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Ecological Review and Demographic Study of Carapa guianensis

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Ecological Review and Demographic Study of Carapa guianensis
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KLIMAS, CHRISTIE ANN ( Author, Primary )
Copyright Date:
2008

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Demography ( jstor )
Diameters ( jstor )
Ecology ( jstor )
Forest growth ( jstor )
Forest trees ( jstor )
Forests ( jstor )
Seedlings ( jstor )
Species ( jstor )
Timber ( jstor )
Trees ( jstor )

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University of Florida
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University of Florida
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Copyright Christie Ann Klimas. Permission granted to the University of Florida to digitize, archive and distribute this item for non-profit research and educational purposes. Any reuse of this item in excess of fair use or other copyright exemptions requires permission of the copyright holder.
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5/31/2007
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496626245 ( OCLC )

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ECOLOGICAL REVIEW AND DEMOGRAPHIC STUDY OF Carapa guianensis By CHRISTIE ANN KLIMAS 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 2006

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Copyright 2005 by Christie A. Klimas

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This document is dedicated to my husba nd, my parents and my grandfather Frank Klimas. Thank you for everything.

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iv ACKNOWLEDGMENTS First of all, I would like to offer my h eartfelt thanks to my advisor, Dr. Karen Kainer, for her continual suppor t and patience. I thank her fo r her constructive criticism and motivation in reading seemingly unending iterations of grant proposals and this thesis. I also want to express gratitude to my Brazilian “co-advisor” Dr. Lucia H. Wadt for her hospitality, her assist ance with my proposal, her sugg estions and support with my field research. Her support helped me br idge cultural divides and pushed me intellectually. I want to thank the members of my supervisory committee: Dr. Wendell Cropper and Dr. Emilio Bruna for their support and thorough evaluation of my thesis. I also offer my sincere thanks to Meghan Brennan and Christine Staudhammer for their statistical sugges tions and advice. I want to thank the Rotary Internatio nal Fellowship program for their initial support of my research. My experience as a Rotary Ambassadorial Fellow was a turning point in my life, both academically and pers onally. It was what helped me develop the interests that I pursued in this thesis and led me to the graduate program at Florida. I want to gratefully thank those w ho provided essential assistance during this exploratory period. My sincere thanks go to Dr. Foster Brown w ho served as my research mentor and still supports and challenges me as I progress in my research. I also want to thank Francisco (Magnesio) and Marisa dos Sant os who helped me adjust to life in Acre and introduced to me an incredible community of family and friends. I want to thank my Rotary host families, especially Maria das Graas, Jo s Eduardo, Rodrigo, Ricardo and Roberto

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v Moura Leite; Mariano, Socorro, Andr and Danielle Marques; and Marcos, Ana and Felipe Mendes. I express my heartfelt thanks to Neuza Boufleuer and Cristina Lacerda, my mentors and collaborators in Brazil. I also want to thank Andra Raposo for her collaboration and support in linke d research efforts. I also wi sh to specially thank Sumaia Vasconcelos, Paulo Wadt, Sabrina Gaspar, Elsa Mendoza, Monica de los Rios and Francisco Carlos. My project benefited from feedback fr om my friends and colleagues at the University of Florida. I would like to speci ally thank Amy Rosen, Kelly Keefe, Amanda Holmes, Chris Barloto, Cara Rockwell, Am y Duchelle, Joanna Tucker, Jamie Cotta, Wendy Pond, Monica Morris, Juliana Azoubel, Simone Athayde, Geraldo Silva, Valerio Gomes, Jensen Montambault, Roberta Velu ci, Eben Broadbent, Angelica Almeyda and Arthur Wong. I gratefully acknowledge financial support from a Tropical Conservation and Development Assistantship, an Environmen tal Protection Agency (EPA) Science to Achieve Results (STAR) master’s fellowship, an IFAS Travel Grant, a Woods Hole Field Research Grant and a Garden Club of America Caroline Thorn Kissel Summer Environmental Studies Scholarship. I cannot express how grateful I am to my family. They continue to be my biggest fans. They believe in my dreams even before I form them and are always cheering me on at the finish line. I thank my brother Geoff for your consta nt questioning. You keep me on my toes. I thank my sister Kim for your unconditional love and the generous person that you are. I thank my parents for your fina ncial support, your words of encouragement, your unwavering belief in my poten tial and the thousands of ot her things that I don’t have

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vi room to mention. I would like to thank Mark Potosnak both for helping me through this thesis and also for pulling me away fro m it. I thank him for walking me through programming basics, constantly checking on my progress, making me smile and offering words of encouragement at opportun e times. His support was invaluable.

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vii TABLE OF CONTENTS page ACKNOWLEDGMENTS.................................................................................................iv LIST OF TABLES.............................................................................................................ix LIST OF FIGURES.............................................................................................................x ABSTRACT.......................................................................................................................xi CHAPTER 1 INTRODUCTION.....................................................................................................1 2 ECOLOGY AND MANAGEMENT OF Carapa guianensis AUBLET ..................3 Abstract....................................................................................................................... ..3 Introduction...................................................................................................................3 Ecology........................................................................................................................ .5 Taxonomy and Species Description......................................................................5 Geographic Distribution........................................................................................7 Genetics.................................................................................................................8 Population Dynamics.............................................................................................9 Reproductive Ecology.........................................................................................11 Flowering and fruiting..................................................................................11 Germination..................................................................................................14 Uses.....................................................................................................................15 Seed oil.........................................................................................................15 Timber..........................................................................................................17 Management Considerations...............................................................................18 Wood quality................................................................................................18 Drought tolerance.........................................................................................19 Effects of parasites.......................................................................................20 Growth and maturation.................................................................................21 Reforestation................................................................................................22 Avenues for Future Research......................................................................................23 3 POPULATION STRUCTURE OF Carapa guianensis IN TWO FOREST TYPES IN THE WESTERN BRAZILIAN AMAZON.............................................26

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viii Introduction.................................................................................................................26 Species Description....................................................................................................28 Study Site....................................................................................................................29 Methods......................................................................................................................30 Plot Installation....................................................................................................30 Mapping Adult Individuals..................................................................................30 Estimating Regeneration.....................................................................................31 Data Analysis.......................................................................................................32 Results........................................................................................................................ .34 Adult Structure....................................................................................................34 Seedling and Sapling Structure...........................................................................36 Spatial Distribution..............................................................................................40 Discussion...................................................................................................................43 Density and Distribution......................................................................................43 Size-class Structure.............................................................................................43 Spatial Distribution..............................................................................................46 Management Implications...................................................................................47 4 CONCLUSION........................................................................................................49 APPENDIX A PLOT INSTALLATION.........................................................................................52 B RANDOM SELECTION OF REGENERATION SUBPLOTS.............................53 LIST OF REFERENCES...................................................................................................54 BIOGRAPHICAL SKETCH.............................................................................................66

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ix LIST OF TABLES Table page 2-1: A list of some common names for C. guianensis in different countries......................6 2-2: While there is little information on seed production per tree, re ported values vary widely. Below is a compilation of research on seed production..............................13 3-1: Descriptive results comparing Carapa guianensis populations in occasionally inundated and terra firme forests. Overall, comparisons between these two forest types were not different except for the per centage of individuals in one dbh class and the percentage of indi viduals in one crown class within a given dbh class.......37 3-2: Total number (N) of seedlings (indivi duals < 1.5 m tall) and saplings (individuals 1.5 m tall and < 10 cm dbh) of Carapa guianensis observed in 2004 and 2005. Number and percent of seedling and sap ling mortality and new recruits between these two periods (10 months) is also shown...........................................................39 3-3: Spatial distribution values for reproductive adults (dbh > 20 cm) and nonreproductive sub-adults (10 cm < dbh < 20 cm) in occasionally inundated and terra firme forests based on the applicati on of Donnelly’s (1978) nearest neighbor method.......................................................................................................41

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x LIST OF FIGURES Figure page 2-1: The geographic distribution of Carapa guianensis ......................................................7 2-2: Spatial distribut ion of individuals 10 cm dbh in four study plots in the Northwestern Amazon (Acre, Brazil). Th e size of the circle is directly correlated to the measured diameter of the individual it is representing.................10 3-1: Location of the study pl ots at the Brazilian Agricultur al Research Corporation’s experimental forest in Acre, Brazil (Figure adapted from Gomes 2001, with permission)...............................................................................................................29 3-2: Density of C. guianensis individuals (dbh > 10 cm) in four 16-ha study plots in two contrasting forest types......................................................................................35 3-3: Size-class distribution of C. guianensis trees > 10 cm dbh in two contrasting forest types...............................................................................................................35 3-4: Correlograms show no co rrelation between tree diam eter and nearest neighbor distance, allowing use of anova for testing statistical differences...........................38 3-5: Density of C. guianensis seedlings and saplings in tw o contrasting forest types......39 3-6: Spatial distribution of individuals > 10 cm dbh in each of the four study plots. The size of the circle is directly corre lated to the measured diameter of the individual it is representing......................................................................................40 3-7: Ripley’s K(r) analyses confirm a clumpe d adult distribution in all four plots. When L(r) (continuous line) is outside the c onfidence interval (dotted lines), it is possible to reject the complete spat ial randomness hypothesis (with a risk of =10%) in favor of regularity or clustering at distance r. Plot 2 displays middlerange (30m) and long-range aggregation (60m)......................................................42

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xi 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 ECOLOGICAL REVIEW AND DEMOGRAPHIC STUDY OF Carapa guianensis By Christie Ann Klimas May, 2006 Chair: Karen A. Kainer Major Department: Forest Resources and Conservation Both NTFPs and logging play important economic roles in Amazonian development, though not without environmenta l costs. Effective forest management, however, can mitigate some of these associated costs, particularly when based on an understanding of the ecologica l parameters under which sustainable harvest can exist. Carapa guianensis Aublet. is a tropical tree with str ong multiple-use characteristics, valued for both the high quality oil extracted fr om its seeds and as a timber resource. This thesis first provides a synthetic review of the most relevant ecol ogical and management literature of Carapa guianensis . Then, I compare population structure of this economically important rainforest tree in two contrasting forest types. Main study objectives were (a) to assess the density, distribution, and size class structure of C. guianensis in occasionally inundated and terra firme forests; (b) to compare spatial distribution between the two fo rest types, using two differe nt methodologies; and (c) to use patterns of abundance, di stribution and demographic st ructure to help infer key demographic stages or ecological variables th at merit special focus when implementing a

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xii management scheme. Four 400 x 400 m plots, two in each forest type, were established to determine distribution an d density patterns of C. guianensis 10 cm at the landscape level, and 32 10 x 10 m subplots were randomly nested within each of the larger plots to measure individuals < 10 cm dbh. Carapa guianesnsis was found at higher densities in occasionally inundated forest than in terra firme forest: 25.7 trees ha-1 and 14.6 trees ha-1, respectively. Mean density of C. guianensis individuals < 10 cm dbh was 413 trees ha-1 in occasionally inundated forests and 154 trees ha-1 in terra firme forests, but interplot variation of regeneration de nsity was high. Annual recrui tment was 94 seedlings (41%) in occasionally inundated fore st and 26 seedlings (39%) in terra firme . Both spatial distribution methods revealed a tendency to ward clumping in both forest types. High densities and clumped distributions in bot h forest types are indices favorable for sustainable species management, though reporte d growth rates for this species merit further attention. Finally, seve ral ecological variables were sufficiently different between terra firme and occasionally inundated forests to r ecommend stratification by forest type for further studies on growth and yield.

