-1). Extractivis ts in the two study sites subsist on agricultural and cattle production, rubbe r tapping and Brazil nut collectio n (Gomes 2001), and some timber extraction in PAE Chico Me ndes (personal obs.).

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21 In my study sites, 0.5 1.5 ha of mature or older secondary forest found in close proximity to households was manually cleared of large trees to initiate the shiftin g cultivation cycle. Clearings were created either in areas void of valuable Brazil nut and rubber ( Hevea brasiliensis ) trees, or great care was taken to protect indivi duals of these species. Clearings were burned once, first cultivated for corn and rice, followed the next year by beans, and then manioc. After a cultivation period of two to three years, agri cultural sites were abandoned, and regenerating secondary forests formed in situ. Young sec ondary forests such as these are typically characterized by lower stand basal area than surrounding mature forest (Saldarriaga et al. 1988). Although these swidden fallows ar e often surrounded by other fallo ws or secondary forests of varying ages, mature forest borde red at least one length of each fallow border studied. Thus, the study landscape consisted of a forest matrix w ith small fallows and pastures embedded within mature forest expanses. Methods Plot Installation and Sampling Scheme Mature forest plots (300 x 300m or 9 ha each) in RESEX Chico Mendes and PAE Chico Mendes were originally demarcated and delin eated in 2002 (Serrano 2005) using transects opened every 25m and extending fo r 300m (Appendix A). Transect s produced 25 x 25m grids of 9-hectare survey plots (Fig. B-1[ A]). From May to July 2005, tr ansects within three of these original plots in RESEX Chico Mendes were reopened to evaluate all B. excelsa individuals > 10cm dbh by searching to the right and left of systematically placed parallel lines spaced 25m apart. To survey 25 % of the area of each mature forest plot for recruitment, four mature forest subplots (25 x 25 m each) were randomly selected w ithin each hectare. Thus randomization was restricted to each one-hectare plot, gene rating a total of 36 su bplots (Fig. B-1[A]).

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22 Six fallows formed in RESEX Chico Mendes (0.5 1.0 ha each), and two in PAE Chico Mendes (1.0 1.5 ha each), were also delineate d in 2005 (Fig. B-1[B]), using transects opened every 10m. Fallow ages ranged from 5 to 12 y ears, based upon year of abandonment of the agricultural site and, due to their variable sizes and shapes, were delineated and demarcated with a Garmin 12XL3 GPS unit. All individuals > 1.5 m in height were located in fallows by searching to the right and left of parallel lines spaced 10 m ap art. Recruitment subplots in fallows (10 x 10 m) were randomly assigned acro ss the extent of each fallow, and comprised approximately 25% of each fallow area (Fig. B-1[B]). Regeneration Environments by Forest Type To evaluate the influence of seed sources on recruitment, all B. excelsa individuals > 10 cm dbh were tallied within ea ch 9 ha mature forest and 0.5 1.5 ha fallow, as well as within a 50m border strip of each plot. Dbh, XY coordinate s, and reproductive status were recorded for each individual. To compare understory light availability am ong 2 mature forest plots and 4 fallows in 2005, photosynthetically active radi ation (PAR) was estimated by averaging photosynthetic photon flux density (PPFD) measurements taken at 10 equidistant sample points running through the center of each plot, from one plot edge to the opposite edge (Appendix B). Measurements were taken every 30 m in mature forest plots a nd measurement spacing varied in fallow sites, due to variation in fallow size. To capture day time variations, PPFD was measured in each plot every 2.5h from 7:30 to 15:00 (ie. four times per day) for three days. PPFD was measured using 10 gallium arsenide phosphide (GaAsP) photocells mounted on a 10-cm disk and connected to a multimenter (Micronta LCD Digital Multimet er 22-185A; Tandy Corporation, Fort Worth, Texas). Sensors were calibrated with a standa rd LI-COR quantum sensor (model LI190SA) and a Campbell CR10x data logger.

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23 To determine soil properties within each fallow and mature forest plot, soil samples were collected in 2005 below the litter layer at two depths (0 20 cm and 20 40 cm). In the fallows, soil cores were collected at two depths within on e randomly selected subplot in each of the four corners of the fallow, as well as the center-most subplot, to cr eate one composite of five soil cores for each depth (Appendix B). In the 9 ha mature forest plots, soil cores were collected within the four hectares located at the corners of each plot, as well within the center-most hectare (Appendix B). Within each of these five sample d hectares, a composite of five soil cores was created for each depth, using the same sampling scheme described for soil collection in the fallows. Two replicates of each composite were dried for 4 days at 65C, and passed through a 2 mm stainless steel sieve. Soil pH was measured at a 1:2.5 soil to water ratio. Extractable P and K were processed using a dilute double acid extracti on (Mehlich-1), with c oncentrations of P determined colorimetrically using the molybda te blue method (Murphy and Riley 1962), and K concentrations were determined using flame emission spectrophotometry. To determine total potential acidity, H+ and Al3+ were extracted with a buffered so lution of calcium acetate at pH 7, and then titrated with a 0.1 N NaOH. Oxidizab le organic carbon was determined on soils passed through a 1 mm screen, ground in a porcelain mort ar, and then digested in a potassium dichromate acid medium with external heat. All soil analyses were conducted at the Soils Laboratory of Embrapa-Ac re, Brazil (EMBRAPA 1997). The temperature and moisture environment in fallow and mature forest were evaluated between July 2005 and June 2006. Mean monthly temperature and percent relative humidity were determined for two fallow sites and one matu re forest site based on measurements taken at 30 minute intervals using HOB O ProSeries data loggers.

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24 Seedling and Sapling Densities Seedling densities were assessed w ithin all subplots installed in mature forest and fallows. Sapling densities in mature forest also were asse ssed within subplots. Du e to the ease of locating saplings in fallow, however, sapling densities we re assessed over entire fallow areas, rather than in subplots only. Individuals < 1.5 m in height were defined as seed lings and individuals between 1.5 m in height and 10 cm dbh were defined as saplings. This classification scheme was selected not to differentiate betw een those individuals still using their seed reserves for growth and those individuals relying on e nvironmental resources, but rather to look at general classes of recruitment. Within each subplot, XY location c oordinates, height, seed ling basal diameter, and sapling dbh were recorded for all individuals. B. excelsa Survival and Growth Survival and annual growth (seedling height and basal diameter and sapling dbh) were evaluated from June to July 2006. To evaluate the effect of PAR on seedling survival and growth, PPFD was measured directly above seed lings in two mature forest plots and four fallows. To capture daytime variations and to m easure the full complement of all seedlings in each selected plot, seedling PPFD was sampled on a rotating basis every 2.5h from 6:45 to 16:45 (ie. five sampling times per day) for three da ys. To assess the influence of photosynthetic capacity on survival and growth of all seedlings, leaf area was evaluated for seedlings in three mature forest plots and four fa llows. When leaf number totaled 5, all observed leaves were sampled, which was the case for the majority of s eedlings. In all other cases, 5 leaves were sampled, representing the full range of leaf sizes To avoid destructive sampling, leaves were traced in the field, and average leaf area was measur ed in Gainesville, Florida, with a LiCor, Inc. Model 3000A Leaf Area Meter with Transparent Conveyor Belt Acce ssory (LiCor, Inc., Lincoln,

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25 Nebraska). Total leaf area was then estimated by multiplying total number of leaves per seedling by average leaf area. The influence of invertebrate herbivory on s eedling survival and growth was also assessed by quantifying the percentage of leaves per plant th at showed signs of attack in 2005. Vertebrate leaf herbivory was not assessed because it wa s not possible to observe or account for the significant portion of predation which is comprised of leaves that are completely removed by this type of herbivore. Data Analysis All statistical analyses were performed usi ng SAS software (Version 9.1, SAS Institute), excepting Students t-tests, which were pe rformed using SPSS software (Version 10.0, SPSS Inc.). Mature forest plots and fallows were tr eated as statistical bloc ks for all analyses, and seedling and sapling densities were determined on a per hectare basis. Average soil values were included as random effects, with pl ot nested within forest type, a nd soil nested within plot, for all analyses. PPFD values were log transformed prior to all analyses to improve normality and homoscedasticity of residuals, which were eval uated visually. PPFD levels were compared between forest types using a repeated measur es ANOVA, and soil nutrients were compared between forest types using a two-way ANOVA. All soil nutrient values were averaged on a plot basis, and average soil pH, potassium, and percen t organic carbon at two depths (0 20 cm and 20 40 cm) were included in explanatory dens ity and growth models. Correlation analysis, using the PROC CORR procedure, revealed hi gh correlations between the two depths for phosphorous and potential acidity (H+Al3+), therefore these soil nutri ent values were collapsed into one depth for inclusion in the explanatory models. Since B. excelsa densities demonstrated a Poisson dist ribution, they were modeled using a general linear mixed model a nd the GLIMMIX procedure. Two-way correlations were

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26 calculated between relevant variable pairs using PROC CORR in order to determine relationships which merited inclusion in the m odel. I explored model building by conventional methods of sequentially dropping no n-significant interactions and c ovariates based on p-values. In this case, I used a less restrictive significance level ( = 0.10) than the usual arbitrary 0.05 level (Johnson 1999, Burnham and Anderson 2002) to allow all possible significant effects to remain for the purpose of building the best ex planatory model (Bancroft 1968). Because a primary purpose of data analysis was to evaluate differences between matu re forest and fallow, the indicator variable for forest type was considered fundamental and, therefore, retained in the density models without regard to its significan ce level (Bancroft 1968). Densities were first compared for all plots, with forest type (mature ve rsus fallow) as the only retained fixed effect. Seedling and sapling densities of mature fore st plots previously surveyed in 2004 in PAE Chico Mendes (Serrano 2005) were included in the comparative density anal ysis to increase my sample size. These data were collected using th e same sampling scheme described herein, and it was assumed that no event had occurred that significantly affected these seedling and sapling densities in this mature forest site (PAE Chico Mendes) between 2004 and 2005. To test this assumption, 2004 seedling and sapling density data collected mature forest in RESEX Chico Mendes, using the same sampling scheme, were compared with my 2005 seedling and sapling density data from this same forest and plot s and no statistical diffe rences were detected (p = 0.57). Finally, no site (P AE Chico Mendes versus RESEX Chico Mendes) interaction was found when comparing B. excelsa densities; therefore, obser vations from both sites were included in the analysis, with site dropped as a fixed effect from the model. To explain observed densities, a subset of da ta (2 mature forest plots and 4 fallows) was analyzed. Tested fixed effects included forest type, number of reproductive adults within 50 m

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27 of each subplot, all plot soil content averages, and plot PPFD averages; insignificant variables were sequentially removed from the final model. All possible predictor variables were included in the final model to provide the best expl anatory model; however, re siduals were not well distributed. Students t-tests were used to compare initial height, initial basal diameter, height and basal diameter growth, number of leaves, total leaf area, and leaf herbivory between fallow and mature forest for all seedlings; all data were normally distributed. Sa pling diameter growth, which was likewise normally distributed, was also compared using a t-test. Logistic regression was used to analyze differences in seedling surviv al; tested fixed effects included initial height, initial diameter, height growth, diameter growt h, forest type, all plot soil content averages, PPFD, number of leaves per seedlin g, total leaf area, and percent leaf herbivory. Insignificant variables were sequentially removed from the final model. Initial seedling height and diameter were highly correlated with total leaf area, theref ore, interactions were explored between total leaf area and initial seedling size. The GLIMMIX procedure was used to test se edling and sapling growth. Growth values, some of which were negative, were log transfor med (log 10 [1+x]) prior to analyses to improve normality and homoscedasticity of residuals, wh ich were observed visually. Because data related to more mechanistic causes of growth (ie. light observed directly above seedlings) were available, the forest type variable was intentiona lly excluded from the seedling growth analysis. However, because these data were unavailable for saplings, forest type was retained for analysis in the sapling diameter growth model. Tested fixed effects included in initial seedling growth models were initial height or diameter, forest type, all plot soil content averages, PPFD, number of leaves per seedling, total leaf area, and percent leaf herbivor y, while initial sapling diameter

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28 growth models included initial dbh, forest type, all plot soil content averages, and plot PPFD averages Again, all possible predictor vari ables were included in the final models to provide the best explanatory models of growth, and insignifi cant variables were sequentially removed from the final model; residuals were fairly well dist ributed. Sapling height growth could not be compared between forest types, as sa pling heights were estimated visually. Differences in seedling leaf herbivory per centage were analyzed with the GLIMMIX procedure, accounting for a log-no rmal distribution of data. L eaf herbivory percentages were arcsin transformed prior to analysis in order to improve the normality of the response variable. Tested fixed effects included initia l height or diameter, forest type all plot soil content averages, PPFD, number of leaves per seed ling, total leaf area; insignifi cant variables were sequentially removed from the final model. Normality and homoscedasticity of residuals were observed visually and the residuals we re fairly well distributed. Results Regeneration Environments by Forest Type Of all variables measured to characterize re generation environments (plot light levels, density of nearby seed sources, te mperature and relative humidity, a nd soil nutrient stores), only light levels differed significan tly between the two forest type s. A repeated measures ANOVA revealed higher average PPFD levels in fallow th an mature forest (p = 0.02) (Fig. 2-2). There were no significant differences between forest types in number of adults within 50 m of evaluated subplots, nor in observe d daily temperature and percent relative humidity monitored in two fallows and one mature forest site between July 2005 and June 2006. Mean monthly temperature in the three sites ranged betw een 23.5 and 24.2C and mean relative humidity ranged between 90.3 and 92.7%. Although twoway ANOVAs revealed no soil property

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29 differences between forest types (Table 2-1), soil P was significantly different by depth (p = 0.0006). Seedling and Sapling Densities Observed mean seedling densities of 12.7 and 5.3 trees ha-1 in fallow and mature forest (Fig. 2-3), respectively, were diffe rent at p = 0.07, according to the general linear mixed model. Mean sapling densities of 5.2 and 1.3 trees ha-1 in fallow and mature forest (Fig. 2-3), respectively, were also different, again at p = 0.07. Based upon an analysis of the data subset, seedling density differences were somewhat expl ained by the number of a dults within 50m of each subplot (p = 0.06), such that seedling dens ities increased with number of adults, while sapling densities were best expl ained by percent organic carbon at the 0 20 cm depth (p = 0.02) (Table 2-2). B. excelsa Survival and Growth Seedlings After one year, 26% of surveyed seedlings in fa llow and 19% in mature forest (21% of all seedlings) were either dead or unable to be lo cated, with no survivorship differences between these two forest types (Table 23). Seedling survival was possibl y explained by total leaf area (p = 0.08) and an interaction betw een percent leaf herbivory an d PPFD (p = 0.08) (Table 2-2); improved survival was associated with gr eater leaf area and hi gher leaf herbivory. Initial seedling height was not significantly different betwee n forest types (Table 2-3), and, across forest types, seedling height in 2006 was best explained by PPFD measured in 2005 (p = 0.0009) (Fig. 2-4); seedling height increased with PPFD increase. Seedling height growth was best explained by an inter action between percent leaf herb ivory and PPFD (p = 0.02) and percent leaf herbivory alone (p = 0.02) (Table 2-2). Height growth increased with PPFD, and increased with herbivory decrease. Seedling grow th appeared to increase more with PPFD at

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30 intermediate herbivory levels. A t-test revealed no significant difference in initial seedling basal diameter between forest types (T able 2-3), however, basal diameter growth was greater in fallow than mature forest (p = 0.05), according to a t-test. According to the general linear mixed model, seedling basal diameter growth wa s best explained by total leaf ar ea (p = 0.05) (Table 2-2), such that seedlings with more leaf area grew mo re. Seedling height and diameter growth demonstrated positive relationships to all test ed factors except percent leaf herbivory. Total leaf area was highly correla ted with seedling height, diam eter and number of leaves and, as a result, less significant variables sequent ially fell out of the growth models. Total leaf area was strongly related to PPFD (p = 0.0004) (Fi g. 2-5). Observed herbivory level was much higher in mature forest than fallows (p < 0.0001) (Table 2-3). The mixed model corroborated that percent seedling leaf herbivory was explaine d by forest type alone (p = 0.05) (Table 2-2), but also showed that herbivory was best explai ned by an interaction between forest type and number of leaves (p = 0.006) (Table 2-2), such that herbivory decrease d with leaf number in mature forest, but increased w ith leaf number in fallow. Saplings Estimated mean sapling heights were 5.51 a nd 2.99 m, in mature forest and fallow, respectively, which was significan tly different (p = 0.02) (Table 2-3). Initial sapling dbh was also greater in mature forest than fallow (p = 0.05) (Table 2-3). Maximum dbh values were 9.5 cm in mature forest and 7.0 cm in fallow. N early 100% sapling survival was observed in both forest types (only one sapling died in a single fallow site), and th erefore no significant difference in sapling survival between forest types was de tected. Sapling dbh growth was much greater in fallow than mature forest (p = 0.02), and was poten tially explained only by forest type (p = 0.07) (Table 2-2).

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31 Discussion Densities Explained Bertholletia excelsa seedling densities were over two times greate r in anthropogenicallydisturbed fallows than mature fore st, and sapling densities were f our times greater in fallow (Fig. 2-3). Many factors influence esta blishment, survival, and growth of young individuals, including proximity and density of seed sources (Janzen 1 970), seed disperser and pr edator behavior (Sork 1987, Forget 1994, Molofsky and Fisher 1993, Asquith et al 1997), light availability (Augspurger 1984a, Augspurger 1984b, Poorter 1999) and nutrient availability (Ceccon et al. 2003), some of which may be related to microhabitat, or forest type. My data suggest that the influence of density and proximity of seed sour ces and light levels, at various stages in the B. excelsa recruitment process, play a significant role in shaping the observed densities of young B. excelsa individuals. Differences in densities betw een forest types, however, could also be a reflection of additional factors not measured, such as seed dispersal. These factors, which affect species establishment to differing degrees at vari ous stages in the recrui tment process, will be discussed below. Seed sources and dispersal Based on a deeper analysis of a subset of data, which included number of seed sources within 50m of surveyed subplots, seedling densi ties were mildly related to the number of nearby seed sources (p = 0.06). These findings are consis tent with scientific evidence that seedling densities decline with increased distance from parent tree s (Janzen 1970). Therefore, higher recruitment would be expected in fallows that either contain adult B. excelsa trees or are located adjacent to mature forests with high adult densi ties, compared to those with no adults nearby. Although density of nearby adults ex plained relative densities, seed lings and saplings were often found in areas with neither a repr oductive adult within 50m, nor tra ces of potential seed sources

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32 that may have suffered mortality prior to my study, confirming seed dispersal distances well beyond the typical dispersal maximum of 25m (Peres and Baider 1997). These dispersal distances may be explained by secondary dispersal (Mori and Prance 1990, Peres et al. 1997). Residuals in these explanatory density models were poorly distributed, with high density observations being grossly underestimated, sugges ting that relevant factors have not been included in the models. These underestimati ons could be due to a clumped, and possibly directed, seed dispersal pattern th at has not been accounted for. Bertholletia excelsa seed dispersal beyond the adult crown is determined nearly entirely by agouti scatterhoarding behavior (Prance and Mori 1978), and local extractivis t reports suggest th at agouti dispersal behavior may be non-random. Extractivists re port preferential seed dispersal in fallow environments (Clay 1997, personal obs.). Th ese accounts are corroborat ed by observations of seed burial at clearing edges (Peres and Baid er 1997) and in young seco ndary forest (Forget et al 2000), perhaps related to agouti predator avoi dance in areas composed of limb and vine tangles (Cintra and Horna 1997). Agouti scatterhoarding activity may potentially increase the proportion of seeds in favorable microenvironmen ts (Jansen and Zuidema 2001), which has been proposed as a more critical infl uence on regeneration than post-di spersal predation escape or successful germination (Rey and Alcntara 2000). The importance of disperser site preference and directed dispersal has been de monstrated for species other than B. excelsa (Uhl et al. 1981, Schupp et al. 1989, Forget 1994). Although disperser beha vior was not tested in this study, it likely had a significant effect on s eed availability for recruitment in fallow and mature forest. Equally important is the influence of differen tial seed removal rates (due to retrieval and predation) between different forest ty pes, which may further affect observed B. excelsa densities. Some authors report higher seed removal rates of vari ous tropical species in gap environments

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33 (Schupp 1988, Schupp and Frost 1989, Cintra and Horna 1997), while Sanchez-Cordero and Martinez-Gallardo (1998) report greater tropical s eed removal in mature forest than gaps. Tabarelli and Mantovani (1996), did not find seed removal of B. excelsa to be different between treefall gaps and undisturbed forest, however, cl earings created by anthropogenic disturbances were not assessed. It is clear that seed disperse r and seed predator activities are essential factors to include in future density evaluations. Seedling survival and growth Once seeds are dispersed and young seedlings establish, individuals may demonstrate differential growth (Augspurger 1984a, Po pma and Bongers 1991, Brown and Whitmore 1992, Poorter 1999) and/or survival (Augspurger 1984b, Cintra and Horna 1997, Brown and Whitmore 1992) in different forest types, which are often di fferentiated in the litera ture between understory and gap environments. In my study, however, fo rest type explained only sapling diameter growth (sapling height growth could not be eval uated). Other measured variables, some of which may be associated with forest type, demo nstrated stronger influences on observed survival and growth of B. excelsa seedlings. To begin with, total leaf area, which is correlated with leaf number, was a mild indicator of seedling survival (p = 0.08). Leaves fac ilitate the acquisition of carbon by means of photosynthesis; an enhanced carbon gain, due to greater leaf area, may confer relatively higher fitness (and ultimately survival) to seedlings by va rious means, including the provision of more structural material a nd energy, an increase in nutrient cont ent, and a better tolerance to environmental stresses (Chabot and Hicks 1982) PPFD, although not highly significant on its own, may contribute to increase d seedling survival, as PPFD was strongly positively correlated with total leaf area (p = 0.0004), which did expl ain seedling survival at p = 0.08. Although some studies have demonstrated greater seedling surviv al in gaps than understory for some species

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34 (Augspurger 1984b, Cintra and Horna 1997), my results do not dir ectly support these conclusions, as I found one-year B. excelsa survivorship to be no different between mature forest and fallow. Kainer et al (1998) also found no relationship between seedling survival and comparative light levels in pasture, fore st gap, and shifting cultivation sites (Kainer et al. 1998). The positive effect of leaf herbivory on survival is not so easily explaine d. Contrary to my findings, numerous studies demonstrate the negative effect of leaf herbivory on seedling survival (Howe 1990, Hulme 1996, Vasconcelos and Cherrett 1997). In this study, the influence of herbivory level on survival may be coincidental Invertebrate herbivory was not unusually high in either forest type, and is probably not sufficient to lead to seedling mortality under normal conditions. In addition, it was not possible to quantify vertebrate herbivory on seedling survival in this study. Nevertheless, it is likely that a greater percentage of mortality was represented by completely uprooted seedlings (Ortiz 1995, Zu idema 2003, personal obs.), resulting from seed predators unearthing and consuming seeds at the ba se of the plant. Furthermore, only 14 of 94 total seedlings died, therefor e concrete differences between factors relating to seedling survivorship were difficult to dete ct. Although forest type was not a significant factor in survival probability, in light of other anal yzed effects, it is possible that the forest type variable was partially masked by other variables such as pe rcent leaf herbivory or PPFD (due to multiple colinearity), which were both si gnificantly different by forest type (PPFD was higher in fallow and leaf herbivory was lower in fallow than mature fo rest). If forest type is partially included in one of these variables, it may play an undetected role in determining seedling survival probability. Seedling heights in 2006 were highly positively correlated with PPFD levels measured directly above seedlings in 2005, which implies a significant advantage to height growth in

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35 higher light environments. In my study, one-year seedling height growth was influenced by an interaction between PPFD and percent leaf herb ivory (p = 0.02), such that height growth declined with increased leaf herbivory and, while generally increasing wi th PPFD, was greatest at intermediate light levels. Previous studies have shown a positive relationship between seedling height growth and li ght availability in the tropics (Augspurger 1984a, Denslow et al 1990, Popma and Bongers 1991), and similar results have been reported for B. excelsa (Poorter 1999, Zuidema 2003), again, with highest growth rates at intermediate light levels. Uhl and Jordan (1984) found that soil nutrient concentrations did not vary significantly between agricultural sites and mature forest 5 years after cutting and burning. Similarly, I found no difference in soil nutrient availability between mature forest and fallows. As a result, soil nutrients did not explain B. excelsa survival and growth as well as other measured variables, such as light availability. Numerous studies have described the negativ e effects of herbivor y on seedling growth (Marquis 1984, Howe 1990, Hulme 1996, Vasconcel os and Cherrett 1997). In my study, seedling leaf area did not differ significantly by forest type, but was highly correlated with seedling height, and the two vari ables exhibited a significant in teraction effect for seedling height growth. Height growth was positively rela ted to both variables, similar to other studies which have demonstrated increased growth with greater initial height (Zuidema 2003) and leaf area (Poorter 1999). Similar to he ight growth, seedling diameter growth was explained by total leaf area (p = 0.05), and possibly by an interaction be tween leaf herbivory and PPFD (p = 0.07), with a positive relationship to each factor. Zuidema et al. (2003) report a similar relationship between B. excelsa biomass growth and light availabil ity. Although not directly evaluated, seedling survival and growth in my study sites ma y also have been influenced by ant presence.

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36 Leaf cutter ants ( Atta spp .) are known to modify soil nutrient concentrations (Lugo et al. 1973, Haines 1978), which may directly influence the development of young B. excelsa individuals, as well as influence the competitive environment with ot her species. In additi on, extractivists assert that leaf cutter ants preferentially predate both B. excelsa seeds and seedlings. The potential importance of leaf cutter ants in fallow environmen ts is worth particular attention as leaf-cutting ant populations have been shown to increase af ter forest clearing (Vas concelos and Cherrett 1995). Blanton and Ewels (1985) fi ndings suggest that leaf-cutting sp ecies continue to thrive as succession proceeds, representing the most importan t herbivore in the successional forests they studied. I found one of my six surveyed fallows was c overed to a great extent in leaf cutter ant nests (species unknown). The relationship between forest type and ant density, intensity of ant seed and seedling predation in fallow and mature forest, and the effects of ant populations on soil composition all merit further consideration, to be tter understand the role of ant populations in B. excelsa recruitment and regeneration. Sapling survival and growth High sapling survival rates in both mature forest and fallo w suggest that, once seedlings have been well-established, they develop and surv ive equally well in either forest type. Zuidema and Boot (2002) also found extremely low mort ality rates (1%), over a two-year period, for B. excelsa individuals > 1 cm dbh. Initial sapling diameter was greater in mature forest, but this can be explained by the young age of the fallows (5 12 years), as fallow saplings have not had the opportunity to attain dbh sizes near 10cm within this time frame. No factor included in statistical analyses, aside from forest type explained higher diameter grow th rates in fallow. However, saplings in fallow may also be able to invest more in biomass growth than mature forest saplings, since height growth is greatly e nhanced by higher light levels in fallow. Light is cited as the most frequently limiting resource for seedli ng growth (Kitajima 1996), a major component of

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37 tropical plant regenera tion processes (Chazdon et al. 1996). Increased light availability is likely one of the most significant advantages confe rred upon well-established seedlings in fallow, as observed PPFD was significantly grea ter in fallow than mature fore st (p = 0.02), and the benefits of light availability are likely great for individuals up to 10 cm dbh, which continue to compete to reach the canopy up to, and even beyond, this 10 cm limit. Secondary forests are, by nature, dynamic and, as a result, are associ ated with an everchanging microenvironment of vegetation, water, and soil nutrients. Therefor e, the suitability for seedling and sapling growth and survival may vary with forest successional stage. However, the observed higher densities of young indi viduals in fallow indicate that th is forest type, in general, constitutes a favorable microhabitat for bot h seedling and sapling development. Greater Ecological and Management Implications This research provides a new, and perhaps mo re accurate, depiction of current Brazil nut regeneration in tropical forest landscapes, as secondary forest environments have previously been ignored. The high densitie s of young Brazil nut individuals in secondary forests created through shifting cultivation (fallows) clearly dem onstrate that Brazil nut regeneration may have been underestimated in previous studies which we re conducted solely within mature forest. In light of these findings, fallows could pot entially be managed fo r enhanced Brazil nut densities, which may provide an opportunity fo r greater income for extractive families while contributing to the sustainability of Brazil nut extraction in the long term. In the past, some extractivists have experimented with small-scal e Brazil nut enrichment plantings, as a means to economically enrich thei r landholdings (Kainer et al. 1998). The benefits of fallow management may also provide an additional incentive for extractivists to allow fallows to regenerate, rather than convert them to pasture, which is an incr easingly common practice in much of the Amazon.

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38 Shifting cultivation has previously been rec ognized for its role in shaping Amazonian ecosystems (Dufour 1990). Pre-Columbian human in terventions, in the form of active planting, were previously considered by Posey (1985) and Bale (1989) as a possible explanation for the presence of Brazil nut groves in parts of the Amazon. Based on the results of this study, however, it is possible that higher de nsities are a result of less di rect human interventions, such as shifting cultivation. Not only can anthropogen ic disturbance, in the form of shifting cultivation, play a positive role in Brazil nut regeneration, it could explain current Brazil nut densities and distributions.