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1 CHAPTER 1 INTRODUCTION Carapa guianensis Aublet. (Meliaceae) is a ke y Amazonian species with both current and future economic potential. As with other specie s in the Meliaceae ( Swietenia macrophylla and Cedrela odorata ), C. guianensis is an important timber tree in the Neotropics (McHargue & Hartsh orn 1983, Mabberley 1987, Dayanandan et al. 1999). It is also valued for the high quality oil ex tracted from its seeds (Shanley 2005). Pure Carapa seed oil is used for medicinal applic ations (Rodrigues 1989), with value-added products including soaps, shampoos, candles a nd repellent torches (Shanley 2005). This species is considered to have such great ec onomic potential that th e Amazonian State of Acre in Brazil has identified it as one of six priority species for extraction research (Acre 2000). The oil has an international de mand. Between 1974 and 1985, Brazil exported between 200 – 300 tons of oil annually (Clay et al. 2000). In a market in Belm, the oil sells for R$15 per liter (approximately US $7). As a valuable timber and non-timber forest product (NTFP), resource exploitation is already occurring and promises to expand as demand for seed oil and timber advance into the interior of the Amazon via federallyfunded highway development projects (Fearnside 2005). Despite its economic importance, howeve r, there is limited information about C. guianensis in the peer-reviewed literature. While there are research gaps in the study of every species, there is a paucity of basic ec ological information available for this species and information on management is concentrat ed in theses and technical reports. In addition, very little is known about C. guianensis population structure, and whether

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2 structure changes across forest types. In this thesis, I first provide a synthetic review of the most relevant ecological and management literature of Carapa guianensis and comment on possible avenues for future resear ch. In the subsequent chapter, I compare the demographic structure and seedling recruitment of C. guianensis in occasionally inundated and terra firme , or upland, forests. Main study objectives were: (a) to assess the density, distribution, a nd size class structure of C. guianensis in occasionally inundated and terra firme forests; (b) to compare spatia l distribution analyses using two different methodologies; and (c) to see wh ether measured patterns of abundance, distribution and demographic stru cture can be used to help infer key demographic stages or ecological variables that merit special focus when implementing a management scheme. Measuring population structure in differen t habitats can be a first step towards assessing sustainability of ha rvesting in different habitat types (Peters 1996). Quantifying species demographic structure provides the unde rlying data for use in ecological models for assessing long-term populati on viability (Alvarez-Buylla et al. 1996). Since vital demographic rates are normally stage-depe ndent (Harper 1977), the structure of a population can be indicative of its demographic future and can be the basis for immediate management decisions when long-term demogr aphic monitoring is not feasible (Bruna & Kress 2002). This thesis is structured so that the second and third chapters are independent articles ready for submission to peer-reviewed journals. As such, relevant conclusions are found at the end of these two chapters and I articulate overa ll conclusions in the final chapter (Chapter 4).

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3 CHAPTER 2 ECOLOGY AND MANAGEMENT OF Carapa guianensis AUBLET Abstract Carapa guianensis Aublet. (Meliaceae) is a ke y Amazonian species with both current and future economic potential. As with other specie s in the Meliaceae ( Swietenia macrophylla and Cedrela odorata ), C. guianensis is an important timber tree in the Neotropics. It is also valued for the high quality oil extracted from its seeds. Despite its economic importance, however, ther e is limited information about C. guianensis in the peer-reviewed literature. The objective of this paper is to provide a synthetic review of the most relevant ecological and management literature of Carapa guianensis and comment on possible avenues for future research. Introduction Carapa guianensis Aublet. (Meliaceae) is a ke y Amazonian species with both current and future economic potential. As with other specie s in the Meliaceae ( Swietenia macrophylla and Cedrela odorata ), Carapa guianensis is an important timber tree in the Neotropics (McHargue & Hartsh orn 1983, Mabberley 1987, Dayanandan et al. 1999). It is also valued for the high quality oil extr acted from its seeds (Shanley 2005), and in a local market in Belm, Brazil, sells for R$15 per liter (approximately US $7). The oil also has international demand; Brazil exporte d between 200 – 300 tons of oil annually between 1974 and 1985 (Clay et al. 2000). Pure Carapa seed oil is used for medicinal applications (Rodrigues 1989), with valueadded products including soaps, shampoos, candles and repellent torches (Shanley 2005). This species is considered to have such

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4 great economic potential that the Amazonian Stat e of Acre in Brazil has identified it as one of six priority species fo r extraction research (Acre 2000) . As a valuable timber and non-timber forest product (NTFP), resource exploitation is already occurring and promises to expand as demand for seed oil a nd timber advance into the interior of the Amazon via federally-funded highway de velopment projects (Fearnside 2005). Despite its economic importance, howeve r, there is limited information about C. guianensis in the peer-reviewed literature. There are crucial gaps in the basic ecological information available for this species, and in formation on management is concentrated in theses and technical reports. This paucity of published ecological data on tropical species, even economically important ones, is not uncommon (Wilson 1988). For example, Swietenia macrophylla King, or mahogany, is one of the best known tropical trees, yet little is known about the ecology of Amazoni an populations. Even ba sic inventories are lacking for mahogany over most of its range (Gullison et al. 1996). A search of Web of Science’s Science Citations Index found only 128 results for Swietenia macrophylla , and an even smaller number of 47 citations for C. guianensis . In contrast, a search for loblolly pine ( Pinus taeda ), an important temperate timb er tree, produced 1,568 results. While this search was likely limited to recent literature with an English-language bias, it still demonstrates that research on C. guianensis is lacking. The objective of this paper is to provide a synthetic revi ew of the most relevant ecological and management literature of Carapa guianensis . I provide a brief description of the species drawn mostly from the work of Terence D. Pennington, a taxonomic authority for C. guianensis . I then review data from peer-reviewed publications, the relatively rich “gray” literature (Canhos et al. 1996, Lancanilao 1997), and unpublished

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5 reports and data to synthesize information on the ecology, uses and management of the species. I conclude with potential avenues for future research. Ecology Taxonomy and Species Description The first botanical description of C. guianensis was from a specimen collected in Guiana by Aublet (Aublet 1775). C. guianensis belongs to the Meliaceae family, and is a large evergreen tree. Buttresses can be abse nt or as high as 1 to 2.5 m (Fournier 2003). The bark has characteristically wide fissures with horizontal splits, and on peeling reveals a pink-red underbark (Jankowsky 1990). In oldgrowth forests at La Selva, Costa Rica, C. guianensis can attain 2 m in diameter and 45 m in height (McH argue & Hartshorn 1983). Fournier (2003) reported that C. guianensis may reach a maximum height of 60 m, though he did not cite details of this observation. More typically, however, C. guianensis reaches heights of 25-30 m (Pennington 1981). Leaf morphology of C. guianensis is well described by Pennington (1981). Leaves are paripinnate, withou t stipules and clustered at the e nd of branchlets. They usually have an apical dormant or gl andular leaflet, wh ich sometimes results in an odd pinnate leaf. Leaflets are opposite (Pennington 1981, Ferraz 2003, F ournier 2003). Its large leaves have a distinctive texture: coriace ous with a dull smooth surface and slightly intricately impressed fine venation belo w (Pennington 1981, Gentry 1993). Some sesquaterpenes (Andrade et al. 2001) and other compounds (Shu-Hua 2003) have been isolated from C. guianensis leaves and flowers, sugges ting some protection against herbivory (Trapp & Croteau 2001, Shu-Hua 2003). Though C. guianensis has been cited as Carapa macrocarpa (Ducke 1922) and Carapa nicaraguensis , Pennington (1981) and Holridge & Poveda (1975) note that

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6 neither is taxonomica lly distinct from C. guianensis . C. guianensis is morphologically similar to the taxonomically distinct C. procera , and the two can be confused as seedlings and adults. In both species, seedling germin ation is hypogeal and cr yptocotylar, and fused cotyledons and unfused petio les are present (Fisch et al. 1996). Though both have compound leaves when adult, C. procera puts out an average of six simple leaves at germination, while leaves of C. guianensis are compound at all stag es, thus providing the best distinguishing characteri stic for seedlings (Fisch et al. 1996). Adult plants are botanically distinguishabl e by inflorescence and leaf let morphology, and Pennington (1981) outlines these differences: 1. C. guianensis has sessile, subsessile flowers or very rarely short stout pedicellate flowers, predominantly 4-merous with 8 anthers, a 4-locular ovary, and (2-)3-4(-6) ovules per loculus; leaflets + elliptic, with an acute or acuminate apex. 2. C. procera has flowers that are always slende r pedicellate, predominantly 5-merous with 10 anthers (rarely with 6 petals and 12 anthers), a 5(-6)-locular ovary, and (2)3-6(-8) ovules per loculus: leaflets generally oblong with a rounded or apiculate apex. Pennington (1981) listed local names for C. guianensis , mentioning that there is not always a clear distinction between tree and timber (Table 2-1). Table 2-1: A list of some common names for C. guianensis in different countries. Country Names England, US Carapa, Crabwood Belize Bastard mahogany, Warawere Brazil Andiroba Colombia Masbolo, Tangarillo, Tangar Costa Rica Cedro bateo, cedro macho Cuba Najes Dominica Acajou Dominican Republic Cabirma de Guinea French Guiana Carapa blanc, Carapa rouge Grenada Crappo, Crappowood Guyana Carapa, Crabwood Surinam Krappa, Krappaboom Venezuela Carapa

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7 Carruyo (1972), Prance & Silva (1 975) and Fournier (2003) pr ovide more variations and additional names for these and other countries. Geographic Distribution Carapa guianensis is widely distributed from Belize along the Atlantic coast of Central America (also on the Pacific slope of Costa Rica) to S outh America throughout the Amazon Basin (Carruyo 1972, Pennington 1981, McHargue & Hartshorn 1983). The species is also found in eastern Cuba, the Dominican Republic, the Windward Islands and Trinidad and Tobago (Figure 2-1) (Pe nnington 1981, Standley & Steyermark 1946, Mhecha et al. 1984) Figure 2-1: The geograp hic distribution of Carapa guianensis . C. guianensis occupies a wide range of niches within this geographic distribution. While most individuals are restricted to altitudes between 0 and 350 m (Carruyo 1972, Pennington 1981), Pennington (1981) st ated that in Ecuador and Venezuela, especially in

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8 the state of Yaracuy, C. guianensis also occurs on mountain slopes at elevations up to 1400 m. Fournier (2003) added that C. guianensis is found above 1000 m on Guadaloupe Island. Rainfall requirements also vary widely as C. guianensis has been reported on sites receiving 1,743 mm yr-1 (UFAC 2005) and above 3000 mm yr-1, with a temperature range of 20 to 35 C (Fournier 2003). Most authors agree that C. guianensis is predominantly a species of swampy or periodically inundated land, pref erring marsh edges, swamp fo rests, alluvial riverbanks and periodically flooded plains, occasionally forming nearly pure stands under these conditions (Rizzini 1972, Pennigton 1981, McHa rgue & Hartshorn 1983, Fournier 2003). Gerry and Kryn (in Carruyo 1972) state that C. guianensis can grow in a range of sites with different soils, as long as the site is not too dry. Magalhes et al. (1986/1987), examining factors associated with species de velopment in experimental plots, however, recorded greater C. guianensis heights in areas with more argisols. In adult C. guianensis trees, extremely wet and extr emely dry periods can induce a cambial dormancy and the formation of termin al parenchyma bands; th is indicates that wet sites without inundation and without extrem ely dry periods may offer the best growth conditions for this species (Dnisch et al. 2002b). Genetics The genetic diversity of C. guianensis as measured by percent polymorphism (35%) or heterozygosity (0.12) is low in co mparison with tropical tree species (Hall et al. 1994). Even taking into account the limitati ons of their study (small number of loci, exclusion of complex loci and use of the mo st parsimonious approach in considering the number of total loci), Hall et al. (1994) considered the genetic variability of Carapa guianensis low for a highly outcrossed, long-lived tree with wide seed dispersal (Hall et

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9 al. 1994). Dayanandan et al. (1999) found no inbreeding in eith er saplings or adults of C. guainensis , but they did see a trend toward decrea sed allelic richness in the sapling cohort of an isolated fragment population. As Carapa guianensis grows predominantly in periodically inundated tropical lowland sw amps interconnected by watercourses, the movement and change of river courses over ti me would aid in the migration of floating seeds to distant populations (Rdnen et al. 1992). Dayanandan et al. (1999) developed and charac terized three polymorphic microsatellite markers for C. guianensis. Vinson et al. (2005) developed five additional microsatellite markers to use in investigati on of mating system, gene flow and paternity in C. guianensis in Par, Brazil. There are limited genetic studies of C. guianensis and a more thorough study of paternity, mating patte rns and gene flow is recommended for a better understanding of management needs. The continuation of work by Vinson et al. (2005) should provide an important first step in addressing this understanding. Population Dynamics Densities of C. guianensis can vary widely between and within regions. In Northeastern Brazil, adult densities (dbh > 15.9 cm) were estimated at 137 individuals ha1 (Sousa 1997). The median adult diameter wa s 34.8 cm and 31% of th e adults exceeded 44.9 cm (Sousa 1997). Plowden (2004) found maximu m densities of 16 trees ha-1 (dbh > 10 cm), with densities ra nging from 0 to 20 trees ha-1 depending on the habitat type. A population of C. guianensis in Acre, Brazil showed an ove rall density of 20.1 trees (dbh > 10 cm) ha-1 with a higher density (25.7 trees ha-1) in occasionally i nundated forest and a lower density (14.6 trees ha-1) in terra firme forest (Klimas, unpublished data 2004). Figure 2-2 shows the distribution of trees in the four plots fr om this study. In an unlogged population of trees > 10 cm diameter at breast height (dbh) in Acre, Brazil, mean, median

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10 and mode dbh were 25.6 cm, 22.2 cm and 11.1 cm, respectively, and no trees were found with a dbh above 80 cm (Klimas unpublished data from the Western Amazon). The 050150250350A B 050150250350 050150250350C 050150250350DMetersMeters Figure 2-2: Spatial dist ribution of individuals 10 cm dbh in four study plots in the Northwestern Amazon (Acre, Brazil). Th e size of the circ le is directly correlated to the measured diameter of th e individual it is representing. Panels A and C represent occasionally inunda ted forest and B and D represent terra firme forest. frequency distribution of this same populati on in the Western Amazon is characterized by a type-I size class distributi on, one that displays a greater number of small trees than

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11 large trees, and an almost constant propor tional reduction from one size to the next (Klimas unpublished data from the Western Amazon). Research in Northwestern Brazil found a clumped dispersion pattern for C. guianensis (Henriques & Sousa 1989) and similar research in Acre, Brazil revealed a tendency toward clumping that was more aggregated for juveniles (10 cm < dbh < 20cm) (Klimas, unpublished data, 2004). A rigorous study by Vieira et al. (2005) found that C. guianensis is older and grows more slowly than previous resear ch had suggested. Authors found that C. guianensis was among the oldest and slowest-growing of th e approximately 50 species measured. One individual of C. guianensis with a dbh of 17 cm was carbon-dated at 785 years old. Individuals that measured 37.5, 55.0, 56.0 a nd 84 cm were found to be 172, 180 + 120, 187 + 145 and 277 + 75 years old based on radiocarbon measurements (Vieira et al. 2005). Ages without error bars were determined by extrapolating growth rates determined by multiple radiocarbon measurements (Vieira et al. 2005). Reproductive Ecology Flowering and fruiting C. guianensis inflorescences are large, 20 80 cm long, branched, and axillary or subterminal. Flowers have a delicate mu sky fragrance (Pennington 1981). Petals are white or creamy with a light pink color ex ternally (Fournier 2 003). No observational research on pollinators was uncovered. In plantings at an experimental stati on, fruit production began at 10 years. In Surinam C. guianensis trees generally begin to flower when they are 6-8 years old and begin fruiting when 10-12 years old (Wille mstein 1975 from Plowden 2004). The months of flowering and fruitfall vary greatly by la titude/longitude and even within the same region (Rizzini 1971, Pennington 1981, McHar gue & Hartshorn 1983, Viana & Silveira

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12 1996, Raposo 2002, Ferraz 2003, Boufleuer et al. 2005). Fournier (2003) reports that fruits mature in eight months, although no expe rimental or observational data was cited to back up this statement. No research on envi ronmental cues that tr igger pollination or fruitfall were observed, although Pennington (1981) stated that in areas where fruitfall coincides with the rainy season, water b ecomes an important dispersal agent. The fruit is a large globulose to slightly 4-angled dehiscent cap sule with 4 valves (quadrants) that separates when the fruit falls to the ground, freeing the seeds (Gentry 1993). Each valve is woody or subwoody 5 12 cm long and 6 10 cm in diameter. There are 1-2 seeds per valve, which are a ngular due to mutual compression (Pennington 1981). Impressions on the valve wall indicat e the number of seed s contained in the segment (McHargue & Hartshorn 1983). In Brazil, Plowden (2004) found that whole fruits contained an average of 8.7 + 0.5 seeds, while at La Selva, Costa Rica, McHargue and Hartshorn (1983) report 6-7 seeds per fruit. Seeds. Individual fresh seeds usually wei gh between 25-35 grams, but seeds weighing over 100 grams have also been recorded (Hall et al. 1994). McHargue & Hartshorn (1983) reported an average dry seed weight of 15.6 grams. Despite their weight, seeds are very buoyant. Seeds l oose their viability rapidly (McHargue & Hartshorn 1983, Connor et al. 1998, Ferraz 2003). Sampaio (1999) recommends storing seeds in plastic bags in a temperature-c ontrolled area (14C and 80% RH; or 12C and 30% RH) where viability can be maintained for up to seven months. For information on seed composition, see Loureiro et al. (1979), Sampaio (1999), Revilla (2000) and Andrade et al. (2001).

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13 Production. An adequate estimate of the (potential) productivity of C. guianensis in a certain forest area requires informa tion on the seed production per tree. As C. guianensis is a masting species and variability of seed production between trees in high (Plowden 2004) (Table 2-2). McHargue and Hartshorn (1983b) reported that the C. guianensis population at La Selva produces good seed crops a bout every other year; in 1971, 1974, and 1976 almost all trees produced abundant seeds, but in 1973 and 1975 very few trees fruited. Realistic production estim ates must include a wi de cross-section of trees measured over a number of consecutive ye ars. Even these data should be replicated across regions before scaling up (or using local results that may not be relevant in distant regions). Table 2-2: While there is little informati on on seed production per tree, reported values vary widely. Below is a compilati on of research on seed production. Seed production Total Seed weight (kg) Authors Observations 754 – 3,944 11.8 – 61.5 (dry) McHargue & Hartshorn 1983 74 1.2 Plowden 2004 A few trees dominated total seed production Not reported 50 200 Shanley & Medina 2005 No studies cited to obtain this estimate Dispersal. Carapa guianensis has two predominant methods of secondary seed dispersion: water dispersal and frugivor es (McHargue & Hartshorn 1983b, Plowden 2004). When fruitfall occurs during the rainy season, this allows for seed dispersal via water. C. guianensis is preferentially found in areas that are subject to flooding, but variability in the timing of fruitfall does not allow for quantif ication of the overall importance of water as a dispersal agent. After fruitfall, McHargue and Hartshorn ( 1983b) report that 80 to 90% of the seeds are removed or eaten. The major vertebrate predators of Carapa are collared peccaries