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39 CHAPTER 3 CONCLUSION The objective of this study was to compare B. excelsa recruitment and regeneration in regenerating swidden fallows and mature forest. My results suggest th at fallows constitute favorable microhabitats for both seedling and sapl ing development. Perhaps most importantly, survival and growth of young B. excelsa individuals appear to be enhanced by the higher light levels found in fallow. Greater light availability is positively related to seedling height and height growth, seedling leaf number and leaf ar ea, and seedling survival, and may indirectly enhance sapling growth and survival. This enha nced survival and growth likely contributes to the observed higher densities of seedlings and saplings in fallow sites. At the conclusion of this study it became clear that seed dispersal and seed predation should be compared between forest types in future regeneration studies, in order to better assess the various recruitment advantages, and possi bly disadvantages, associated with fallow environments. In addition, herbi vore activity, and its effects on B. excelsa survival and growth, should be evaluated in greater detail, quantif ying the proportion of herb ivory exercised by both vertebrates and invertebrates, in both forest types. Leaf cutte r ant behavior in regenerating fallow fields merits particul ar consideration. Future studies should also compare B. excelsa regeneration between different secondary forest types, as well as between fallows of varying ages, as various successional forest types and suc cessional stages may contribute to differential survival and growth of young individuals. At th e time of thesis submission, soil analyses were incomplete, therefore conclusions regarding the effects of so il content were limited. Upon publication of Chapter 3 in a scie ntific journal, the incorporati on of additional soil data should yield further insights.

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40 Although some significant factors in B. excelsa recruitment processes, such as disperser activity, were not evaluated in the study, th e importance of the results should not be underestimated. My research focuses on a widespr ead forest type which ha s not previously been included in assessments of B. excelsa population dynamics therefore conclusions regarding regeneration in fallows may better explain current B. excelsa densities and distributions than other studies to date. An understa nding of the role of fallows in B. excelsa regeneration may also provide insight into the regenera tion requirements of other valuab le gap-loving species, such as those in the Cedrela and Swietenia genera

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41 Figure 2-1: Location of study sites PAE Chico Mendes and RESEX Chico Mendes in Acre, Brazil (Figure adapted from Go mes 2001, with permission). 0 0.5 1 1.5 2 2.5 7:3010:0012:3015:00 TimeMean PPFD ( molm-1s-1) Fallow Mature Figure 2-2: Mean ( SE) PPFD at four times of da y in four fallows and two mature forest plots Data are log transformed.

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42 0 2 4 6 8 10 12 14 16 18 SeedlingsSaplings Size ClassDensity (trees ha-1) Fallow Mature Forest Figure 2-3: Bertholletia excelsa seedling and sapling densitie s (means SE) in fallow and mature forest. 0 0.5 1 1.5 2 2.5 050100150200250PPFD ( mol*m-1*s-1)2006 Seedling height (m) Figure 2-4: Relationship between B. excelsa seedling height and photon flux density (PPFD) measured in 2005. Each point represents one observed B. excelsa seedling. Regression line fitted: y = 0.005x + 0.52 R2 = 0.31.

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43 0 500 1000 1500 2000 2500 3000 3500 4000 050100150200250PPFD ( mol*m-1*s-1)Seedling Total Leaf Area (cm2) Figure 2-5: Relationship between B. excelsa seedling total leaf ar ea and photon flux density (PPFD) measured in 2005. Each point represents one observed B. excelsa seedling. Regression line fitted: y = 7.93x + 294.97, R2 = 0.23. Table 2-1: Mean ( sd) soil pr operties observed with in fallows and mature forest plots in RESEX Chico Mendes and PAE Chico Mendes. Soil property Fallow Mature Forest pH 5.5 ( 0.2) 5.1 ( 0.3) K (cmol*kg-1) 0.4 ( 0.03) 0.3 ( 0.1) P (mg*kg-1) 2.0 ( 0.3) 2.7 ( 0.4) Potential Acidity (H+Al3+) (cmol*kg-1) 4.0 ( 0.7) 5.7 ( 1.0) Organic Carbon (%) 0.5 ( 0.1) 0.6 ( 0.1)

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44 Table 2-2: Mixed model results based on B. excelsa seedling and sapling data subsets. Model fixed effects F-statistic p-value Seedlings Density (ha-1) Forest type 1.09 0.33 Number of adults within 50m of subplots 3.71 0.06 Survival Total leaf area 3.22 0.08 Percent leaf herbivory*PPFD 3.26 0.08 Height growth (cm) Initial height 4.99 0.08 Percent leaf herbivory 13.09 0.02 PPFD*Percent leaf herbivory 11.93 0.02 Potential acidity (0 40cm) 7.55 0.04 Soil K (0 20cm) 6.21 0.06 Basal diameter growth (cm) Total leaf area 4.26 0.05 Percent leaf herbivory 3.42 0.08 Percent leaf herbivory*PPFD 3.58 0.07 Percent leaf herbivory Forest type 6.96 0.05 Number of leaves*forest type 5.84 0.006 Saplings Density Forest type 1.80 0.23 Percent organic carbon 9.25 0.02 Growth (dbh) Forest type 4.55 0.07

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45 Table 2-3: Comparison of B. excelsa seedling and sapling characteristics in fallow and mature forest (all p-values based upon t-tests, ex cept one-year survivorsh ip, which is based upon logistic regression). Characteristic N Fallow Mature Forest p-value Seedlings Initial seedling height (m) 76 0.67 ( 0.42) 0.59 ( 0.38) 0.45 Initial seedling basal diameter (cm) 76 0.64 ( 0.34) 0.55 ( 0.21) 0.15 Seedling height growth (m) 62 0.14 ( 0.28) 0.12 (0.22) 0.82 Seedling basal diameter growth (cm) 62 0.19 ( 0.25) 0.10 (0.11) 0.05 Number of leaves per seedling 75 12.8 ( 9.4) 9.7 (7.6) 0.13 Total leaf area (cm2) 68 1003.4 ( 915.4) 710.5 ( 807.2) 0.20 Leaf herbivory (%) 74 16 ( 24) 58 ( 28) < 0.0001 One-year survivorship (%) 66 74 81 0.79 Saplings Initial height (m) 31 2.99 ( 1.95) 5.51 ( 3.74) 0.02 Initial dbh (cm) 31 2.23 ( 2.07) 4.45 ( 3.32) 0.05 Dbh growth (cm) 25 0.47 ( 0.39) 0.11 (0.32) 0.02 Data reported as means ( sd), excepting percentage data. Reporte d sapling heights are estimated.

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46 APPENDIX A TRANSECT CREATION To open transects, a compass was placed on a level-cut branch. Ne xt, two sapling poles where placed in the line of sight of the compass to begin a straight line. Field technicians and local assistants referred to this line in order to open a straight transect through the forest with a machete. Reference poles were placed no more th an every 15 m apart, with poles placed closer together in hilly or overgrown areas. Distance was measured with a metric tape to create entire 300 m transects. Each transect was re-measured with metric tape after clearing was completed. This method facilitated searching and locating B. excelsa individuals over large areas.

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47 APPENDIX B SCHEMATIC OF PLOT INSTALLATION AND SAMPLING SCHEME A B Figure B-1: Mature forest and fallo w plot design. A) Nine ha plot of mature forest, with four 25 x 25m subplots randomly selected within each ha, for a total of 36 subplots B) 1.1 ha fallow with 27 10 x 10m subplots. Distan ce is indicated in meters. represents recruitment subplot. Light environments within two mature forest plots a nd four fallows were ch aracterized by photon flux density (PPFD) measurements; represents PPFD measurement. Soil samples within each fallow and mature forest plot were collected belo w the litter layer at two depths (0-20 cm and 2040 cm). Sampling scheme is depicted above for each forest type; represents soil sample.

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48 LIST OF REFERENCES Asquith, N.M., Wright, S.J., Clauss, M.J., 1997. Does mammal community composition control recruitment in Neotropical forests? Ev idence from Panama. Ecology 78(3), 941-946. Augspurger, C.K., 1984a. Light requirements of neotropical tree seedlings: A comparative study of growth and survival. J. Ecol. 72(3), 777-795. Augspurger, C.K., 1984b. Seedling survival of tr opical tree species : Interactions of dispersal distance, light gaps, and pa thogens. Ecology 65(6), 1705-1712. Bale, W., 1989. The culture of Amazonian forests. In: Posey, D.A., Bale, W. (Eds.), Resource Management in Amazonia: Indigenous and Fo lk Strategies. Advances in Economic Botany No. 7, New York Botanical Ga rden, New York, NY, pp. 1-21. Bancroft, T.A., 1968. Topics in intermediate sta tistical methods. Iowa State University Press, Ames, Iowa, USA. Blanton, C.M., Ewel, J.J., 1985. Leaf-cutting ant herbivory in successional and agricultural tropical ecosystems. Ecology 66(3), 861-869. Borges, S.H., Stouffer, P.C., 1999. Bird communitie s in two types of anthropogenic successional vegetation in Central Amazonia. The Condor 101, 529-536. Brokaw, N.V.L., 1982. The definition of treefall gap and it s effect on measures of forest dynamics. Biotropica 14(2), 158-160. Brokaw, N. V. L., 1985. Gap-phase regeneration in a tropical forest. Ecology 66(3), 682-687. Brown, N.D., Whitmore, T.C., 1992. Do dipterocarp seedlings really partition tropical rain forest gaps? Phil. Trans: Biol Sci. 335(1275), 369-378. Burnham, K.P., Anderson, D.R., 2002. Model sel ection and multi-model in ference: a practical information-theoretic approach. Second edit ion. Springer-Verlag, New York, New York, USA. Ceccon, E., Huante, P., Campo, J., 2003. Effects of nitrogen and phosphorus fertilization on the survival and recruitment of seedlings of do minant tree species in two abandoned tropical dry forests in Yucatn, Mexico. For. Ecol. Manage. 182(1), 287-402. Chabot, B.F., Hicks, D.J., 1982. The ecology of leaf life spans. Ann. Rev. Ecol. Syst. 13, 229259. Chazdon, R.L., Pearcy, R.W., Lee, D.W., Fetcher, N., 1996. Photosynthetic responses of tropical forest plants to contrasti ng light environments. In: Mulkey, S.S., Chazdon, R.L., Smith, A.P. (Eds.), Tropical Forest Plant Ecophysio logy Chapman and Hall, New York, NY, pp. 5-55.

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49 Cintra, R., Horna, V., 1997. Seed and seedling survival of the palm Astrocaryum murumuru and the legume Dipteryx micrantha in gaps in Amazonian forest. J. Trop. Ecol. 13(2), 257277. Clay, J. W., 1997. Brazil nuts: the use of a keysto ne species for conservation and development. In: Freese, C.H. (Ed.), Harvesting Wild Species: Implications for Biodiversity Conservation. The John Hopkins Univer sity Press: Baltimore, MD, pp. 246-282. Da Silva, J.M.C., Uhl, C., Murray, G., 1996. Pl ant succession, landscape management, and the ecology of frugivorous birds in abandoned Am azonian pastures. Cons. Biol. 10(2), 491503. Denevan, W.M., 1992. The pristine myth: The landscap e of the Americas in 1492. Annals of the Association of American Geographers 82(3), 369-385. Denslow, J.S., Schultz, J.C., Vitousek, P.M., Strain, B.R., 1990. Growth response of tropical shrubs to treefall gap envi ronments. Ecology 71(1), 165-179. Dufour, D.L., 1990. Use of tropical rainforest s by Native Amazonians. BioScience 40(9), 652659. EMBRAPA (Empresa Brasileira de Pesquisa Agropecuria), 1997. Manual de Mtodos de Anlise de Solos. Segunda edio. Centro Naci onal de Pesquisa de Solos, Rio de Janeiro, Brazil. Ferguson, B.G., Vandermeer, J., Morales, H., Gr iffith, D.M., 2003. Post-agricultural succession in El Petn, Guatemala. Cons. Biol. 17(3), 818-828. Forget, P-M., 1994. Recruitment pattern of Voucapoua americana (Caesalpiniaceae), a rodentdispersed tree species in Fren ch Guiana. Biotropica 26, 408-419. Forget, P-M., Milleron, T., Feer, F., Henry, O., D ubost, G., 2000. Effects of dispersal pattern and mammalian herbivores on seedling recruitment for Virola michelii (Myristicaceae) in French Guiana. Biotropica 32(3), 452-462. Fujisaka, S., Escobar, G., Veneklaas, E., 1998. Pl ant community diversity relative to human land uses in an Amazon forest colony. Bi odiversity and Conservation 7, 41-57. Gomes, C.V.A., 2001. Dynamics of land use in an Amazonian Extractive Reserve: The Case of the Chico Mendes Extractive Reserve in Acre, Brazil. Thesis, University of Florida, Gainesville, Florida. Guariguata, M.R., Ostertag, R ., 2001. Neotropical secondary forest succession: changes in structural and functional characteri stics. For. Ecol. Manage. 148, 185-206. Haines, B. L., 1978. Element and energy flows through colonies of the leaf-cutting ant, Atta colombica in Panama. Biotropica 10(4), 270-277.

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50 Heckenberger, M.J., Kuikuro, A., Kuikuro, U.T ., Russell, J.C., Schmidt, M., Fausto, C. Franchetto, B., 2003. Amazonia 1492: Pristine fore st or cultural pa rkland? Science 301 (5640), 1710-1714. Holdridge, L.R., 1978. Ecologa basada en zonas de vida. Instituto Interamericano de Ciencias Agrcolas, San Jos, Costa Rica. Howe, H.F., 1990. Survival and growth of juvenile Virola surinamensis in Panama: Effects of herbivory and canopy closure. J. Trop. Ecol. 6(3), 259-280. Hulme, P.E., 1996. Herbivores and the performa nce of grassland plants: A comparison of arthropod, mollusc and rodent he rbivory. J. Ecol. 84(1), 43-51. Huber, J., 1910. Mattas e madeiras amaznicas Bol. Mus. Paraese Hist. Nat. 6, 91-225. IMAC Instituto de Meio Ambiente do Acre 1991. Atlas geogrfico ambiental do Acre. IMAC, Rio Branco, Acre, Brasil. ITTO, 2002. ITTO guidelines for the restoration, management, and rehabilitation of degraded and secondary tropical forests. ITTO Po licy Development Series No.13. International Tropical Timber Organization. Jansen, P.A., 2003. Scatterhoarding and tree re generation: Ecology of nut dispersal in a neotropical rainforest. Ph D Thesis, Wageningen University, The Netherlands. Jansen, P.A., Zuidema, P.A., 2001. Logging, seed dispersal by vertebra tes, and the natural regeneration of tropical timber trees. In: Fimbel, R.A., Robinson, J.G., Grajal, A. (Eds). The cutting edge. Conserving wildlife in logge d tropical forests. Columbia University Press, New York, pp. 35-59. Janzen, D.H., 1970. Herbivores and the number of tr ee species in tropical forests. The American Naturalist. 104 (940), 501-528. Janzen, D.H., 1990. An abandoned field is not a tree fall gap. Vida Silvestre Neotropical 2(2), 64-67. Johnson, D.H., 1999. The insignificance of statistical significance testing. Journal of Wildlife Management 63(3), 763-772. Kainer, K.A., Duryea, M.L., de Macedo, N. C., Williams, K., 1998. Brazil nut seedling establishment and autecology in extractive reserves of Ac re, Brazil. Ecol. Appl. 8, 397410. Kitajima, K., 1996. Ecophysiology of tropical tree seedlings. In: Mulkey, S.S., Chazdon, R.L., Smith, A.P. (Eds.), Tropical Forest Plan t Ecophysiology Chapman and Hall, New York, NY, pp. 559-592.