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14 ( Tayassu tajacu ), white-lipped peccaries ( Tayassu pecari ), and large rodents such as agoutis ( Dasyprocta punctata ) and pacas ( Agouti paca ) (McHargue and Hartshorn, 1983). Based on a study by Guariguata et al. (2002), seed removal rates were uniformly high irrespective of forest site. Small excl usion cages were successf ul in reducing seed predation, suggesting that la rger animals were probabl y responsible for removing Carapa seeds. Germination Scarano et al. (2003) report phsyiolog ical variation regarding dormancy in response to seed flotation. Results showed that seed responses to floating were two-fold: they either germinated while floating, growing both shoots and roots, or germination was inhibited. As the length of floating increased, maintenance of seed viability diminished, and after 2-2.5 months, most seed s were no longer viable (Scarano et al. 2003). The germination capacity of C. guianensis seedlings is high. Published values range from 41 to 94% germination after varying time periods (Sampaio 1999, Connor et al. 1998, Guariguata et al. 2002). Germination begins 6 to 10 days from when the seeds fall from the tree, and seeds remain viable fo r 2 to 3 months (Sampaio 1999). It has been hypothesized that the sp ecies’ large seeds prov ide the energy rese rves necessary to produce a tall shoot to raise th e leaves above the normal seasonal or annual flood waters (McHargue & Hartshorn 1983b). Carapa seeds are capable of germinating in the shade, and seedlings are able to establish and grow under a cl osed canopy (Clark and Clark 1985). Seed germination studies have not cl early determined the most favorable environment for germination. McHargue and Ha rtshorn (1983) found that in well-drained soils, half-embedded and completely buried seeds had higher rates of germination. In

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15 poorly-drained swamp soils, seeds on th e surface or half-embedded had 90% germination. Neither buried seeds in the swamp soils nor surface -sowed seeds in the well-drained soils germinate d. Results from Guariguata et al. (2002) also showed a significant treatment effect on the number of Carapa seedlings that established (41% from buried and 59% from surface sown seeds). The benefits of burial by terrestrial mammals may be important for reducing dens ity-dependent mortal ity by moving seeds away from maternal fruiting trees where seed and seedling density is high (Guariguata et al. 2002), but whether it plays a si gnificant role in increasing ge rmination is still unclear. Uses C. guianensis seed oil is widely used in fo lk medicine in Brazil and other countries in the Amazon basin (Rodrigues 1989, Penido et al. 2005), with value-added products including soaps, shampoos , candles and as a repellent (Shanley 2005). The oil is also used to prepare cosmetics and as fuel for lanterns in rubber tapping communities (Boufleuer et al. 2005). Even with its value as a NTFP, Plowden (2004) asserts that Carapa ’s major economic use today is its insect -resistant reddish timber often used in place of its heavily explo ited relative, mahogany ( Swetenia macrophylla ). Seed oil Ethnographic questionnaires applied to nonindigenous Brazilian forest communities indicated that C. guianensis seed oil was used to treat arthritis; throat inflammations; to prevent insect bites; to he al insect bites, cuts , sores and bruises; diarrhea; diabetes; ear infection; as a digestive stimulant and to treat ce rvical cancer (treat pain in the vaginal region) (listed in orde r of frequency encountered) (Hammer & Johns 1993). Ferraz (2003) also reports that some indigenous groups use the bitter yellow oil from C. guianensis seeds as an insect repellent, esp ecially against mosquitoes carrying

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16 dengue and malaria. A method of obtaining a lipid extract from C. guianensis for cosmetic treatment of cellulite ha s even been patented (Rouillard et al. 1999), although no published research verifies its veracity. Hammer and Johns (1993) found significant general bioactivity in the seed of C. guianensis . The presence of active principles su ch as alkaloids, triterpenoids, cardiac glycosides, carbohydrates and ta nnins support the notion that C. guianensis is an important source of pharmacologically act ive compounds and merits further study (Hammer & Johns 1993). Mendona et al. (2005) showed that crude oil preparations from C. guianensis had high insecticidal activity with LC50 values of 57 g/l against Aedes aegypti , although identification of the component s present in the active samples that might be responsible for the larvicidal activity against A. aegypti is still needed. Miot et al. (2004) found that C. guianensis oil (100%) showed a superior profil e of repellence compared to product absence, but its repellent effect was signi ficantly inferior to DEET. These results, however, are based on only 4 subjects and open the doors for more expansive and rigorous testing of the oi l’s effectiveness as an insect repellent. Miot et al. (2004) cited research from Gilbert et al. (1999) indicating that re pellent candles made from C. guianensis oil burned for 48 hours protect ed 100% against bites of Aedes aegypti in a closed environment of up to 27 + 10 m2. The citation from Gilbert et al. (1999), however, only stated that toxicology measurements of C. guianensis candles have been made at Alfenas University, Minas Gerais, Brazil. Mo re rigorous studies are still necessary to confirm results from these studies and bette r analyze the effectiveness of the oil’s repellent strength.

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17 To obtain the oil, seeds are boi led in a large pot of water and left for 8 to 15 days to induce fermentation. Seeds are then shelled, and the pulp (seed endosperm) is mashed and kneaded by hand. The seed mass is then set out on a sloping sheet. Over the course of a week, the oil drips down a cloth wick into a collecti ng jar (Plowden 2004, Boufleuer et al. 2005). In addition to this method, extraction of oil can be done using small presses and filters more commonly used to extract oil from Brazil nuts or can be done on an industrial scale. Plowden ( 2004) found that it would take 14.43 kg of seeds to produce one liter of oil. Timber While the oil market is well-developed, C. guianensis ’s major economic use today is its timber (Plowden 2004). The wood of Carapa is moderately heavy (0.70 to 0.75 g/cm3) with bright reddish-brown coloring. It is similar in many respects to that of true mahogany, but it is harder and heavier and lacks the lustre and color of mahogany (Pennington 1981). According to Pennington (1981), two types of C. guianensis timber are recognized by foresters: red and white. Red or Hill “crabwood” is said to be superior to white and is obtained from trees growi ng on higher land. White “crabwood” is derived from those growing on swampy flat ground. Carruyo (1972) and Jankowsky (1990) provide a more complete list of the physic al properties of the wood. Timber from C. guianensis is considered variable; laboratory test s report both high and low resistance to brown and white rot fungi. This variability ma y stem from the two different timber types. The wood is easy to work and allows for a good finish. It is much sought after for construction of furniture, detailed boxes for jewelry , construction, window frames and doors , dividing walls, shutters, doorframes, molding , thin wood leaf , chipboard , and the final woodwork for boats and ships (SUDAM, 1979 from Sampaio 1999; Souza, 1997).

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18 The Swedish Forest Products Research La boratory has conducted a wide array of studies on the wood of C. guianensis and other species and is a valuable starting point for research on the wood properties of C. guianensis (Boutelje 1980). Fina lly, the wood is an excellent fuel source due to a high ignition temperature a nd therefore slow combustion (SUDAM, 1979 from Sampaio 1999; Sousa, 1997). Management Considerations As with most tropical species, knowle dge about the site demands and the appropriate management of timber pl antations is still limited (Dnisch et al. 2002b), and this is also true for management of C. guianensis in natural forest. Studies in plantations and natural forest have begun to identify fact ors that may affect future management of C. guianensis . Wood quality The low variability in wood density, fl avonoid content that gives rise to C. guianensis’s decorative color, and a high fibre content all contribute to C. guianensis ’s wood quality. Bauch & Dnisch (2 000) found that these charact eristics are maintained in plantation grown trees. C. guianensis produces mature wood (hear twood) at a very early age (four years in plantations), compared with development of adult wood in other tropical hardwoods (Bauch & Dnisch 2000). As heartwood is very important for furniture and veneer wood, this finding is promising for high-quality timber production under suitable plantation conditions (B auch and Dnisch 2000), though seeds and seedlings are susceptible to attack by parasi tes and these attacks ar e more prevalent when grown under plantation conditions (see below).

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19 Drought tolerance C. guianensis , a pioneer species, exhibited diffe rent functional gas exchange and hydraulic traits than later successional lowland rainforest species in French Guiana (Huc et al. 1994). The authors believe that these re sults may reflect a competitive ability for water and nutrient uptake in the absence of soil drought. Maximum water uptake in C. guianensis was found in soils with a soil wate r potential of to kPa (Dnisch et al. 2002c). C. guianensis utilizes a strategy that increa ses its net assimilation rate under drought conditions compared to Swietenia macrophylla and Cedrela odorata . Trees use stored xylem water for transpiration during drier periods. While the absolute amount of stored xylem water is sufficient to compensate only short-term daily water deficits, good conditions for refilling xylem water at night of ten exist in tropical forests. Xylem water storage thus positively influences photosynthe sis during drier periods (Dnisch & Morais 2002). Dnisch and Puls (2003) found that C. guianensis favors the compensation of unfavorable hydrological conditions to mainta in growth during the entire year while related species ( Swietenia and Cedrela ) depend on a high water supply of the soil. Morphological and anatomical studies on the structure of the roots and leaves also showed a better capacity for the regula tion of water and nutrient uptake of C. guianensis compared to Swietenia and Cedrela (Dnisch et al. 1999). Noldt et al. (2001) were able to demonstrate special strategies to resist drought in by analyzing the anatomical structure and chemical characteristics of the fine roots of C. guianensis . They found that C. guianensis exodermis develops cells especi ally well adapted to drought. These adaptations to drought are very impor tant for long-term survival of C. guianensis and with increasing evidence of current and future droughts in the Amazon (Nepstad 1999), these adaptations may select for survival of this species under changing climate

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20 conditions. They may also facilitate mana gement under a variety of conditions and facilitate management in areas wh ere water limitation is a factor. Effects of parasites C. guianensis seeds and seedlings are susceptible to attack by the parasite ( Hypsipyla sp.) (Pennington 1981). Becker (1973) records larvae of H. grandella Zell. from the shoots and those of H. ferrealis (Hamp.) from the seeds of C. guianensis . Plowden (2004) also mentions that seeds are susceptible to attack by fly maggots. Attack by H. ferrealis causes forking in seedlings a pr oblem for timber production, especially since this parasite is more prevalent when trees are planted in full sunlight (Carruyo, 1972). While full sunlight has been stated as the cause for attack, no experiments have been conducted to determine percent attack with different light availabilities. Other confounding variables such as soil moisture or planted seedling density may have more of an effect on seed and seed ling susceptibility to attack. Infested seeds are recognized by the sawdustlike substance coming out of 1-3 mm diameter holes in the seed coat through which larvae emerge. Seeds can sustain some seed damage by moths and still germinat e (McHargue & Hartshorn 1983). Although I am not aware of any literature that speaks to th e effect of seed infestation on oil quality, depending on how long the larvae had been feed ing on the seed, larval infestation could reduce the quantity of seeds available for oil production. Soaking seeds in water for 14 days (changing the water daily ), then germinating them in sealed clear plastic bags drowns Hypsipyla sp. and allows the seeds to become turgid (Fisch et al. 1996, Ferraz 2003).