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51 Lugo, A.E., Farnworth, E.G., Pool, D., Jerez, P., Kaufman, G., 1973. The impact of the leaf cutter ant Atta colombica on the energy flow of a tropi cal wet forest. Ecology 54(6), 1292 1301. Marquis, R.J., 1984. Leaf herbivores decrease fi tness of a tropical plant. Science 226 (4674), 537-539. Mesquita, R.C.G., 1995. The effects of diffe rent proportions of canopy opening on the carbon cycle of a central Amazonian secondary forest. Ph D. thesis, University of Georgia, Athens. Mesquita, R.C.G., Ickes, K., Ganade, G., Williamson, G.B., 2001. Alternative successional pathways in the Amazon Basin. J. Ecol. 89, 528. Molofsky, J., Fisher, B.L., 1993. Habitat and predati on effects on seedling survival and growth in shade-tolerant tropical trees. Ecology 74(1), 261-265. Montagnini, F., Mendelsohn, R. O., 1997. Managing forest fallows : Improving the economics of swidden agriculture. Ambio 26(2), 118-123. Mori, S.A., Prance, G.T., 1990. Taxonomy, ecol ogy, and economic botany of the Brazil nut ( Bertholletia excelsa Humb. and Bonpl.: Lecythidaceae). Adv. Econ. Bot. 8, 130-150. Murphy, L.J., Riley, J.P., 1962. A modified sing le solution method for determination of phosphate in natural waters. Anal. Chim. Acta 27, 31. Myers, G.P., Newton, A.C., Melgarejo, O., 2000. The influence of canopy gap size on natural regeneration of Brazil nut ( Bertholletia excelsa ) in Bolivia. For. Ecol. Manage. 127, 119128. Nascimento, H.E.M., Andrade, A.C.S., Camargo, J.L.C., Laurance, W.F., Laurance, S.G., Ribeiro, J.E.L., 2006. Effects of the surrounding matrix on tree recruitment in Amazonian forest fragments. Conser vation Biology 20(3), 853-860. Nations, J.D., Nigh, R.B., 1980. The evolutionary potential of Lacandon Ma ya sustained-yield tropical forest agriculture. Journa l of Anthropological Research 36, 1-30. Ortiz, E.G., 1995. Survival in a nutshell. Amricas 47, 7. Pea-Claros, M., De Boo, H., 2002. The effect of forest successional stag e on seed removal of tropical rain forest tree sp ecies. J. Trop. Ecol. 18, 261. Peres, C. A., Baider, C., 1997. Seed dispersal, spatial distribution and population structure of Brazil nut trees ( Bertholletia excelsa ) in Southeastern Amazoni a. J. Trop. Ecol. 13(4), 595616.

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52 Peres, C.A., Baider, C., Zuidema, P.A., Wadt L.H.O., Kainer, K.A., Gomes-Silva, D.A.P., Salomo, R.P., Simes, L.L., Franciosi, E.R. N., Valverde, F.C., Gribel, R., Shepard Jr., G.H., Kanashiro, M., Coventry, P., Yu, D.W ., Watkinson, A.R., Freckleton, R.P., 2003. Demographic threats to the sustainability of Brazil nut exploitation. Science 302, 21122114. Peres, C.A., Schiesari, L.C., Dias-Leme, C. L., 1997. Vertebrate pred ation of Brazil-nuts ( Bertholletia excelsa Lecythidaceae), an agouti-disperse d Amazonian seed crop: A test of the escape hypothesis. J. Trop. Ecol. 13(1), 69-79. Poorter, L., 1999. Growth responses of 15 rain-fores t tree species to a ligh t gradient: The relative importance of morphological and physiol ogical traits. Func Ecol. 13(3), 396-410. Popma, J., Bongers, F., 1991. Acclim ation of seedlings of three Mexi can tropical rain forest tree species to a change in light avai lability. J. Trop. Ecology 7(1), 85-97. Posey, D.A., 1985. Indigenous management of tropical forest ecosystems: The case of the Kayap Indians of the Brazilian Amazon. Agroforestry Systems 3, 139-158. Prance, G.T., Mori, S.A., 1978. Observations on the fruits and seeds of neotropical Lecythidaceae. Brittonia 30(1), 21-23. Rey, P.J., Alcntara, J.M., 2000. Recruitmen t dynamics of a fleshy-fruited plant ( Olea europaea ): Connecting patterns of seed dispersal to seedling establishment. J. Ecol., 88(4), 622-633. Saldarriaga, J.G., West, D.C., Tharp, M.L., Uh l, C., 1988. Long-term chronosequence of forest succession in the upper Rio Negro of Colomb ia and Venezuela. J. Ecology 76, 938-958. Sanchez-Cordero, V., Martinez-Gallardo, R., 1998. Postdispersal fruit and seed removal by forest-dwelling rodents in a lowland rainfore st in Mexico. J. Trop. Ecol. 14(2), 139-151. SAS Institute, Inc., 2004. SAS. Version 9.1. SAS, Cary, North Carolina, USA. Schupp, E.W., 1988. Seed and early seedling predation in the forest unders tory and in treefall gaps. Oikos 51, 71-78. Schupp, E.W., Frost, E.J., 1989. Differential predation of Welfia georgii seeds in treefall gaps and the forest understory. Biotropica 21(3), 200-203. Schupp, E.W., Howe, H.F., Augspurger, C.K., Leve y, D.J., 1989. Arrival and survival in tropical treefall gaps. Ecol ogy 70(3), 562-564. Serrano, R.O.P., 2005. Regenerao e estrutura populacional de Bertholletia excelsa (H.B.K.) em reas com diferentes histricos de ocupa o, no Vale do Rio Acre (Brasil). Dissertao, Universidade Federal do Acre, Acre, Brasil.

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53 Sork, V.L., 1987. Effects of predation and light on seedling establishment in Gustavia superba Ecology 68(5), 1341-1350. SPSS Inc., 1999. SPSS Version 10.0. SPSS, Inc., Chicago, IL. Tabarelli, M., Mantovani W., 1996. Remoo de sementes de Bertholletia excelsa (Lecythidaceae) por animais em uma floresta de terra firme na Amaznia Central, Brasil. Revista Brasileira de Biologia 56(4), 755-760. Uhl, C., Buschbacher, R., Serro, E. A. S., 19 88. Abandoned pastures in Eastern Amazonia. I. patterns of plant succession. J. Ecol. 76, 663-681. Uhl, C., Clark, K., Clark, H., Murphy, P., 1981. Early plant succession after cutting and burning in the upper Rio Negro region of the Amazon Basin. J. Ecol. 69(2), 631-649. Uhl, C. Clark, H., Clark, K., Maquirino, P., 1982. Su ccessional patterns associated with slashand-burn agriculture in the upper Rio Negro re gion of the Amazon Basin. Biotropica 14(4), 249-254. Uhl, C., Jordan, C., 1984. Succession and nutrien t dynamics following forest cutting and burning in Amazonia. Ecology 65(5), 1476-1490. Vasconcelos, H.L., Cherrett, J.M., 1995. Changes in leaf-cutting ant populations (Formicidae: Attini) after the clearing of mature forest in Brazilian Amazonia. Studies on Neotropical Fauna and Environment 30, 107-113. Vasconcelos, H.L., Cherrett, J.M., 1997. Leaf-cut ting ants and early fore st regeneration in Central Amazonia: Effects of herbivory on tree seedling establishment. J. Trop. Ecol. 13(3) 357-370. Viana, V.M., Mello, R.A., Moraes, L.M., Mende s, N.T., 1998. Ecologia e manejo de populaes de castanha-do-Par em reservas extrativ istas Xapur, Estado do Acre. In: Gascon, C., Moutinho, P. (Eds.), Floresta Amaznica: Di nmica, Regenerao e Manejo. Instituto Nacional de Pesquisa Amazni a, Manaus, Brazil, pp. 277-292. Vieira, S., Trumbore, S., Camargo, P.B., Selhorst D., Chambers, J.Q., Higuchi, N., Martinelli, L.A., 2005. Slow growth rates of Amazonian trees: Consequences for carbon cycling. Proceedings of the National Academy of Sciences 102, 18502-18507. Wadt, L.H.O., Kainer, K.A., Gomes-Silva, D.A. P., 2005. Population structur e and nut yield of a Bertholletia excelsa stand in Southwestern Amazonia. For. Ecol. Manage. 211, 371-384. Whitmore, T.C., 1989. Canopy gaps and the two majo r groups of forest trees. Ecology 70(3), 536-538. ZEE Zoneamento Ecolgico-Econmico do Acre 2000. Recursos Naturais e Meio Ambiente, vol. I, Secretaria de Estado de Cincia, T ecnologia e Meio Ambiente, Rio Branco, Acre, Brasil.

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54 Zuidema, P.A., 2003. Ecology and Mana gement of the Brazil nut tree ( Bertholletia excelsa ). PROMAB Scientific Series 6, PROMAB, Utrecht, Netherlands. Zuidema, P.A., Boot, R.G.A., 2002. De mography of the Brazil nut tree ( Bertholletia excelsa ) in the Bolivian Amazon: Impact of seed extr action on recruitment and population dynamics. J. Trop. Ecol. 18, 1-31

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55 BIOGRAPHICAL SKETCH Jamie Nicole Cotta was born and raised in Rive rside, California, and began her career in tropical conservation at the University of Califor nia, San Diego, where she received a B.S. in Ecology, Behavior and Evolution. During her studies she conduc ted behavior observations of bonobo chimpanzees ( Pan paniscus ), at the San Diego Zoo, and pygmy lorises ( Nycticebus pygmaeus ), at the Center for Reproduction of Endangere d Species. She next assisted a Cornell post-doc with behavioral research on orange-fronted parakeets ( Aratinga canicularis ) in Santa Rosa National Park, Costa Rica. Her undergradu ate studies concluded with a semester of Tropical Biology in Monteverde, Costa Rica, where she first became interested in sustainable tropical forest management. She is most proud of her work as a research assistant for the School for Field Studies, in Yungaburra, Queensland, Australia, where she mana ged a tropical plant nursery and helped to design a restoration pl anting with a local community tree planting organization (TREAT). Her tropical research experiences fostered an interest in tropical ecology and forest restoration in the tr opics, leading her to pursue a Masters of Science degree in Interdisciplinary Ecology at the University of Fl orida. Upon graduation she plans to apply her experience and education with in an NGO setting, helping businesses and communities adopt more sustainable practices and lifestyles.


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Title: Shifting Cultivation Effects on Brazil Nut (Bertholletia excelsa) Regeneration
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Copyright Date: 2008

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Material Information

Title: Shifting Cultivation Effects on Brazil Nut (Bertholletia excelsa) Regeneration
Physical Description: Mixed Material
Copyright Date: 2008

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Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
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SHIFTING CULTIVATION EFFECTS ON BRAZIL NUT (Bertholletia excelsa)
REGENERATION





















By

JAMIE NICOLE COTTA


A THESIS PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
MASTER OF SCIENCE

UNIVERSITY OF FLORIDA

2007

































Copyright 2007

by

Jamie Nicole Cotta





















































To my parents, who made it all possible.










3









ACKNOWLEDGMENTS

This thesis could not have been completed without the steadfast support of my ever

encouraging advisor, Dr. Karen Kainer, who generously shared her expertise and provided

counsel throughout the research and writing processes. I am equally grateful to my committee

member, Dr. Lucia Wadt, who welcomed me into her home in Brazil three years in a row,

offering not only research advice, but personal and professional guidance that contributed to the

success of my project in Brazil. I also thank the members of my supervisory committee: Dr.

Emilio Bruna and Dr. Kaoru Kitajima, for their comments and suggestions during the writing of

my thesis. I am immensely thankful for Christine Staudhammer's statistical counsel; she has a

true gift for communicating her statistical expertise with forestry professionals. I also thank

Meghan Brennan for sharing her statistical knowledge.

I sincerely thank my international sponsor, EMBRAPA-Acre (Empresa Brasileira de

Pesquisa Agropecudria) in Rio Branco, Brazil, for generously supporting this research. A

number of Brazilian colleagues and assistants provided essential insights and offered their talents

to this research project, including Paulo Wadt, Paulo Rodrigues de Carvalho, Pedro Raimundo R.

de Araujo, Aldeci da Silva Oliveira, Airton, Freire, Sergio, and Rivelino. I would like to

sincerely thank the extractivist families, specifically Valderi, Maria Alzenira, Duda and Bilu

Mendes, and Amarilzo da Rocha Bento, for generously sharing their insights, and providing me

with a true home away from home. Their joyful, affectionate children will also never be

forgotten. I would also like to thank Tim Martin, Christie Klimas, Cara Rockwell, Chris

Baraloto, Marisa Tohver, Roberta Veluci, Alexander Cheesman, Skya Rose Murphy, Shoana

Humphries, Valerio Gomes, and Marco Lentini for their contributions to my final draft.

This research would not have been possible without the support of my Department, the

School of Natural Resources and Environment, and a generous scholarship from the School of









Forest Resources and Conservation. I am also grateful for summer research grants from the

University of Florida Graduate School and the Tropical Conservation and Development

Program. The Institute of Food and Agricultural Sciences, the School of Natural Resources and

Environment, and the UF Graduate Student Council also generously provided me with finds to

return research results to communities.