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21 Growth and maturation Due to formation of vessel bands in the juvenile wood (false rings), C. guianensis is not suited to dendroeco logical studies (Dnisch et al. 2002b, Dnisch et al. 2002a). This makes determining tree age difficult, part icularly since growth rates are variable between regions and forest habitats. A group of planted trees at La Selva, Costa Rica measured 10-15 cm in diameter, about 7-10 m tall after 6 years of growth, the same age as a group of shaded seedlings in a nearby sw amp, most of which are below 1 m in height (McHargue & Hartshorn 1983b). In plantings done at the Experimental Station of CuruUna/PA, in full sunlight, with 2.5 x 2.5 m spaci ng and 80% survival, trees averaged a 1.8 m yr-1height increase and a 1.10 cm annual dbh increase (Willemstein 1975 from Plowden 2004). Growth rates also differ between forest and plantations. At about four years of age, growth rates of C. guianensis in primary forest was only appr oximately half that of trees from unfertilized plantations. Plantation grown C. guianensis exhibited rapid growth throughout the year with annual growth of about 1.14cm. The intra-annual pattern of C. guianensis in the primary forest differed with an annual diameter growth of only 0.246 cm (Bauch and Dnisch 2000). The high variab ility of individual growth rates makes it difficult to use published growth rates from one region in another region, or even a neighboring site. This difficulty is enhanced if there are large differences between the sites (i.e. plantation vs. natural forest). Measurements of the soil water supply in primary forest indicate that high soil water content can lead to growth depressi on or even short dormancy periods (Dnisch et al. 1999), although no dormancy was observed during the dry s eason (Bauch and

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22 Dnisch 2000). Positioning plantations in areas that are not subject to periodic inundation may result in increased annual growth. Variation in diameter and height increm ent due to light environment and other microenvironmental conditions is problematic for estimating timber yield and age to first reproduction in natural forest. As determined in Vieira et al. (2005), small diameter trees can survive in the understory for hundreds of years, making it difficult to estimate cutting cycles and timber yield. This environmental variation also makes it difficult to predict when trees will reach a size that supports seed production. More research is necessary to better determine average growth rates under a forest canopy and limiting microenvironmental factors that may limit growth. Reforestation Carapa guianensis was suggested as a species that holds promise for reforesting secondary forests (Sampaio 1999). Dnisch et al. (2002c) recommend it for the restoration of degraded areas. The physiol ogical adaptations to drought mentioned above (Huc et al. 1994, Dnisch et al. 1999, Dnisch & Morais 2002, Dnisch & Puls 2003), the high germination rates (Connor et al. 1998, Sampaio 1999, Guariguata et al. 2002) and its high survival in plantations (Willems tein 1975 from Plowden 2004) are suggest its potential for establishment and survival. It also showed promise for reforestation in pasture and grasslands. Carapa guianensis had fast germination, high survival and relatively high performance when planted with the exotic grass Saccharum spontaneum . It also had the ability to resprout after fire (Hooper et al. 2002). While the authors did not mention attack by H. ferrealis , it is possible that they were concerned more with rapid establishment and growth than with adult tree form.

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23 Dnisch et al. (2003) do not recommend monocu ltures of plants with low transpiration rates, such as C. guianensis , in the Central Amazon due to high water loss from interception and water runthrough in young monoculture stands. Authors instead recommend enrichment plantations where th e positive effect of secondary vegetation stabilizes water fluxes. Avenues for Future Research While authors have touted the economic potential of C. guianensis in the literature, very little research has been done to quantify this potential or to look at sustainable harvest limits/challenges. This se ction outlines data still lacking to make informed decisions about sust ainable management of this species as both a timber and non-timber resource. According to Hall and Bawa (1993), knowledge of the natural distribution, abundance, populati on structure and dynamics, and variation of these factors across a landscape is required in order to assess the sustaina bility of resource harvesting. Permanent plot networks in tropical fore sts may provide some information on these variables for C. guianensis (Losos & Leigh Jr. 2004, Viera et al. 2005), but in these cases, C. guianensis may form only a small percentage of the inventoried trees. A recent article examines the population structure and spat ial structure of this species in two forest types in Acre, Brazil (Klimas, in preparat ion). This study, however, fails to provide information on the size-specific growth rate, fecundity and mortality of this species or otherinformation on species dynamics, inform ation that is essential for simple demographic models (Leftkovitch 1965). Using genetic and evolutionary models in combination with demographic models may provide a more realistic projection of th e efforts necessary for species management. These models incorporate loss of genetic variability for adaptive evolution, random

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24 fixation of deleterious mutations or allele s by genetic drift and inbreeding depression (Hedrich & Miller 1992, Ells tand & Elam 1993). Only preliminary genetic studies exist for C. guianensis . The continuing work of Vinson et al. (2005) will allow a better understanding of management needs, and recent genetic surveys of C. guianensis in Acre, Brazil (Raposo) will provide valuable information on paternity and mating patterns. To truly capture the risks of management, howev er, estimates of effective population size, inbreeding coefficients and the genetic variation are important (Alvarez-Buylla et al. 1996). Basic ecological information on this speci es is still unavailable. Research on pollinators, a key management variable is lacking. Silvicultural experiments to ascertain methods for improving growth rates, tree fo rm and increasing seed production per tree would also provide valuable management data for both plantation and forest-grown Carapa guianensis . Methods of controlling H. ferrealis and H. grandella infestations in seedlings and saplings, respectively, and ways to promote/improve survival and growth of young C. guianensis individuals may also bear inves tigation. Experiments that explore underlying causes for seedling susceptibility to parasite attack would permit better management of early plantations. Seed collection will likely affect seed pr edators. More research determining the broader ecosystem impacts of seed and timbe r collection may be beneficial. Similarly, broader faunal resources such as game meat are critically important for human inhabitants (Hill et al. 2003). The linkages between seed populations and seed predators is important to quantify (Forget 1996), both fo r ecologists and forest residents who rely on these animals for sustenance.

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25 Economic and social analyses of co mmercialization barriers for small scale landowners is a key step toward utilization and management of this resource. Even with preliminary information on the market value of seed oil in different areas of the Brazilian Amazon (Shanley 2005), the costs of transp ortation to the nearest market, labor, investment in equipment and the trade-offs with current economi c activities have not been fully analyzed, though see Plowden (2004) for preliminary labor estimates. From the forest to the final product, there is still a wealth of information yet to be discovered to better advise ecosystem managers and commercial producers of this species.

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26 CHAPTER 3 POPULATION STRUCTURE OF Carapa guianensis IN TWO FOREST TYPES IN THE WESTERN BRAZILIAN AMAZON Introduction The Amazon basin contains over half of th e world’s remaining tropical rainforest, and is facing unprecedented changes that will have major impacts on biodiversity, regional hydrology and the gl obal carbon cycle (Nepstad 2001, Fearnside 2005). Humans play a significant role in these changes, of ten as part of a stru ggle to improve their standard of living (Schmink 1994, Wood 2002). Us e of forest resources is frequently necessary for improving the social and econom ic living conditions of forest residents (reviewed in Ticktin 2004) and may add to the perceived value of standing tropical forest (Arnold & Perez 2001). Logging and non-timbe r forest product (NTFP) extraction, however, incurs associated environmenta l costs (Peters 1996, Nepstad 1999, Peres et al. 2003). Effective forest management can mitigate these costs, particularly when based on an understanding of the ecological parameters under which sustaina ble harvest can exist (Putz et. al 2000). In contrast, management activi ties that ignore the regeneration and growth requirements of the species under exploitation have little chance of long-term success (Hartshorn 1995, Peters 1996). While there is still considerable scientific debate over whether natural resource extraction can be sustained over the long te rm and how to accomplish this (Nepstad 1999, Peres et al. 2003, Pearce et al. 2003, Fearnside 2005), the debate is hindered by the lack of demographic information for most tropical species (Gullison et al. 1996, Zuidema

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27 2003). To sustain, or even increase the a bundance of an extracted product, detailed knowledge of species life history and demogr aphic behavior is e ssential (Sunderland and Dransfield 2002). Peters (1996) specifies four key ecological parameters for guiding sustainable management of a ny given species: life cycle characteristics, multiplicity of uses and types of resources produced, abundance in different forest types, and size-class distribution of populations. Patte rns of abundance, distributio n and demographic structure can be used to help infer key demographic stages or ecological variables that merit special focus when implementing a management scheme (Bruna & Ribeiro 2005). Since these variables can differ by habitat type (Bruna & Kress 2002, Wa gner & Fortin 2005), they should be evaluated for any given species in the multiple habitats or forest types in which they occur. Carapa guianensis is valued for both the high quality oil extracted from its seeds (Shanley 2005) and as a timber resource (Dayanandan et al. 1999). Pure C. guianensis seed oil is used for medicinal purposes (R odrigues 1989), as well as in products such as soaps, shampoos, candles and insect-repel lent torches (Boufleuer 2001, Shanley 2005). C. guianensis ’s major economic use today, however, is as insect-resistant timber often used in place of its heavily exploited relative, Swietenia macrophylla King, commonly known as mahogany (Plowden 2004). Very little, however, is known about C. guianensis population structure, and whether struct ure changes across forest types. Measuring population structure in different habitats is a first step towards assessing sustainability of harvesting in different habi tat types (Peters 1996) . Population structure in most forestry and ecological studies has been defined in terms of the size-class or diameter distribution of indi viduals, with frequency histog rams showing the number or

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28 percentage of individuals in each size class (Knight 1975, Peters 1996). Quantifying species demographic structure provides data for ecological models assessing long-term population viability (Alvarez-Buylla et al. 1996). Since vital demographic rates are normally stage-dependent (Harper 1977), the st ructure of a populati on can be indicative of its demographic future and can be the basis for immediate management decisions when long-term demographic monitoring is not feasible (Bruna & Kress 2002). We compared the demographic structure and seedling recruitment of C. guianensis in occasionally inundated and terra firme , or upland, forests. Main study objectives were: (a) to assess the density, distri bution, and size class structure of C. guianensis in occasionally inundated and terra firme forests; (b) to compare spatial distribution between the two forest types, using two diffe rent methodologies; and (c) to use patterns of abundance, distribution and demographic structure to help in fer key demographic stages or ecological variable s that merit special focus when implementing a management scheme. Species Description C. guianensis is a medium to large hardwood tree that can attain 2 m in diameter and 50 m in height (Pennington 1981, McHar gue & Hartshorn 1983). Most authors agree that C. guianensis is a predominantly pioneer species of wet areas, although it is also found on a variety of drier sites (Pennigt on 1981, McHargue & Ha rtshorn 1983, Fournier 2003). It is found in the West Indes, Antill es, Central America south of Honduras, many parts of the Amazon region, and trop ical Africa (Smith 1965, Pennington 1981, McHargue & Hartshorn 1983). In Br azil, it is more commonly referred to as “andiroba”, and Pennington (1981) provides a reference to other common names for this species. The fruit is a spherical or subspherical capsule of dry dehiscent fiber w ith 4 valves (quadrants)

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29 that separate and open upon falling to the ground, freeing the seeds. It has two predominant methods of secondary seed di spersal: water and frugivores (McHargue & Hartshorn 1983b, Plowden 2004). Each valve co ntains between 1 and 4 seeds (Smith 1965, McHargue & Hartshorn 1983, Sampaio 2000). Annual seed production in this species is variable. Based on a 5-year study, McHargue and Hart shorn (1983) reported that C. guianensis produces good seed crops almost ev ery other year. In Acre, forest residents and technicians re port a 2-3 year masting cycle (personal communication). Study Site Field surveys were carried out within the 1,200 hectare e xperimental forest of the Brazilian Agricultural Research Corporation (Embrapa) in the northeastern portion of the state of Acre, Brazil (Figure 3-1). The study region has lightly undulating topography, with dominant vegetation classified as humi d, moist tropical forest (Holdridge 1978). The region has a pronounced 3-mont h dry season from June to August. The mean annual temperature is 24.5. Brief intrusions of cold air from the South occasionally drop ACRE BRAZIL Study site ACRE BRAZIL Study site Figure 3-1: Location of th e study plots at the Brazili an Agricultural Research Corporation’s experimental forest in Acre, Brazil (Figure adapted from Gomes 2001, with permission).