Most importantly, I thank my family. My parents and sister represent the most admirable

of role models and, through their guidance and living examples, I have learned how to face great

challenges and accomplish personal goals, while never losing focus of others. Because of them I

understand true compassion and generosity of spirit. My grandparents, aunts, uncles, and

cousins have each shared a special part of themselves, which have all contributed to my success

and, without their presence in my life I would not have been able to pursue and complete this

degree.









TABLE OF CONTENTS

page

A CK N O W LED G M EN T S ................................................................. ........... ............. .....

L IST O F T A B L E S ............. ..... ............ ................................................................... . 8

LIST OF FIGURES .................................. .. ..... ..... ................. .9

A B S T R A C T ................................ ............................................................ 10

CHAPTER

1 IN T R O D U C T IO N ....................................................................................... .................... 12

2 SHIFTING CULTIVATION EFFECTS ON BRAZIL NUT (Bertholletia excelsa)
REGENERATION ................................. .. ... .... ................... 17

Intro du action ............... ......................... .. ............................... ......................................17
Species Description .................................. ... .. .... ...... ................. 19
Study Area ........................................ 20
M methods ........................ .. ....................................................... 21
Plot Installation and Sampling Scheme........................................................... 21
Regeneration Environments by Forest Type........................................................22
Seedling and Sapling D ensities ............................................... ............................ 24
B excelsa Survival and G row th ........................................................... .....................24
D ata A n a ly sis ............................................................................................................. 2 5
Results .............................. .................. ............. 28
Regeneration Environments by Forest Type ........................................................28
Seedling and Sapling D ensities ............................................... ............................ 29
B excels Survival and G row th ........................................................... .....................29
S e e d lin g s ................................................................2 9
S a p lin g s ................................................................3 0
D iscu ssio n ................... ................... ...................1..........
D en site s E x p lain ed ................................................................................................ 3 1
Seed sources and dispersal ................................. ................................. 31
Seedling survival and growth ................................. .............................33
Sapling survival and growth.....................................36
Greater Ecological and Management Implications ........................................ ...37

3 C O N C L U S IO N ................................................................................................................. 3 9

APPENDIX

A T R A N SE C T C R E A T IO N ................................................................................................ 46

B SCHEMATIC OF PLOT INSTALLATION AND SAMPLING SCHEME ..........................47









L IST O F R E F E R E N C E S ............................................................................... ...........................48

B IO G R A PH IC A L SK E T C H ............................................................................... .....................55









LIST OF TABLES


Table page

2-1 Mean ( sd) soil properties observed within fallows and mature forest plots in
RESEX Chico Mendes and PAE Chico Mendes. ............. ........................................43

2-2 Mixed model results based on B. excelsa seedling and sapling data subsets. ..................44

2-3 Comparison ofB. excelsa seedling and sapling characteristics in fallow and mature
fo re st ........................................................................................ . 4 5









LIST OF FIGURES


Figure page

2-1 Location of study sites PAE Chico Mendes and RESEX Chico Mendes in Acre,
B razil ............. .......... .. ......... ............... ................................... 4 1

2-2 Mean ( SE) PPFD at four times of day in four fallows and two mature forest plots.......41

2-3 Bertholletia excelsa seedling and sapling densities (means SE) in fallow and
m atu re forest. ........................................................... ................. 42

2-4 Relationship between B. excelsa seedling height and photon flux density (PPFD)
m e a su re d in 2 0 0 5 ...............................................................................................................4 2

2-5 Relationship between B. excelsa seedling total leaf area and photon flux density
(PPFD ) m measured in 2005 .......................................................... ........ .......43

B-1 M ature forest and fallow plot design ........................................ .......................... 47









Abstract of Thesis Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Master of Science

SHIFTING CULTIVATION EFFECTS ON BRAZIL NUT (Bertholletia excelsa)
REGENERATION

By

Jamie N. Cotta

May 2007

Chair: Karen Kainer
Major Department: School of Natural Resources and Environment

Brazil nut (Bertholletia excelsa), has emerged as the cornerstone of the extractive economy

in much of the Amazon, but the debate continues as to whether or not current harvest levels have

a detrimental effect on Brazil nut seedling recruitment. Regeneration studies to date have been

conducted solely within mature forest, but my study provides further insight into current Brazil

nut regeneration dynamics, with a unique first look at regeneration in swidden fallows within

two multiple-use areas in Acre, Brazil. Recruitment of individuals < 10 cm dbh was evaluated

within 25 x 25 m and 10 x 10 m subplots in four 9 ha mature forest plots and six fallows (0.5-1.0

ha each), respectively. Individuals < 10 cm dbh were grouped into two size classes: seedlings

(individuals < 1.5 m in height) and saplings (individuals between 1.5 m in height and 10 cm

dbh). General linear mixed model analyses revealed higher densities ofB. excelsa in fallow than

mature forest. Survival and growth of young B. excelsa individuals appear to be enhanced by

higher light levels found in fallows. Greater light availability is positively related to seedling

height and height growth, seedling leaf number and leaf area, and seedling survival, and may

indirectly enhance sapling growth and survival. This enhanced survival and growth likely

contributes to the observed higher densities of seedlings and saplings in fallows. Not only can

anthropogenic disturbance, in the form of shifting cultivation, play a positive role in Brazil nut









regeneration, it could explain current Brazil nut densities and distributions. Finally, in light of

these findings, swidden fallows could potentially be managed for enhanced Brazil nut densities,

which may provide an opportunity for greater income for extractive families while contributing

to the sustainability of Brazil nut extraction in the long term.









CHAPTER 1
INTRODUCTION

Degraded and secondary forests are widespread throughout the tropics, representing 60%

of remaining tropical forests (ITTO 2002). In the Brazilian Amazon alone, over 12% of the

forest mosaic is classified as degraded and secondary forest (ITTO 2002). As these forests

undergo succession they are capable of conferring significant environmental and livelihood

benefits. Secondary forests provide people with food resources, timber and non-timber forest

products, including fuelwood, and quality hunting sites (ITTO 2002). They also regulate water

regimes, protect soils from erosion, store carbon, and serve as biological corridors and refuges

for species (ITTO 2002). In addition, regenerating secondary forests are especially important in

the life histories of a number of tropical plant species, including some that are of particular

socioeconomic and ecological importance, such as Cedrela spp. and Swietenia spp. (Whitmore

1989). Similarly, two other valued species, Astrocaryum murumuru and Dipteryx micrantha,

benefit from seed dispersal to treefall gaps (Cintra and Horna 1997). All four of these species

exhibit gap-opportunistic traits, achieving enhanced growth in higher light conditions found in

canopy openings or young regenerating forest environments.

Brazil nut (Bertholletia excelsa, H.B.K.) is another highly valued tropical tree reported to

be a gap-opportunistic species (Mori and Prance 1990, Myers et al. 2000), benefiting from

canopy openings during early stages of its life history. Brazil nut has emerged as the cornerstone

of the extractive economy in much of the Amazon (Clay 1997), and the seeds, commonly

referred to as nuts, are used for a variety of products, including raw and dried nuts, oils, flour,

medicines, and personal care products (Ortiz 1995), making Brazil nut a versatile and valuable

non-timber forest product (NTFP). To assess recruitment status as related to long-term harvest

sustainability of this valuable species, several regeneration studies have been conducted in recent









years (Viana et al. 1998, Zuidema and Boot 2002, Peres et al. 2003, Serrano 2005), yielding

differing reports on the impact of nut harvest on Brazil nut population structures. However,

studies to date have been conducted solely within mature forest sites. The modern Amazonian

landscape consists of a mosaic of mature forest, regenerating secondary forest, swidden fallow,

and pasture; therefore, in light of the increasing extent of secondary forests in the region, the role

of these sites in critical economic and conservation species recruitment and regeneration

processes should not be underestimated.

In Acre, Brazil, local extractivists report relatively high levels of Brazil nut recruitment in

regenerating swidden fallows, which may be due to specific biotic and abiotic characteristics

associated with these sites, including differences in seed disperser activity (Clay 1997, personal

obs.), light availability (Kainer et al. 1998, Myers et al. 2000, Zuidema 2003), and nutrient

availability (Kainer et al. 1998), as well as increased resistance to pathogens in higher light

environments (Augspurger 1984b). The aim of my study is to assess to what extent swidden

fallows constitute favorable regeneration environments for B. excelsa. Although various studies

have assessed the performance of tropical seedlings in natural gaps (Popma and Bongers 1991,

Cintra and Horna 1997, Myers et al. 2000), typically formed by treefalls, conclusions regarding

recruitment, survival and growth in these canopy openings should not be applied to all cases of

canopy disturbance. For the purpose of this study, I highlight some characteristics unique to

secondary fallow forests, in comparison with treefall gaps, as well as other regenerating forest

types, which may each contribute considerably different influences on B. excelsa regeneration.

Fallows forests differ from treefall gaps in many ways. To begin with, the most common

treefall gap size is < 20 m2 (Brokaw 1982), while swidden plots average 2 hectares (equivalent to

20,000 m2) (Fujisaka et al. 1998). Abiotic conditions, including light and temperature levels,









vary within gaps (Denslow et al. 1990). As a result, one might observe a larger range of light

and temperature levels in larger gap areas, such as swidden plots. Also, herbivore activity may

differ as a result of this difference in gap size (Janzen 1990), as some vertebrates prefer

secondary forest environments (Pefia-Claros and De Boo 2002), but others avoid crossing large

clearings (da Silva et al. 1996).

What may differentiate fallows most from other secondary forests is the influence of land

use history on succession and regeneration (Uhl et al. 1988, Guariguata and Ostertag 2001,

Mesquita et al. 2001). In general, swidden fallows develop after 2 to 4 years of cultivation, and

are left to regenerate for a period of 5 to 15 years (Montagnini and Mendelsohn 1997). In fields

which have been subjected to little burning, Cecropia spp. are often the dominant pioneers to

establish in young fallows, however, where burning has been more frequent, Vismia spp. often

dominate (Borges and Stouffer 1999). The dominance of these early pioneers directly influences

subsequent vegetation succession (Nascimento et al. 2006). For example, in Cecropia-

dominated areas heavy litterfall and increased shading can inhibit seedling establishment

(Mesquita 1995, cited in Nascimento et al. 2006). As succession proceeds, differences in forest

structure, stem density, and species composition can be observed. Fujisaka et al. (1998) found

species richness to be higher in 3 5 year old fallows than in 1 2 year old fallows, and higher

in fallows than in pasture sites. Also, vines were more abundant in mature forest and fallows of

3 5 years than in cultivated sites and pasture (Fujisaka et al. 1998). Furthermore, grass and

weed dominance was reduced in fallows, compared to pasture and agroforestry sites, resulting in

less competitive suppression (Ferguson et al. 2003). More shrub species were found in cropped

and fallow sites than in mature forest and pasture (Fujisaka et al. 1998), and herbaceous plants

were most common in cropped fields and pasture. Finally, as a result of burning and crop









harvesting, nutrient availability may be significantly altered in young fallows (Uhl and Jordan

1984).

After a few years of succession, large regenerating clearings may harbor increased

numbers of vertebrates due to greater food availability for certain species than that of mature

forest (Janzen 1990). Composition of seed dispersers may also differ between gap types, as a

result of hunting activities that often coincide with crop cultivation (Janzen 1990).

In large man-made clearings, new recruitment is dependent upon seed dispersal (Ferguson

et al. 2003), if cropping activities are carried out such that adults of desired species are removed

or avoided, and no seeds are present in the seed bank. In this case, nearby patches of mature

forest are required for forest trees to colonize abandoned plots (Uhl et al. 1982). On the other

hand, recruitment may result from both dispersal and adult seed rain in a treefall gap. Although

swidden fallows are often surrounded by other fallows or secondary forests of varying ages

(Ferguson et al. 2003), recruitment processes may not be drastically compromised relative to

those in treefall gaps, since mature forest is often left intact along at least a portion of the

swidden edge (personal obs.). All of the aforementioned differences associated with forest gap

type are relevant to any analysis of recruitment and regeneration processes of species such as B.

excelsa in disturbed areas.

My study offers a new look at the seedling ecology of a valuable NTFP, through a focused

study on the post-germination phase of B. excelsa regeneration within two multiple-use areas in

the southeastern portion of Acre, Brazil. I predicted that densities of B. excelsa individuals

< 10 cm diameter at breast height would be greater in fallow than mature forest for two main

reasons: 1) local extractivists report relatively high agouti seed dispersal in fallows (Clay 1997),

and 2) survival and growth of young B. excelsa individuals is reported to be enhanced in the









presence of higher light availability in forest gaps (Mori and Prance 1990, Myers et al. 2000). In

addition to evaluating seeding and sapling densities, light levels, soil nutrient content, leaf

herbivory level, and number of nearby seed sources were evaluated in both fallow and mature

forest to determine their respective influences on B. excelsa survival and growth. Chapter 2 is

presented as a single, independent chapter ready for submission to a peer-reviewed journal.

Relevant conclusions and management implications are presented in Chapter 3.









CHAPTER 2
SHIFTING CULTIVATION EFFECTS ON BRAZIL NUT (Bertholletia excelsa)
REGENERATION

Introduction

Brazil nut (Bertholletia excelsa, H.B.K.), has emerged as the cornerstone of the extractive

economy in much of the Amazon (Clay 1997), which has given rise to the debate over whether

or not current harvest levels have a detrimental effect on Brazil nut seedling recruitment. Some

scholars report that current harvest levels have minimal to no detrimental effect on seedling

recruitment in selected Brazil nut populations (Viana et al. 1998, Zuidema and Boot 2002), while

others report that regeneration is rare or nonexistent in over-exploited populations (Peres et al.

2003, Serrano 2005). All previous Brazil nut regeneration studies have been conducted solely

within mature forest, however, species regeneration is not limited to this forest type. The

modern Amazonian landscape is a mosaic of mature forest, regenerating secondary forest,

swidden fallow, and pasture; which comprise a continuum of microhabitat suitability for B.

excelsa recruitment and regeneration. An evaluation of Brazil nut regeneration ecology is

needed across the entire mosaic of forest types in order to provide an accurate picture of current

B. excelsa population structures.

Bertholletia excelsa may regenerate more successfully in secondary than mature forest,

based on reports that it is a gap-opportunistic species (Mori and Prance 1990, Myers et al. 2000).

Bertholletia excelsa is one of a number of shade-tolerant species which experience enhanced

seedling survival and growth in canopy openings, characterized by higher light levels (Brokaw

1985, Whitmore 1989, Molofsky and Fisher 1993, Cintra and Horna 1997). According to Myers

et al. (2000), B. excelsa seedlings not only benefit from forest gaps, they require a minimum gap

size of> 95 m2 or > 10.4% global site factor in order to reach sapling size. One large-scale

disturbance yet unexplored for B. excelsa recruitment is that of recent anthropogenic secondary









forests, which, in addition to being widespread throughout Amerindian history (Denevan 1992,

Heckenberger et al. 2003), are becoming increasingly common in the Amazonian landscape

(ITTO 2002).