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30 temperatures to 10C during the dry season (ZEE 2000). During the 2004-2005 study period, the maximum and minimum temperatur es, respectively, were 35C and 16C; relative humidity ranged from 71 to 91% with an average of 86%; and total rainfall was 2,089 mm in 2004 and 1,743 mm in 2005 (UFAC 2005). Weather data were collected 25 km from the study site. Soils in the occasionally inundated forest were red and yellow ultisols; soils in the terra firme forest were plintosoils (Rodrigues et al. 2001). Methods Plot Installation Four 400 x 400 m (16-ha) plots were esta blished from June through July 2004 to determine distribution an d density patterns of C. guianensis at the landscape level. Two plots were installed in areas where the majo rity of the environment was classified as “ terra firme ”, or upland forest and two in occasiona lly inundated forest, defined as level or concave areas on poorly drained ground or areas subject to periodic flooding. Terra firme was considered to be convex or level ground in well-drained areas (Azevedo 2005). Internal transects were installed every 50 m to create eight 400 x 50 m rectangles. A central transect was used to bisect these rectangles. These internal divisions created a grid system used to map C. guianensis individuals. Appendix A contains additional information on the sampling design. Mapping Adult Individuals C. guianensis trees > 10 cm diameter at breast height (dbh) were inventoried in all 4 plots. A minimum of two researchers loca ted trees by systematically walking parallel to the transect lines. After locating a tree, the exact y-coordinate was recorded based on a tape-measured distance. The x-coordinate was visually estimated based on distance to the proximate transect. This accuracy of th e x-coordinate was tested by measuring the

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31 distance with a meter tape and an error of + 5 m was attained before continuing. In addition to the coordinates, plot numbe r, canopy position (dominant, co-dominant, intermediate or suppressed) from Smith et al. (1997), dbh, microenvironment ( terra firme or occasionally inundated) and reproductive status were recorded. Reproductive status was positive if there were seeds or associated dehiscent capsules located either on the tree or on the ground underneath the tree or if there was a seedling bank from a former year that could only be clearly attributed to that tree. While at the landscape scale, each of the 4 plots was assigned to one of two forest types ( terra firme or occasionally inundated forest), we also assessed the “microenvi ronment” immediately surrounding each tree (approximately a 1-2 m diameter), assigning each individual to a microenvironment (again, either terra firme or occasionally inundated). Estimating Regeneration Thirty-two 10 x 10 m subplots were random ly nested within each of the larger plots to measure individuals < 10 cm dbh in August and October, 2004 (Appendix B). Subplots were only selected fo r study if the entire 100 m2 area belonged to the same microenvironment as the larger, designated fo rest type. For example, only subplots that were clearly and totally defined as terra firme were accepted for study as a terra firme regeneration subplot. If a selected subplot fe ll in a lowlying area or transition zone, it was removed from the sample and another subplot was randomly selected for consideration. Selected subplots with trails or rivers within their boundaries were also discarded. Within each subplot, all seedlings (ind ividuals < 1.5 m tall) and saplings (all individuals 1.5 m tall and < 10 cm dbh) were tagged and x,y coordinates noted. For seedlings, basal diameter (at the level of the soil) and height were measured. For saplings, dbh was measured and height was estimated. While the seedling classification

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32 may have captured small saplings and could have been further divided, my goal was not to differentiate between those individuals stil l using their seed reserves for growth and those individuals relying on environmental re sources, but to look at general classes of recruitment. All tagged seedlings and sapli ngs were remeasured to determine 10-month mortality, and diameter (basal or dbh) and height growth. Data Analysis Since analysis of variance (Anova) cannot be used if autocorrelations are present, we used correlograms from the ‘spatial’ package in R to test for spatial autocorrelation of tree diameter as a function of distance (Ven ables & Ripley 2002). Correlograms divide a range of data into bins and computes the average squared differences for pairs with separation in each bin. Results are returned only for those bins with 6 or more pairs; 50 bins were selected per plot. A logit mode l, in R described as binomial proportion comparisons, was used to test for differe nces in the binary production values and differences between size classes in the different fo rest types. Following this test, Anova was used to test for differences in adult, sapling and seedling densities and adult diameter dist ribution. Forest type and microenvironment were treated as fixed effects. Nesting wa s used for the seedling and sapling anovas. Tukey’s HSD was used for all applicable multiple comparisons. A two-sample test of proportions was used to compare percentage values for tree reproduction of the various dbh classes. In this case, a bonferroni co rrection was used on the alpha level. R programming language was used for all anal yses and a p-value of 0.05 was considered statistically significant. I characterized the spatial distribution of C. guianensis using the aggregation index R (Clark & Evans 1954), corrected for edge effects (Donnelly 1978). The aggregation

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33 index is based on measurement of nearest neighbor distances for each individual and provides an indication of whether a popul ation has a clumped, random or uniform distribution. Spatstat, an R pack age for spatial point pattern analysis (Baddeley & Turner 2005) was used to calculate nearest nei ghbor distances based on the x,y coordinate reference system. I used the calculated zvalue and associated p-value to determine whether the observed distributi on was significantly different than the expected random pattern. This analysis was comp leted for individuals (10 cm < dbh < 20 cm), hereafter termed sub-adults, and reproductive adults (dbh > 20 cm) in each experimental plot for the purpose of exploring whether sub-adults have a different spa tial distribution than reproducing adults. These subdivi sions were used since results showed that the majority of individuals dbh > 20 cm had evidence of reproduction, while this was not the case for individuals < 20 cm dbh. Plow den (2004) found similar size-based production estimates. Anovas tested significant differences in neares t neighbor distances be tween forest types. The use of first nearest neighbors, howeve r, does not differentiate an aggregated distribution from an even distribution of regul arly sized clumps, and information is lost (Cressie 1993, Dale 1999). Therefore, Ripley’s K(r) function (Ripley 1977 from Goreaud et al. 1999) and the edge correction factor propos ed by Ripley were also applied to the data. Ripley’s K(r) function determines the ex pected number of neighbors in a circle of radius ds centered on an arbitrary tree in the point pattern. This circle begins at a specified radius and is increased until it encompasses the entire study region. The expected number of neighbors in the circle is defined as *K(r). The intensity, , is the expected number of points per unit area = N/S; where N is the number of points in the pa ttern and S is the study region area.

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34 K(r) = 1/ * 1/N * kij; where kij=1 if the distance between i and j is less than r, and 0 otherwise. Since the number of trees should increase with an increasing circle radius, the linearized function L(r) proposed by Besag (1977) was used to simplify pattern interpretation such that: L(r) = (K(r)/(1-r)). For a Poisson pattern, L(r) = 0 at every dist ance r; for clustered patterns at distance r, L(r) > 0; and in the case of regul arity at distance r, L(r) < 0 (Goreaud et al. 1999). This analysis included all trees > 10 cm dbh. Confidence intervals were estimated usi ng the Monte Carlo method. One thousand Poisson patterns were simulated and the confid ence interval was defined for each r so that only the highest 5% and the lowest 5% of L(r) values were outside the interval (Goreaud et al. 1999). Results for L(r) were graphed w ith their corresponding confidence interval for all plots. If the graphed L(r) functions fo r field data remained within the confidence intervals, it demonstrated a random distri bution of individuals. Spatstat was also employed for all calculations of Ripley’s K(r) with the “iso” edge-co rrection factor used for the adjustment of edge eff ects (Baddeley & Turner 2005). Results Adult Structure Almost twice as many C. guianensis trees > 10 cm dbh were encountered in occasionally inundated versus terra firme forests (822 and 466, respectively). Thus, tree densities were higher in the fo rmer than the latter (25.5 + 0.2 and 14.5 + 2.7 trees ha-1, respectively) (p = 0.056) (Figure 3-2). Fo r both forest types the distribution of

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35 0 5 10 15 20 25 30 Plot 1Plot 3Plot 2Plot 4 Occasionally inundated Terra firme Forest typeAdult density (trees > 10 cm ha-1) Figure 3-2: Density of C. guianensis individuals (dbh > 10 cm) in four 16-ha study plots in two contrasting forest types. individuals in dbh classes reve aled a classic j-distribution with a higher number of subadults (10 cm < dbh < 20 cm) and smaller diameter tree s and a decrease in the number of individuals in the larger di ameter classes (Figure 3-3) . Further binomial proportion 0 50 100 150 200 250 300 350 400 203040506070> 70U pp er limit of dbh classes ( cm ) Frequency (number of individuals) Occasionally inundated Terra firme Figure 3-3: Size-cla ss distribution of C. guianensis trees > 10 cm dbh in two contrasting forest types.

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36 comparisons indicated that the percentage of individuals per fore st type only differed with respect to forest type in the 10 to 20 cm dbh class (p < 0.001) (Table 3-1). Correlograms showed no correlation between tree diameter and neares t neighbor distance (Figure 3-4). Anova tests were thus appropriate for data analysis, and revealed that average dbh was significantly higher in the occasionally inundated versus terra firme forest (p = 0.031). The number of reproductive trees was significantly higher in occasionally inundated forest than in the terra firme forest (p = 0.003). There was, however, no significant difference in the percenta ge of reproducing individuals of C. guianensis in each dbh class between forest type s and the majority of trees < 20 cm in both forest types did not show evidence of reproduction (Table 3-1). In the occasionally inundated and terra firme forest plots, respectively, 52 and 69% of all adult trees > 10 cm dbh were in the dominant or co-dominant categories, and it was almost entirely these trees that were reproductive. Finally, assuming reproduction at dbh > 20 cm, the non-reproductive to potentially-reproductive ratio in the occas ionally inundated forest was significantly different than in the terra firme forest (8.44 and 5.79, respec tively) (p = 0.025). Thus, compared with the occasionally-inundated forests, for every reproductive adult in terra firme forest,, there are fewer corre sponding non-reproductive sub-adults. Seedling and Sapling Structure As with individuals > 10 cm dbh, C. guianensis seedling densities were significantly higher (almost triple) in occasionally inundated plots than in terra firme plots (p = 0.006), while sapling densities di d not differ by forest type (Figure 3-5).

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37 Table 3-1: Descriptive results comparing Carapa guianensis populations in occasionally inundated and terra firme forests. Overall, comparisons between these two forest types were not different except fo r the percentage of individuals in one dbh class and the percentage of individua ls in one crown class within a given dbh class. Diameter class Crown class a (Percentage in each diameter class) Forest type Percent of individuals (%) D CD I S Reprod. indiv.b (%) 10 < dbh < 20 40.0*** 1 12 45 42 11 20 < dbh < 30 30.3 4 49*** 39 8 74 30 < dbh < 40 18.7 14 73 11 2 98 40 < dbh < 50 7.8 28 69 2 2 98 50 < dbh < 60 2.7 55 45 0 0 95 60 < dbh < 70 0.4 67 33 0 0 100 Occasionally inundated dbh > 70 0.1 100 0 0 0 100 Percentage of all individuals > 10 cm dbh 20 32 8 40 56 10 < dbh < 20 50.2 1 8 46 45 7 20 < dbh < 30 24.7 1 26 56 17 74 30 < dbh < 40 16.5 10 64 21 5 97 40 < dbh < 50 5.2 33 46 21 0 100 50 < dbh < 60 2.4 82 18 0 0 100 60 < dbh < 70 0.9 75 25 0 0 100 Terra firme dbh > 70 0.2 100 0 0 0 100 Percentage of all individuals > 10 cm dbh 28 41 7 24 53 aCrown position: D=Dominant; CD=Co-dom inant; I=Intermediate; S=Suppressed bReproductive individuals *** p = 0.0005

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38 0100200300400500 -1.0-0.50.00.51.0 Distance (m)Correlation A 0100200300400500 -1.0-0.50.00.51.0 Distance (m)Correlation B 0100200300400500 -1.0-0.50.00.51.0 Distance (m)Correlation C 0100200300400500 -1.0-0.50.00.51.0 Distance (m)Correlation D Figure 3-4: Correlograms show no correla tion between tree diameter and nearest neighbor distance, allowing use of anova for testing statistical differences. Panels A and C represent occasionally inundated forest and B and D represent terra firme forest.

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39 Figure 3-5: Density of C. guianensis seedlings and saplings in two contrasting forest types. Seedlings or saplings were found in 81% of the occasionally inundated subplots, but only 51% of the terra firme subplots. Occasionally inundated fo rest also had greater rates of seedling recruitment (p = 0.001) and mortal ity (p = 0.017) after a 10-month period, but percent recruitment and mortality were not di fferent between forest types (Table 3-2). These same sapling parameters did not differ wi th respect to forest type (Table 3-2). Table 3-2: Total number (N) of seedlings (individuals < 1.5 m tall) and saplings (individuals 1.5 m tall and < 10 cm dbh) of Carapa guianensis observed in 2004 and 2005. Number and percent of s eedling and sapling mortality and new recruits between these two pe riods (10 months) is also shown. Seedlings Saplings Occasionally Inundated Forest Terra Firme Forest Occasionally Inundated Forest Terra Firme Forest Total N (2004) 231** 66 30 32 Total N (2005) 248 73 28 31 Mortality: N (%) 77* (33.3) 19 (28.8) 3 (10.0) 1 (3.1) New Recruits: N (%) 94** (40.7) 26 (39.4) 1 (3.3) 0 * p < 0.05 ** p < 0.01 0 50 100 150 200 250 300 350 400 450 500SeedlingsSaplingsDensity (Individuals ha-1) Occasionally inundated Forest Terra firme Forest

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40 Spatial Distribution Spatial distribution of reproductive adults (dbh > 20 cm) was aggregated in all forest plots (Figure 3-6) (T able 3-3). Based on applicati on of Donnelly’s (1978) nearest neighbor method, the index of aggregation (R) for reproductive adults in occasionally inundated forests was statistically the same as that for the terra firme forests (0.9056 and 050150250350A B 050150250350 050150250350C 050150250350DMetersMeters Figure 3-6: Spatial distribution of individuals > 10 cm dbh in ea ch of the four study plots. The size of the circle is directly corre lated to the measured diameter of the individual it is represen ting. Panels A and C repres ent occasionally inundated forest and B and D represent terra firme forest.

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41 0.8258, respectively). These values indicate a reje ction of the null hypothesis of a strictly random pattern of distribution in all the plots. Table 3-3: Spatial distribution va lues for reproductive adults (dbh > 20 cm) and nonreproductive sub-adults (10 cm < dbh < 20 cm) in occasionally inundated and terra firme forests based on the applicati on of Donnelly’s (1978) nearest neighbor method. Forest type Plot Na Rb p-value Average distance between trees (m) (_x SE) Adults 1 264 0.9010 <0.0002 11.1 + 0.4 Adults 3 229 0.9006 <0.0003 11.9 + 0.5 Sub-adults 1 151 0.7586 <0.0001 12.4 + 0.8 Occasionally inundated Sub-adults 3 178 0.9116 <0.0020 13.7 + 0.6 Adults 2 139 0.9152 <0.0060 15.5 + 0.8 Adults 4 93 0.7256 <0.0001 15.1 + 1.3 Sub-adults 2 138 0.8891 <0.0013 15.1 + 0.8 Terra firme Sub-adults 4 96 0.5942 <0.0001 12.1 + 1.2 a Number of individuals. b Index of aggregation: R = 1 if the spa tial pattern is random; R = 0 when clumping occurs; and R > 2.15 when a uniform dist ribution pattern exists (Krebs 1999). When analyzed separately, sub-adults (10 cm < dbh < 20 cm) exhibited a greater level of aggregation than repr oductive adults (Table 3-3). All plots, irrespective of forest type, were significantly differe nt from a random distribution. Plot 4, representative of a terra firme forest, and which was dominated by a bamboo forest in a pproximately half the plot, had the highest le vel of aggregation. Results from Ripley’s K(r) confirm a clum ped distribution in all four plots (Figure 3-7). For this analysis, sub-a dults and reproductive adults we re analyzed as a unit. When L(r) is outside the confidence interval, it is possible to reject the complete spatial randomness hypothesis (with a risk of =10%) in favor of regularity or clustering at

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42 distance r (Goreaud et al. 1999). In the terra firme forest, plot 2 indicates middle range (30 m) and longer range aggrega tion (60 m) (Figure 3-7). 020406080100 -4-2024 rL(r) A 020406080100 -2-1012 rL(r) B 020406080100 -6-20246 rL(r) C 020406080100 -2001020 rL(r) D Figure 3-7: Ripley’s K(r) analyses confirm a clumped adult distribution in all four plots. When L(r) (continuous line) is outside the confidence interval (dotted lines), it is possible to reject the complete sp atial randomness hypothesis (with a risk of =10%) in favor of regularity or clustering at distance r. Plot 2 (B) displays middle-range (30 m) and long-range a ggregation (60 m). Panels A and C represent occasionally inundated forest and B and D represent terra firme forest.

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43 Both forest type and microenvironment we re significant predictors of nearest neighbor distances (p = 9.1 x 10-6 and p = 0.0001, respectively), with those trees in terra firme being more distant fr om one another (15.3 + 1.1 m) than those in the seasonally inundated forests (11.5 + 0.4 m). Canopy class, production and all possible interaction terms were not significant for nearest neighbor comparisons. Discussion Density and Distribution I found that the density of one of the Amazon’s most important timber trees far exceeds that of most other species. Carapa guianensis had densities of 25.5 and 14.5 trees ha-1 in occasionally inundated and terra firme forests, respectively. In contrast, Gullison et al. (1996) report densities ra nging from 0.31 to 1.6 trees ha-1 for Swietenia macrophylla (dbh > 2.5 cm). In the Western Amazon, Wadt et al. (2005) found densities of only 1.35 trees 10 cm dbh ha-1 for Brazil nut (Bertholletia excelsa), another economically important tropical tree. Furtherm ore, I found extremely high densities in the occasionally inundated forest compared to the terra firme forest. McHargue and Hartshorn (1983) also reported that C. guianensis was predominantly a species of swampy or periodically inundate d land, but is also found in lower densities on higher and better-drained slopes and ridges. The natural distribution and abundance of a species is partly a function of the spatial variability in available habitats and the species capacity to colonize these habitats (Hall & Bawa 1993). Size-class Structure The frequency distribution (Figure 3-3) s hows that populations in both forest types show similar diameter distribu tion patterns. Peters (1996) desc ribes this j-distribution as a “type-I size class distribution”, one that displays a greater nu mber of smaller size-class

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44 trees than larger size-class tr ees, and an almost constant proportional reduction from one size class to the next. This t ype of structure is character istic of shade-tolerant canopy trees that maintain a more or less consta nt rate of recruitmen t (Peters 1996). While C. guianensis is a well characterized shade tolerant (Clark & Clark 1985), it is a masting species, and its ability to maintain constant recruitment over time in this population may speak to the ecological importance of this seed production strategy. The distribution suggests a demographically hea lthy population; many authors c onsider a type I structure the ideal of a stable, self-maintaini ng plant population (Meyer 1952, Leak 1965). Not only were diameter distri butions similar across forest types, but the percentage of reproducing individuals in any given dbh class was also not significantly different between occasionally inundated and terra firme forest. This suggest s that regardless of forest type, individuals in a given diameter class have approximately the same probability of producing seeds. Further res earch is still necessary to de termine whether the quantity of seeds produced differs between forest type s. Similarly, further research may indicate a difference in the time it takes to reach a cer tain dbh between forest types. I found that seed production was initiated in the majority of trees > 20 cm dbh in all plots, yet do not know how long it took for them to attain this diameter. Viera et al. (2005) documented growth rates of five C. guianensis trees growing in open forest with bamboo and dense terra firme forest. They found that individuals of C. guianensis with dbh measurements of 17.0, 37.5, 55.0, 56.0 and 84.0 cm, had ages of 785, 172, 180 + 120, 187 + 145 and 277 + 75, respectively. These data do not incl ude individuals in occasionally inundated forests, but they clearly demonstrate high variab ility in growth rates. This variability in growth rates may prove problematic fo r sustainable harvest projections of C. guianensis