In Acre, Brazil, landholders annually clear 0.5 2 ha patches of older secondary or mature

forest for swidden agriculture. When agricultural sites are abandoned, unique secondary forests

succeed in fallow sites, which may constitute particularly favorable regeneration environments

for B. excelsa. Local extractivists describe fallows as having greater B. excelsa recruitment than

mature forest, associated with a relatively higher incidence of agouti seed dispersal to these

disturbed sites. In addition, seedling survival may also be enhanced in fallow, due to decreased

vulnerability to pathogens in higher light environments (Augspurger 1984b). Subsequent

establishment and growth of B. excelsa may be greater in fallows because of high light levels

(Kainer et al. 1998, Myers et al. 2000, Zuidema 2003) and greater nutrient availability (Kainer et

al. 1998).

An evaluation of the potential of fallow sites to provide favorable B. excelsa regeneration

environments will complement previous assessments of tropical seedling performance in other

disturbed sites, specifically natural gaps (Popma and Bongers 1991, Cintra and Horna 1997,

Myers et al. 2000). Although both types of disturbance increase forest light availability, fallows

represent ecologically unique microhabitats which may differentially affect seedling recruitment,

survival and growth. As a result of the shifting cultivation process, which often includes the

application of fire, fallow forests may be characterized by significantly different nutrient stocks

(Uhl and Jordan 1984), plant assemblages (Fujisaka et al. 1998, Ferguson et al. 2003), and

vertebrate activity (Schupp 1988, Cintra and Horna 1997, Pefia-Claros and de Boo 2002).









This study directly tests the hypothesis, based upon aforementioned scientific evidence and

extractivist observations, that B. excelsa seedling and sapling densities are greater in fallows than

mature forest, by assessing the post-germination phase ofB. excelsa recruitment. To understand

some of the mechanisms behind observed recruitment, effects of light, nutrient availability and

proximity of seed sources are also assessed. In addition, one-year survivorship and growth rates

of young individuals, seedling leaf production, and seedling herbivory, which further shape B.

excelsa recruitment, are compared between fallow and mature forest.

Species Description

Bertholletia excelsa is a monospecific member of the Lecythidaceae family, found in

unflooded (terra fire) forests across the Amazon basin and the Guianas (Mori and Prance

1990). This canopy emergent can reach up to 50 m in height (Mori and Prance 1990), and some

individuals have been recorded at almost 1000 years of age (Vieira et al. 2005). In the two

landholdings surveyed in my study (Filipinas and Cachoeira), densities of Brazil nut trees

> 10 cm diameter at breast height (dbh) were 1.4 individuals ha-1 (Wadt et al. 2005) and 2.5

individuals ha-1 (Serrano 2005), respectively, which are representative of densities reported by

Peres et al. (2003) elsewhere in the study region. Although some authors report that B. excelsa

occurs in groves of 50 100 individuals, with each grove separated from another by distances of

up to 1 km (Mori and Prance 1990), Wadt et al. (2005) reported no existence of groves in my

study area.

Mammal communities play a critical role in the regeneration of many large-seeded tropical

tree species (Sork 1987, Asquith et al. 1997, Sanchez-Cordero and Martinez-Gallardo 1998,

Forget et al. 2000, Jansen 2003), via seed dispersal and seed predation. Brazil nut is no

exception, as the regeneration ofB. excelsa is highly dependent upon the presence of agoutis

(Dasyprocta spp.), the primary dispersers of Brazil nut seeds (Huber 1910, Prance and Mori









1978). Agoutis have been observed burying seeds at clearing edges (Peres and Baider 1997) and

in young secondary forest (Forget et al. 2000), and extractivists have often observed agoutis

carrying seeds to fallows (Clay 1997, personal obs.). Agoutis may prefer fallow sites because of

the protection provided against predators by regenerating vegetation (Cintra and Horna 1997).

These reports suggest a possibility of comparable or greater seed dispersal to fallows than mature

forest. Recruitment ofB. excelsa in both mature forest and fallows may also depend upon

density and proximity of reproductive adults, as the proportion of seeds that arrive at a certain

point in the forest should decline with distance from parent tree (Janzen 1970). Agoutis have

been observed dispersing B. excelsa seeds as far as 100 m from seed sources, with the majority

of seeds being dispersed within 25 m (Peres and Baider 1997). Normal seed shadows can be

extended by secondary dispersal, when seeds are unearthed by seed predators and reburied even

further from the adult tree (Mori and Prance 1990, Peres et al. 1997).

Study Area

Fieldwork was carried out in the southeastern portion of Acre Brazil, between 100 and 110

south of the equator in Acre, Brazil. The region has undulating topography, vegetation classified

as humid, moist tropical forest (Holdridge 1978), and a pronounced three month dry-season from

June to August. Average annual rainfall is from 1600-2000 mm (IMAC 1991). Soils of the

region are classified under the Brazilian classification system as Argissolos (ZEE 2000), which

correspond roughly to U.S. Soil Taxonomy System Ultisols. Recruitment of B. excelsa was

evaluated within RESEX Chico Mendes and the Chico Mendes Agro-Extractive Settlement

Project (PAE Chico Mendes), multiple use areas of 976,570 ha and 24,898 ha, respectively,

separated by 30 kilometers (Fig. 2-1). Extractivists in the two study sites subsist on agricultural

and cattle production, rubber tapping and Brazil nut collection (Gomes 2001), and some timber

extraction in PAE Chico Mendes (personal obs.).









In my study sites, 0.5 1.5 ha of mature or older secondary forest found in close proximity

to households was manually cleared of large trees to initiate the shifting cultivation cycle.

Clearings were created either in areas void of valuable Brazil nut and rubber (Hevea brasiliensis)

trees, or great care was taken to protect individuals of these species. Clearings were burned

once, first cultivated for corn and rice, followed the next year by beans, and then manioc. After a

cultivation period of two to three years, agricultural sites were abandoned, and regenerating

secondary forests formed in situ. Young secondary forests such as these are typically

characterized by lower stand basal area than surrounding mature forest (Saldarriaga et al. 1988).

Although these swidden fallows are often surrounded by other fallows or secondary forests of

varying ages, mature forest bordered at least one length of each fallow border studied. Thus, the

study landscape consisted of a forest matrix with small fallows and pastures embedded within

mature forest expanses.

Methods

Plot Installation and Sampling Scheme

Mature forest plots (300 x 300m or 9 ha each) in RESEX Chico Mendes and PAE Chico

Mendes were originally demarcated and delineated in 2002 (Serrano 2005) using transects

opened every 25m and extending for 300m (Appendix A). Transects produced 25 x 25m grids of

9-hectare survey plots (Fig. B-1[A]). From May to July 2005, transects within three of these

original plots in RESEX Chico Mendes were reopened to evaluate all B. excelsa individuals

> 10cm dbh by searching to the right and left of systematically placed parallel lines spaced 25m

apart. To survey 25 % of the area of each mature forest plot for recruitment, four mature forest

subplots (25 x 25 m each) were randomly selected within each hectare. Thus randomization was

restricted to each one-hectare plot, generating a total of 36 subplots (Fig. B-1[A]).









Six fallows formed in RESEX Chico Mendes (0.5 1.0 ha each), and two in PAE Chico

Mendes (1.0 1.5 ha each), were also delineated in 2005 (Fig. B-1[B]), using transects opened

every 10m. Fallow ages ranged from 5 to 12 years, based upon year of abandonment of the

agricultural site and, due to their variable sizes and shapes, were delineated and demarcated with

a Garmin 12XL3 GPS unit. All individuals > 1.5 m in height were located in fallows by

searching to the right and left of parallel lines spaced 10 m apart. Recruitment subplots in

fallows (10 x 10 m) were randomly assigned across the extent of each fallow, and comprised

approximately 25% of each fallow area (Fig. B-1[B]).

Regeneration Environments by Forest Type

To evaluate the influence of seed sources on recruitment, all B. excelsa individuals

> 10 cm dbh were tallied within each 9 ha mature forest and 0.5 1.5 ha fallow, as well as within

a 50m border strip of each plot. Dbh, XY coordinates, and reproductive status were recorded for

each individual.

To compare understory light availability among 2 mature forest plots and 4 fallows in

2005, photosynthetically active radiation (PAR) was estimated by averaging photosynthetic

photon flux density (PPFD) measurements taken at 10 equidistant sample points running through

the center of each plot, from one plot edge to the opposite edge (Appendix B). Measurements

were taken every 30 m in mature forest plots and measurement spacing varied in fallow sites,

due to variation in fallow size. To capture daytime variations, PPFD was measured in each plot

every 2.5h from 7:30 to 15:00 (ie. four times per day) for three days. PPFD was measured using

10 gallium arsenide phosphide (GaAsP) photocells mounted on a 10-cm disk and connected to a

multimenter (Micronta LCD Digital Multimeter 22-185A; Tandy Corporation, Fort Worth,

Texas). Sensors were calibrated with a standard LI-COR quantum sensor (model LI190SA) and

a Campbell CRO1x data logger.









To determine soil properties within each fallow and mature forest plot, soil samples were

collected in 2005 below the litter layer at two depths (0 20 cm and 20 40 cm). In the fallows,

soil cores were collected at two depths within one randomly selected subplot in each of the four

corners of the fallow, as well as the center-most subplot, to create one composite of five soil

cores for each depth (Appendix B). In the 9 ha mature forest plots, soil cores were collected

within the four hectares located at the covers of each plot, as well within the center-most hectare

(Appendix B). Within each of these five sampled hectares, a composite of five soil cores was

created for each depth, using the same sampling scheme described for soil collection in the

fallows. Two replicates of each composite were dried for 4 days at 650C, and passed through a 2

mm stainless steel sieve. Soil pH was measured at a 1:2.5 soil to water ratio. Extractable P and

K were processed using a dilute double acid extraction (Mehlich-1), with concentrations of P

determined colorimetrically using the molybdate blue method (Murphy and Riley 1962), and K

concentrations were determined using flame emission spectrophotometry. To determine total

potential acidity, H+ and Al3+ were extracted with a buffered solution of calcium acetate at pH 7,

and then titrated with a 0.1 N NaOH. Oxidizable organic carbon was determined on soils passed

through a 1 mm screen, ground in a porcelain mortar, and then digested in a potassium

dichromate acid medium with external heat. All soil analyses were conducted at the Soils

Laboratory of Embrapa-Acre, Brazil (EMBRAPA 1997).

The temperature and moisture environment in fallow and mature forest were evaluated

between July 2005 and June 2006. Mean monthly temperature and percent relative humidity

were determined for two fallow sites and one mature forest site based on measurements taken at

30 minute intervals using HOBO ProSeries data loggers.









Seedling and Sapling Densities

Seedling densities were assessed within all subplots installed in mature forest and fallows.

Sapling densities in mature forest also were assessed within subplots. Due to the ease of locating

saplings in fallow, however, sapling densities were assessed over entire fallow areas, rather than

in subplots only. Individuals < 1.5 m in height were defined as seedlings and individuals

between 1.5 m in height and 10 cm dbh were defined as saplings. This classification scheme was

selected not to differentiate between those individuals still using their seed reserves for growth

and those individuals relying on environmental resources, but rather to look at general classes of

recruitment. Within each subplot, XY location coordinates, height, seedling basal diameter, and

sapling dbh were recorded for all individuals.

B. excelsa Survival and Growth

Survival and annual growth (seedling height and basal diameter and sapling dbh) were

evaluated from June to July 2006. To evaluate the effect of PAR on seedling survival and

growth, PPFD was measured directly above seedlings in two mature forest plots and four

fallows. To capture daytime variations and to measure the full complement of all seedlings in

each selected plot, seedling PPFD was sampled on a rotating basis every 2.5h from 6:45 to 16:45

(ie. five sampling times per day) for three days. To assess the influence of photosynthetic

capacity on survival and growth of all seedlings, leaf area was evaluated for seedlings in three

mature forest plots and four fallows. When leaf number totaled < 5, all observed leaves were

sampled, which was the case for the majority of seedlings. In all other cases, 5 leaves were

sampled, representing the full range of leaf sizes. To avoid destructive sampling, leaves were

traced in the field, and average leaf area was measured in Gainesville, Florida, with a LiCor, Inc.

Model 3000A Leaf Area Meter with Transparent Conveyor Belt Accessory (LiCor, Inc., Lincoln,









Nebraska). Total leaf area was then estimated by multiplying total number of leaves per seedling

by average leaf area.

The influence of invertebrate herbivory on seedling survival and growth was also assessed

by quantifying the percentage of leaves per plant that showed signs of attack in 2005. Vertebrate

leaf herbivory was not assessed because it was not possible to observe or account for the

significant portion of predation which is comprised of leaves that are completely removed by this

type of herbivore.

Data Analysis

All statistical analyses were performed using SAS software (Version 9.1, SAS Institute),

excepting Student's t-tests, which were performed using SPSS software (Version 10.0, SPSS

Inc.). Mature forest plots and fallows were treated as statistical blocks for all analyses, and

seedling and sapling densities were determined on a per hectare basis. Average soil values were

included as random effects, with plot nested within forest type, and soil nested within plot, for all

analyses. PPFD values were log transformed prior to all analyses to improve normality and

homoscedasticity of residuals, which were evaluated visually. PPFD levels were compared

between forest types using a repeated measures ANOVA, and soil nutrients were compared

between forest types using a two-way ANOVA. All soil nutrient values were averaged on a plot

basis, and average soil pH, potassium, and percent organic carbon at two depths (0 20 cm and

20 40 cm) were included in explanatory density and growth models. Correlation analysis,

using the PROC CORR procedure, revealed high correlations between the two depths for

phosphorous and potential acidity (H+Al3+), therefore these soil nutrient values were collapsed

into one depth for inclusion in the explanatory models.

Since B. excelsa densities demonstrated a Poisson distribution, they were modeled using a

general linear mixed model and the GLIMMIX procedure. Two-way correlations were









calculated between relevant variable pairs using PROC CORR in order to determine

relationships which merited inclusion in the model. I explored model building by conventional

methods of sequentially dropping non-significant interactions and covariates based on p-values.

In this case, I used a less restrictive significance level (a = 0.10) than the usual arbitrary 0.05

level (Johnson 1999, Burnham and Anderson 2002) to allow all possible significant effects to

remain for the purpose of building the best explanatory model (Bancroft 1968). Because a

primary purpose of data analysis was to evaluate differences between mature forest and fallow,

the indicator variable for forest type was considered fundamental and, therefore, retained in the

density models without regard to its significance level (Bancroft 1968). Densities were first

compared for all plots, with forest type (mature versus fallow) as the only retained fixed effect.

Seedling and sapling densities of mature forest plots previously surveyed in 2004 in PAE

Chico Mendes (Serrano 2005) were included in the comparative density analysis to increase my

sample size. These data were collected using the same sampling scheme described herein, and it

was assumed that no event had occurred that significantly affected these seedling and sapling

densities in this mature forest site (PAE Chico Mendes) between 2004 and 2005. To test this

assumption, 2004 seedling and sapling density data collected mature forest in RESEX Chico

Mendes, using the same sampling scheme, were compared with my 2005 seedling and sapling

density data from this same forest and plots and no statistical differences were detected

(p = 0.57). Finally, no site (PAE Chico Mendes versus RESEX Chico Mendes) interaction was

found when comparing B. excelsa densities; therefore, observations from both sites were

included in the analysis, with site dropped as a fixed effect from the model.

To explain observed densities, a subset of data (2 mature forest plots and 4 fallows) was

analyzed. Tested fixed effects included forest type, number of reproductive adults within 50 m









of each subplot, all plot soil content averages, and plot PPFD averages; insignificant variables

were sequentially removed from the final model. All possible predictor variables were included

in the final model to provide the best explanatory model; however, residuals were not well

distributed.

Student's t-tests were used to compare initial height, initial basal diameter, height and

basal diameter growth, number of leaves, total leaf area, and leaf herbivory between fallow and

mature forest for all seedlings; all data were normally distributed. Sapling diameter growth,

which was likewise normally distributed, was also compared using a t-test. Logistic regression

was used to analyze differences in seedling survival; tested fixed effects included initial height,

initial diameter, height growth, diameter growth, forest type, all plot soil content averages,

PPFD, number of leaves per seedling, total leaf area, and percent leaf herbivory. Insignificant

variables were sequentially removed from the final model. Initial seedling height and diameter

were highly correlated with total leaf area, therefore, interactions were explored between total

leaf area and initial seedling size.

The GLIMMIX procedure was used to test seedling and sapling growth. Growth values,

some of which were negative, were log transformed (log 10 [1+x]) prior to analyses to improve

normality and homoscedasticity of residuals, which were observed visually. Because data

related to more mechanistic causes of growth (ie. light observed directly above seedlings) were

available, the forest type variable was intentionally excluded from the seedling growth analysis.

However, because these data were unavailable for saplings, forest type was retained for analysis

in the sapling diameter growth model. Tested fixed effects included in initial seedling growth

models were initial height or diameter, forest type, all plot soil content averages, PPFD, number

of leaves per seedling, total leaf area, and percent leaf herbivory, while initial sapling diameter









growth models included initial dbh, forest type, all plot soil content averages, and plot PPFD

averages. Again, all possible predictor variables were included in the final models to provide the

best explanatory models of growth, and insignificant variables were sequentially removed from

the final model; residuals were fairly well distributed. Sapling height growth could not be

compared between forest types, as sapling heights were estimated visually.

Differences in seedling leaf herbivory percentage were analyzed with the GLIMMIX

procedure, accounting for a log-normal distribution of data. Leaf herbivory percentages were

arcsin transformed prior to analysis in order to improve the normality of the response variable.

Tested fixed effects included initial height or diameter, forest type, all plot soil content averages,

PPFD, number of leaves per seedling, total leaf area; insignificant variables were sequentially

removed from the final model. Normality and homoscedasticity of residuals were observed

visually and the residuals were fairly well distributed.

Results

Regeneration Environments by Forest Type

Of all variables measured to characterize regeneration environments (plot light levels,

density of nearby seed sources, temperature and relative humidity, and soil nutrient stores), only

light levels differed significantly between the two forest types. A repeated measures ANOVA

revealed higher average PPFD levels in fallow than mature forest (p = 0.02) (Fig. 2-2). There

were no significant differences between forest types in number of adults within 50 m of

evaluated subplots, nor in observed daily temperature and percent relative humidity monitored in

two fallows and one mature forest site between July 2005 and June 2006. Mean monthly

temperature in the three sites ranged between 23.5 and 24.20C and mean relative humidity

ranged between 90.3 and 92.7%. Although two-way ANOVAs revealed no soil property









differences between forest types (Table 2-1), soil P was significantly different by depth (p =

0.0006).

Seedling and Sapling Densities

Observed mean seedling densities of 12.7 and 5.3 trees ha-1 in fallow and mature forest

(Fig. 2-3), respectively, were different at p = 0.07, according to the general linear mixed model.

Mean sapling densities of 5.2 and 1.3 trees ha-1 in fallow and mature forest (Fig. 2-3),

respectively, were also different, again at p = 0.07. Based upon an analysis of the data subset,

seedling density differences were somewhat explained by the number of adults within 50m of

each subplot (p = 0.06), such that seedling densities increased with number of adults, while

sapling densities were best explained by percent organic carbon at the 0 20 cm depth (p = 0.02)

(Table 2-2).

B. excelsa Survival and Growth

Seedlings

After one year, 26% of surveyed seedlings in fallow and 19% in mature forest (21% of all

seedlings) were either dead or unable to be located, with no survivorship differences between

these two forest types (Table 2-3). Seedling survival was possibly explained by total leaf area (p

= 0.08) and an interaction between percent leaf herbivory and PPFD (p = 0.08) (Table 2-2);

improved survival was associated with greater leaf area and higher leaf herbivory.

Initial seedling height was not significantly different between forest types (Table 2-3),

and, across forest types, seedling height in 2006 was best explained by PPFD measured in 2005

(p = 0.0009) (Fig. 2-4); seedling height increased with PPFD increase. Seedling height growth

was best explained by an interaction between percent leaf herbivory and PPFD (p = 0.02) and

percent leaf herbivory alone (p = 0.02) (Table 2-2). Height growth increased with PPFD, and

increased with herbivory decrease. Seedling growth appeared to increase more with PPFD at









intermediate herbivory levels. A t-test revealed no significant difference in initial seedling basal

diameter between forest types (Table 2-3), however, basal diameter growth was greater in fallow

than mature forest (p = 0.05), according to a t-test. According to the general linear mixed model,

seedling basal diameter growth was best explained by total leaf area (p = 0.05) (Table 2-2), such

that seedlings with more leaf area grew more. Seedling height and diameter growth

demonstrated positive relationships to all tested factors except percent leaf herbivory.

Total leaf area was highly correlated with seedling height, diameter and number of leaves

and, as a result, less significant variables sequentially fell out of the growth models. Total leaf

area was strongly related to PPFD (p = 0.0004) (Fig. 2-5). Observed herbivory level was much

higher in mature forest than fallows (p < 0.0001) (Table 2-3). The mixed model corroborated

that percent seedling leaf herbivory was explained by forest type alone (p = 0.05) (Table 2-2),

but also showed that herbivory was best explained by an interaction between forest type and

number of leaves (p = 0.006) (Table 2-2), such that herbivory decreased with leaf number in

mature forest, but increased with leaf number in fallow.

Saplings

Estimated mean sapling heights were 5.51 and 2.99 m, in mature forest and fallow,

respectively, which was significantly different (p = 0.02) (Table 2-3). Initial sapling dbh was

also greater in mature forest than fallow (p = 0.05) (Table 2-3). Maximum dbh values were 9.5

cm in mature forest and 7.0 cm in fallow. Nearly 100% sapling survival was observed in both

forest types (only one sapling died in a single fallow site), and therefore no significant difference

in sapling survival between forest types was detected. Sapling dbh growth was much greater in

fallow than mature forest (p = 0.02), and was potentially explained only by forest type (p = 0.07)

(Table 2-2).









Discussion


Densities Explained

Bertholletia excelsa seedling densities were over two times greater in anthropogenically-

disturbed fallows than mature forest, and sapling densities were four times greater in fallow (Fig.

2-3). Many factors influence establishment, survival, and growth of young individuals, including

proximity and density of seed sources (Janzen 1970), seed disperser and predator behavior (Sork

1987, Forget 1994, Molofsky and Fisher 1993, Asquith et al. 1997), light availability

(Augspurger 1984a, Augspurger 1984b, Poorter 1999), and nutrient availability (Ceccon et al.

2003), some of which may be related to microhabitat, or forest type. My data suggest that the

influence of density and proximity of seed sources and light levels, at various stages in the B.

excelsa recruitment process, play a significant role in shaping the observed densities of young B.

excelsa individuals. Differences in densities between forest types, however, could also be a

reflection of additional factors not measured, such as seed dispersal. These factors, which affect

species establishment to differing degrees at various stages in the recruitment process, will be

discussed below.

Seed sources and dispersal

Based on a deeper analysis of a subset of data, which included number of seed sources

within 50m of surveyed subplots, seedling densities were mildly related to the number of nearby

seed sources (p = 0.06). These findings are consistent with scientific evidence that seedling

densities decline with increased distance from parent trees (Janzen 1970). Therefore, higher

recruitment would be expected in fallows that either contain adult B. excelsa trees or are located

adjacent to mature forests with high adult densities, compared to those with no adults nearby.

Although density of nearby adults explained relative densities, seedlings and saplings were often

found in areas with neither a reproductive adult within 50m, nor traces of potential seed sources









that may have suffered mortality prior to my study, confirming seed dispersal distances well

beyond the typical dispersal maximum of 25m (Peres and Baider 1997). These dispersal

distances may be explained by secondary dispersal (Mori and Prance 1990, Peres et al. 1997).

Residuals in these explanatory density models were poorly distributed, with high density

observations being grossly underestimated, suggesting that relevant factors have not been

included in the models. These underestimations could be due to a clumped, and possibly

directed, seed dispersal pattern that has not been accounted for. Bertholletia excelsa seed

dispersal beyond the adult crown is determined nearly entirely by agouti scatterhoarding

behavior (Prance and Mori 1978), and local extractivist reports suggest that agouti dispersal

behavior may be non-random. Extractivists report preferential seed dispersal in fallow

environments (Clay 1997, personal obs.). These accounts are corroborated by observations of

seed burial at clearing edges (Peres and Baider 1997) and in young secondary forest (Forget et

al. 2000), perhaps related to agouti predator avoidance in areas composed of limb and vine

tangles (Cintra and Horna 1997). Agouti scatterhoarding activity may potentially increase the

proportion of seeds in favorable microenvironments (Jansen and Zuidema 2001), which has been

proposed as a more critical influence on regeneration than post-dispersal predation escape or

successful germination (Rey and Alcantara 2000). The importance of disperser site preference

and directed dispersal has been demonstrated for species other than B. excelsa (Uhl et al. 1981,

Schupp et al. 1989, Forget 1994). Although disperser behavior was not tested in this study, it

likely had a significant effect on seed availability for recruitment in fallow and mature forest.

Equally important is the influence of differential seed removal rates (due to retrieval and

predation) between different forest types, which may further affect observed B. excelsa densities.

Some authors report higher seed removal rates of various tropical species in gap environments









(Schupp 1988, Schupp and Frost 1989, Cintra and Horna 1997), while Sanchez-Cordero and

Martinez-Gallardo (1998) report greater tropical seed removal in mature forest than gaps.

Tabarelli and Mantovani (1996), did not find seed removal of B. excelsa to be different between

treefall gaps and undisturbed forest, however, clearings created by anthropogenic disturbances

were not assessed. It is clear that seed disperser and seed predator activities are essential factors

to include in future density evaluations.

Seedling survival and growth

Once seeds are dispersed and young seedlings establish, individuals may demonstrate

differential growth (Augspurger 1984a, Popma and Bongers 1991, Brown and Whitmore 1992,

Poorter 1999) and/or survival (Augspurger 1984b, Cintra and Horna 1997, Brown and Whitmore

1992) in different forest types, which are often differentiated in the literature between understory

and gap environments. In my study, however, forest type explained only sapling diameter

growth (sapling height growth could not be evaluated). Other measured variables, some of

which may be associated with forest type, demonstrated stronger influences on observed survival

and growth of B. excelsa seedlings.

To begin with, total leaf area, which is correlated with leaf number, was a mild indicator of

seedling survival (p = 0.08). Leaves facilitate the acquisition of carbon by means of

photosynthesis; an enhanced carbon gain, due to greater leaf area, may confer relatively higher

fitness (and ultimately survival) to seedlings by various means, including the provision of more

structural material and energy, an increase in nutrient content, and a better tolerance to

environmental stresses (Chabot and Hicks 1982). PPFD, although not highly significant on its

own, may contribute to increased seedling survival, as PPFD was strongly positively correlated

with total leaf area (p = 0.0004), which did explain seedling survival at p = 0.08. Although some

studies have demonstrated greater seedling survival in gaps than understory for some species









(Augspurger 1984b, Cintra and Horna 1997), my results do not directly support these

conclusions, as I found one-year B. excelsa survivorship to be no different between mature forest

and fallow. Kainer et al. (1998) also found no relationship between seedling survival and

comparative light levels in pasture, forest gap, and shifting cultivation sites (Kainer et al. 1998).

The positive effect of leaf herbivory on survival is not so easily explained. Contrary to my

findings, numerous studies demonstrate the negative effect of leaf herbivory on seedling survival

(Howe 1990, Hulme 1996, Vasconcelos and Cherrett 1997). In this study, the influence of

herbivory level on survival may be coincidental. Invertebrate herbivory was not unusually high

in either forest type, and is probably not sufficient to lead to seedling mortality under normal

conditions. In addition, it was not possible to quantify vertebrate herbivory on seedling survival

in this study. Nevertheless, it is likely that a greater percentage of mortality was represented by

completely uprooted seedlings (Ortiz 1995, Zuidema 2003, personal obs.), resulting from seed

predators unearthing and consuming seeds at the base of the plant. Furthermore, only 14 of 94

total seedlings died, therefore concrete differences between factors relating to seedling

survivorship were difficult to detect. Although forest type was not a significant factor in survival

probability, in light of other analyzed effects, it is possible that the forest type variable was

partially masked by other variables such as percent leaf herbivory or PPFD (due to multiple

colinearity), which were both significantly different by forest type (PPFD was higher in fallow

and leaf herbivory was lower in fallow than mature forest). If forest type is partially included in

one of these variables, it may play an undetected role in determining seedling survival

probability.

Seedling heights in 2006 were highly positively correlated with PPFD levels measured

directly above seedlings in 2005, which implies a significant advantage to height growth in









higher light environments. In my study, one-year seedling height growth was influenced by an

interaction between PPFD and percent leaf herbivory (p = 0.02), such that height growth

declined with increased leaf herbivory and, while generally increasing with PPFD, was greatest

at intermediate light levels. Previous studies have shown a positive relationship between

seedling height growth and light availability in the tropics (Augspurger 1984a, Denslow et al.

1990, Popma and Bongers 1991), and similar results have been reported for B. excelsa (Poorter

1999, Zuidema 2003), again, with highest growth rates at intermediate light levels.

Uhl and Jordan (1984) found that soil nutrient concentrations did not vary significantly

between agricultural sites and mature forest 5 years after cutting and burning. Similarly, I found

no difference in soil nutrient availability between mature forest and fallows. As a result, soil

nutrients did not explain B. excelsa survival and growth as well as other measured variables,

such as light availability.

Numerous studies have described the negative effects of herbivory on seedling growth

(Marquis 1984, Howe 1990, Hulme 1996, Vasconcelos and Cherrett 1997). In my study,

seedling leaf area did not differ significantly by forest type, but was highly correlated with

seedling height, and the two variables exhibited a significant interaction effect for seedling

height growth. Height growth was positively related to both variables, similar to other studies

which have demonstrated increased growth with greater initial height (Zuidema 2003) and leaf

area (Poorter 1999). Similar to height growth, seedling diameter growth was explained by total

leaf area (p = 0.05), and possibly by an interaction between leaf herbivory and PPFD (p = 0.07),

with a positive relationship to each factor. Zuidema et al. (2003) report a similar relationship

between B. excelsa biomass growth and light availability. Although not directly evaluated,

seedling survival and growth in my study sites may also have been influenced by ant presence.









Leaf cutter ants (Atta spp.) are known to modify soil nutrient concentrations (Lugo et al. 1973,

Haines 1978), which may directly influence the development of young B. excelsa individuals, as

well as influence the competitive environment with other species. In addition, extractivists assert

that leaf cutter ants preferentially predate both B. excelsa seeds and seedlings. The potential

importance of leaf cutter ants in fallow environments is worth particular attention as leaf-cutting

ant populations have been shown to increase after forest clearing (Vasconcelos and Cherrett

1995). Blanton and Ewel's (1985) findings suggest that leaf-cutting species continue to thrive as

succession proceeds, representing the most important herbivore in the successional forests they

studied. I found one of my six surveyed fallows was covered to a great extent in leaf cutter ant

nests (species unknown). The relationship between forest type and ant density, intensity of ant

seed and seedling predation in fallow and mature forest, and the effects of ant populations on soil

composition all merit further consideration, to better understand the role of ant populations in B.

excelsa recruitment and regeneration.

Sapling survival and growth

High sapling survival rates in both mature forest and fallow suggest that, once seedlings

have been well-established, they develop and survive equally well in either forest type. Zuidema

and Boot (2002) also found extremely low mortality rates (1%), over a two-year period, for B.

excelsa individuals > 1 cm dbh. Initial sapling diameter was greater in mature forest, but this can

be explained by the young age of the fallows (5 12 years), as fallow saplings have not had the

opportunity to attain dbh sizes near 10cm within this time frame. No factor included in statistical

analyses, aside from forest type, explained higher diameter growth rates in fallow. However,

saplings in fallow may also be able to invest more in biomass growth than mature forest saplings,

since height growth is greatly enhanced by higher light levels in fallow. Light is cited as the

most frequently limiting resource for seedling growth (Kitajima 1996), a major component of









tropical plant regeneration processes (Chazdon et al. 1996). Increased light availability is likely

one of the most significant advantages conferred upon well-established seedlings in fallow, as

observed PPFD was significantly greater in fallow than mature forest (p = 0.02), and the benefits

of light availability are likely great for individuals up to 10 cm dbh, which continue to compete

to reach the canopy up to, and even beyond, this 10 cm limit.

Secondary forests are, by nature, dynamic and, as a result, are associated with an ever-

changing microenvironment of vegetation, water, and soil nutrients. Therefore, the suitability for

seedling and sapling growth and survival may vary with forest successional stage. However, the

observed higher densities of young individuals in fallow indicate that this forest type, in general,

constitutes a favorable microhabitat for both seedling and sapling development.

Greater Ecological and Management Implications

This research provides a new, and perhaps more accurate, depiction of current Brazil nut

regeneration in tropical forest landscapes, as secondary forest environments have previously

been ignored. The high densities of young Brazil nut individuals in secondary forests created

through shifting cultivation fallowss) clearly demonstrate that Brazil nut regeneration may have

been underestimated in previous studies which were conducted solely within mature forest.

In light of these findings, fallows could potentially be managed for enhanced Brazil nut

densities, which may provide an opportunity for greater income for extractive families while

contributing to the sustainability of Brazil nut extraction in the long term. In the past, some

extractivists have experimented with small-scale Brazil nut enrichment plantings, as a means to

economically enrich their landholdings (Kainer et al. 1998). The benefits of fallow management

may also provide an additional incentive for extractivists to allow fallows to regenerate, rather

than convert them to pasture, which is an increasingly common practice in much of the Amazon.









Shifting cultivation has previously been recognized for its role in shaping Amazonian

ecosystems (Dufour 1990). Pre-Columbian human interventions, in the form of active planting,

were previously considered by Posey (1985) and Balee (1989) as a possible explanation for the

presence of Brazil nut "groves" in parts of the Amazon. Based on the results of this study,

however, it is possible that higher densities are a result of less direct human interventions, such

as shifting cultivation. Not only can anthropogenic disturbance, in the form of shifting

cultivation, play a positive role in Brazil nut regeneration, it could explain current Brazil nut

densities and distributions.









CHAPTER 3
CONCLUSION

The objective of this study was to compare B. excelsa recruitment and regeneration in

regenerating swidden fallows and mature forest. My results suggest that fallows constitute

favorable microhabitats for both seedling and sapling development. Perhaps most importantly,

survival and growth of young B. excelsa individuals appear to be enhanced by the higher light

levels found in fallow. Greater light availability is positively related to seedling height and

height growth, seedling leaf number and leaf area, and seedling survival, and may indirectly

enhance sapling growth and survival. This enhanced survival and growth likely contributes to

the observed higher densities of seedlings and saplings in fallow sites.

At the conclusion of this study it became clear that seed dispersal and seed predation

should be compared between forest types in future regeneration studies, in order to better assess

the various recruitment advantages, and possibly disadvantages, associated with fallow

environments. In addition, herbivore activity, and its effects on B. excelsa survival and growth,

should be evaluated in greater detail, quantifying the proportion of herbivory exercised by both

vertebrates and invertebrates, in both forest types. Leaf cutter ant behavior in regenerating

fallow fields merits particular consideration. Future studies should also compare B. excelsa

regeneration between different secondary forest types, as well as between fallows of varying

ages, as various successional forest types and successional stages may contribute to differential

survival and growth of young individuals. At the time of thesis submission, soil analyses were

incomplete, therefore conclusions regarding the effects of soil content were limited. Upon

publication of Chapter 3 in a scientific journal, the incorporation of additional soil data should

yield further insights.









Although some significant factors in B. excelsa recruitment processes, such as disperser

activity, were not evaluated in the study, the importance of the results should not be

underestimated. My research focuses on a widespread forest type which has not previously been

included in assessments of B. excelsa population dynamics therefore conclusions regarding

regeneration in fallows may better explain current B. excelsa densities and distributions than

other studies to date. An understanding of the role of fallows in B. excelsa regeneration may also

provide insight into the regeneration requirements of other valuable gap-loving species, such as

those in the Cedrela and Swietenia genera.











9y,.-


BRAZIL


x-
L ~ ACRE .


RESEX Chco Mendes r-
-pAE Chico Mendes


Figure 2-1: Location of study sites PAE Chico Mendes and RESEX Chico Mendes in Acre,
Brazil (Figure adapted from Gomes 2001, with permission).


7:30


10:00


12:30


- -*- Fallow
- Mature


15:00


Time

Figure 2-2: Mean ( SE) PPFD at four times of day in four fallows and two mature forest plots.
Data are log transformed.


2.5

2

1.5


1

0.5


--- ..- .
-^f ^ ---
















O Fallow
* Mature Forest


Seedlings Saplings

Size Class



Figure 2-3: Bertholletia excelsa seedling and sapling densities (means + SE) in fallow and
mature forest.


0.5 +


0 50 100 150 200 250

PPFD (pmol*m-*s-1)



Figure 2-4: Relationship between B. excelsa seedling height and photon flux density (PPFD)
measured in 2005. Each point represents one observed B. excelsa seedling. Regression line
fitted: y = 0.005x + 0.52 R2 = 0.31.


gO
* S
0*




* 6
so












4000
3500
3000
2500
2000
1500
1000


PPFD (pmol*m'*s1)


Figure 2-5: Relationship between B. excelsa seedling total leaf area and photon flux density
(PPFD) measured in 2005. Each point represents one observed B. excelsa seedling. Regression
line fitted: y = 7.93x + 294.97, R2 = 0.23.


Table 2-1: Mean ( sd) soil properties observed within fallows and mature forest plots in
RESEX Chico Mendes and PAE Chico Mendes.


Soil property Fallow Mature Forest

pH 5.5 ( 0.2) 5.1 ( 0.3)

K (cmol*kg-1) 0.4 ( 0.03) 0.3 ( 0.1)

P (mg*kg-1) 2.0 (+ 0.3) 2.7 (+ 0.4)

Potential Acidity
(HA1+) (cmol*kgb ) 4.0 (+ 0.7) 5.7 (+ 1.0)

Organic Carbon (%) 0.5 (+ 0.1) 0.6 (+ 0.1)


S

0

0.










Table 2-2: Mixed model results based on B. excelsa seedling and sapling data subsets.


Model fixed effects
Seedlings
Density (ha-1)
Forest type
Number of adults within 50m of subplots


Survival
Total leaf area
Percent leafherbivory*PPFD
Height growth (cm)
Initial height
Percent leafherbivory
PPFD*Percent leaf herbivory
Potential acidity (0 40cm)
Soil K (0 20cm)
Basal diameter growth (cm)
Total leaf area
Percent leafherbivory
Percent leafherbivory*PPFD
Percent leaf herbivory
Forest type
Number of leaves*forest type
Saplings
Density
Forest type
Percent organic carbon
Growth (dbh)
Forest type


F-statistic


p-value


1.09
3.71

3.22
3.26

4.99
13.09
11.93
7.55
6.21

4.26
3.42
3.58

6.96
5.84


0.33
0.06

0.08
0.08

0.08
0.02
0.02
0.04
0.06

0.05
0.08
0.07

0.05
0.006


1.80 0.23
9.25 0.02

4.55 0.07









Table 2-3: Comparison ofB. excelsa seedling and sapling characteristics in fallow and mature
forest (all p-values based upon t-tests, except one-year survivorship, which is based
upon logistic regression).
Characteristic N Fallow Mature Forest p-value
Seedlings
Initial seedling height (m) 76 0.67 ( 0.42) 0.59 (+ 0.38) 0.45
Initial seedling basal diameter (cm) 76 0.64 ( 0.34) 0.55 (+ 0.21) 0.15
Seedling height growth (m) 62 0.14 ( 0.28) 0.12 (+0.22) 0.82
Seedling basal diameter growth (cm) 62 0.19 ( 0.25) 0.10 (+0.11) 0.05
Number of leaves per seedling 75 12.8 ( 9.4) 9.7 (7.6) 0.13
Total leaf area (cm2) 68 1003.4 ( 915.4) 710.5 (+ 807.2) 0.20
Leafherbivory (%) 74 16 ( 24) 58 ( 28) < 0.0001
One-year survivorship (%) 66 74 81 0.79
Saplings
Initial height (m) 31 2.99 (+1.95) 5.51 ( 3.74) 0.02
Initial dbh (cm) 31 2.23 (+2.07) 4.45 ( 3.32) 0.05
Dbh growth (cm) 25 0.47 (+0.39) 0.11 (+0.32) 0.02
Data reported as means (+ sd), excepting percentage data. Reported sapling heights are
estimated.









APPENDIX A
TRANSECT CREATION

To open transects, a compass was placed on a level-cut branch. Next, two sapling poles

where placed in the line of sight of the compass to begin a straight line. Field technicians and

local assistants referred to this line in order to open a straight transect through the forest with a

machete. Reference poles were placed no more than every 15 m apart, with poles placed closer

together in hilly or overgrown areas. Distance was measured with a metric tape to create entire

300 m transects. Each transect was re-measured with metric tape after clearing was completed.

This method facilitated searching and locating B. excelsa individuals over large areas.












APPENDIX B
SCHEMATIC OF PLOT INSTALLATION AND SAMPLING SCHEME


*25 50 75 10 125 150 175 20 225 250 275 300
*
*







** *









N ----- ------
0 25 50 75 100 125 150 175 200 225 250 275 300






CO



0




N-
S. *
Co

CD



0-

0C


CI


0 10 20 30 40 50 60


70 80 90 100


Figure B-1: Mature forest and fallow plot design. A) Nine ha plot of mature forest, with four 25
x 25m subplots randomly selected within each ha, for a total of 36 subplots B) 1.1 ha fallow with
27 10 x 10m subplots. Distance is indicated in meters. U represents recruitment subplot. Light
environments within two mature forest plots and four fallows were characterized by photon flux

density (PPFD) measurements; represents PPFD measurement. Soil samples within each
fallow and mature forest plot were collected below the litter layer at two depths (0-20 cm and 20-
40 cm). Sampling scheme is depicted above for each forest type; @ represents soil sample.









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BIOGRAPHICAL SKETCH

Jamie Nicole Cotta was born and raised in Riverside, California, and began her career in

tropical conservation at the University of California, San Diego, where she received a B.S. in

Ecology, Behavior and Evolution. During her studies she conducted behavior observations of

bonobo chimpanzees (Pan paniscus), at the San Diego Zoo, and pygmy lorises (Nycticebus

pygmaeus), at the Center for Reproduction of Endangered Species. She next assisted a Cornell

post-doc with behavioral research on orange-fronted parakeets (Aratinga canicularis) in Santa

Rosa National Park, Costa Rica. Her undergraduate studies concluded with a semester of

Tropical Biology in Monteverde, Costa Rica, where she first became interested in sustainable

tropical forest management. She is most proud of her work as a research assistant for the School

for Field Studies, in Yungaburra, Queensland, Australia, where she managed a tropical plant

nursery and helped to design a restoration planting with a local community tree planting

organization (TREAT). Her tropical research experiences fostered an interest in tropical ecology

and forest restoration in the tropics, leading her to pursue a Masters of Science degree in

Interdisciplinary Ecology at the University of Florida. Upon graduation she plans to apply her

experience and education within an NGO setting, helping businesses and communities adopt

more sustainable practices and lifestyles.