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45 and other long-lived, slow-growing species. Si nce vital demographic rates are normally stage-dependent (Harper 1977), matrix mode ls that explicitly consider the size (Leftkovitch 1965), dependent fecundity, growth and survival rates of individuals have been used to explore the eff ect of different harvesting regimes on the growth rate and structure of populations (Pin ard 1993, Olmsted & Alvarez Buylla 1996, Zuidema & Boot 2002). High variability in growth rates confound projection matrices. Forest type may influence time to se ed production, but other variables may outweigh forest type effects. For example, light, which ha s been well correlated with relative height and spatial di stribution (Nigh & Love 2004), has been shown to play an important role in stimula ting seed production (Greene et al. 2002). In my study, the occasionally inundated plots consistently had higher numbers and densities across almost all size classes, suggesting a more robust population. In terra firme plots, however, a greater percentage of adults > 10 cm dbh were in the dominant or co-dominant canopy positions, and it was in the terra firme plots where the proportion of reproductive to nonreproductive adults was greater. Perhaps in the terra firme, a greater proportion of adults reach the upper canopy where they have grea ter access to light, creating conditions in which a greater proportion of these adults produce fruits. Forest type may have other effects du e to microclimate variation between the higher terra firme forests and the more swampy occasionally inundated forest (Svenning 1999). Boll et al. (2005) found that soil moisture was the most important environmental predictor of occurrence for the tropical palm Aphandra. Valencia et al. (2004) also found that soil affected abundance differences for a variety of tropical trees. John and Sukumar (2004) found that species richness was w eakly influenced by topography. While not

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46 evident in this data set, intense fires, dur ing the 2005 dry season, in creased mortality in one of the terra firme forest plots (Wadt, personal co mmunication). Recent research has indicated that tropical forest s are becoming more susceptibl e to fire (Nepstad 1999). The effect of forest type may become importa nt in conjunction with these environmental changes. Spatial Distribution While adult and seedlings densities diffe red between forest types, both spatial analyses demonstrated aggreg ation of individuals. Clumped distributions of trees are typical in both tropical dry (Hubbell 1979) and humid fore sts (Clark and Clark 1984). Aggregation seems to be highly linked w ith seed dissemination processes (Goreaud et al. 1999) and aggregation is ecologically benefi cial; aggregation of conspecifics should increase pollination efficiency, outcrossi ng success, and species abundance (Hubbell 1979). Adults occur in relatively large ne ighborhoods, probably due to competitive exclusion as the plants get larger with age. Seedlings, saplings and smaller diameter individuals occur in tighter cl usters, since they are recruite d by mature plants (Gibson 2002). Some differences in levels of aggreg ation were detected. Plot 4 (Figure 3-6), representative of a terra firme forest, was partially dominated by a bamboo. Very few individuals of C. guianensis were found in this area, whic h led to lower densities and higher levels of aggregation for trees in this plot. Another terra firme forest plot (plot 2) showed both middle range and long range aggregation (Figure 3-7). The middle range ag gregation may be due to seed dispersal close the parent tree and the long-range disp ersal could be due to water or animal transport of seeds. Water is a crucial transport agent for C. guianensis as seeds fall during

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47 the rainy season in some areas (Pennington 1981, Raposo 2002). Agoutis (Dasyprocta leporina) and collared peccaries (Tayassu tajacu) also disperse seeds while foraging (McHargue & Hartshorn 1983). Both Clark & Evans and Ripley’s K(r) demonstrated aggregation of individuals. As expected, Ripley’s K(r) was more appr opriate for showing whether aggregation occurred at different distances. Both me thods, however, indicat ed that the local population of C. guianensis was aggregated irrespective of forest type. Aggregation is common in a variety of other tree species (Boll et al. 2005, Svenning & Skov 2005) and has been most often related to dispersal limitation (Condit et al. 2000, Svenning 2001, Svenning & Skov 2005). Management Implications These results highlight several demographi c variables that merit special focus for species management. First and foremost, C. guianensis densities are extremely high compared to almost any tropical species. Low densities of economically important conspecific adult trees in many tropical fore sts are a major constraint to sustainable resource exploitation and a chronic management problem (Peters 1996). My study and others (McHargue & Hartshorn 1983, Plowden 2004), however, suggest that in a variety of environments, C. guianensis is found at relatively high densities, favoring management and potentially suggesting comparatively highe r yields (seed or timber) per hectare than other species. Furthermore, the extremely high densities in the occasionally inundated forest suggest that this forest type in part icular may merit special attention for future management; there were simply more trees present. These high densities which favor manage ment, however, may be counterbalanced with excessively low growth rates. Growth differences between forest types might be the

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48 most important demographic variable for determining timber harvest cycles. The extremely slow C. guianensis growth rates reported by Viera et al. (2005) from terra firme forests serve as cautionary signals if hoping to manage for sustainable timber production. For non-timber resources, differences in key demographic variables, such as age to first reproduction, could play a major role in resource management. My study results do not provide direct data on age to first reproduction, but I did find that proportionally more adults in terra firme forests are reproductivel y mature, perhaps tied to light access. While it is unclear whether these terra firme adu lts reach reproductive maturity more quickly than those in occasionally inundated forests, this merits further study for management purposes. Highly dispersed timber and non-timber re sources present a ma jor challenge to managing tropical forests because it is not econo mically feasible to access these disparate resources. I found that C. guianensis in both forest type s presented a clumped distribution, which could facilita te a concentration of manageme nt activities such as trail construction and seed collection. In conclusion, this study found spa tially similar distributions of Carapa guianensis in both terra firme and occasionally inunda ted forests studied. Tree densities and reproductive potential, however, were sufficien tly different to recommend stratification by forest type for further studies on growth and yield.

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49 CHAPTER 4 CONCLUSION While there is still considerable scien tific debate over whether and how natural resource extraction can be sustained over th e long term from tropical tree populations (Nepstad 1999, Peres et al. 2003, Pearce et al. 2003, Fearnside 2005), the debate is hindered by the lack of demographic inform ation for most tropical species (Gullison et al. 1996, Zuidema 2003), as well as the paucity of published ecologica l data on tropical species. The objective of this thesis was to provide a synthetic review of the most relevant ecological and management literature of Carapa guianensis to identify gaps in current research and recommend avenues for further st udy. In the second chapter, I compared the demographic structure and seedling recruitment of C. guianensis in two forest types, occasionally inundated and terra firme, or upland forests. Main study objectives were: (a) to assess the density, distribut ion, and size class structure of C. guianensis in occasionally inundated and terra firme forests; (b) to compare spatia l distribution analyses using two different methodologies; and (c) to use pa tterns of abundance, distribution and demographic structure to help infer key dem ographic stages or ecological variables that should be the subject of special focus when implementing a management scheme. The synthetic review revealed that al though authors have touted the economic potential of C. guianensis in the literature, very little re search has been done to quantify this potential or to look at sustainable harvest limits/challe nges. Gaps in information on C. guianensis pollinators and species genetics were identified. There is still a need for

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50 silvicultural experiments to ascertain methods for improving growth rates, tree form and increasing seed production per tree. Methods of controlling H. ferrealis and H. grandella infestations in seedlings and saplings, respectively, and ways to promote/improve survival and growth of young C. guianensis individuals may also bear investigation. Seed collection will likely affect seed pr edators. More research determining the broader ecosystem impacts of seed and timbe r collection may be beneficial. Similarly, broader faunal resources such as game meat are critically important for human inhabitants (Hill et al. 2003). The linkages between seed populations and seed predators is important to quantify, both for ecologists and forest residents who rely on these animals for sustenance. Economic and social analyses of comme rcialization barriers for small scale landowners is a key step towa rd utilization and management of this resource. From the forest to the final product, ther e is still a wealth of information yet to be discovered to better advise ecosystem managers and commercial producers of this species. Research on the population structure provi ded encouraging resu lts for a species with such great economic potential. In contra st with many tropical tr ee species, the local population of C. guianensis at the Embrapa forest has a high density of trees per hectare: 20.1. The population survey also confirmed the importance of microhabitat, with seasonally inundated plots having a ~55% highe r density than upland forest plots (25.7 versus 14.6 trees hectare-1). Seedling recruitment in th e study plots slightly exceeded mortality in both habitats, which in dicates a currently healthy population. Based on application of Donnelly’s (1978) nearest nei ghbor method, the index of aggregation (R) for all individuals was 0.9056 in the seasona lly inundated plots and 0.8258 for the upland

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51 dry forest plots, significantly different than a random distribution. All individuals showed a significant tendency toward clumping a nd this was even more pronounced for nonreproductive juveniles. Low density of conspeci fic adult trees is a major constraint to sustainable resource exploitation, but this study and others (McHargue & Hartshorn 1983, Plowden 2004) reveal a high density and clumped distribution, two factors that favor management and enc ourage further research. Measuring population structure in differen t habitats can be a first step towards assessing sustainability of ha rvesting in different habitat types (Peters 1996). Quantifying species demographic structure provides the unde rlying data for use in ecological models for assessing long-term populati on viability (Alvarez-Buylla et al. 1996). Since vital demographic rates are normally stage-depe ndent (Harper 1977), the structure of a population can be indicative of its demographic future and can be the basis for immediate management decisions when long-term demogr aphic monitoring is not feasible (Bruna & Kress 2002). This demographic data coupled w ith tree growth rates, survival and seed production data could form the basis of an ecological model to measure sustainable harvest rates of both the timber and non-timber resources that C. guianensis yields. Unless information on harvest limits and suggest ed management techniques is available for C. guianensis, resource extraction will likely be based on maximizing short-term economic revenue and not on managing a st able or increasing resource base.

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52 APPENDIX A PLOT INSTALLATION To open the 400 m transects that defined the plot boundaries, we determined the initial compass direction (normally 90 or 0 de grees). A support for the level (a branch) was cut and the compass was placed on the suppo rt. Two branches/poles where placed in the line of sight of the compass to begin a st raight line. Field technicians and ‘mateiros’ then used the branches as a reference to cut a straight line through the forest following the determined compass direction. Reference bran ches were placed no more than every 10 m apart (more frequently in hilly terrain) so that the compass direction was apparent. A compass was used to check the placement of the reference branches throughout. A metric tape was used to measure the distance cl eared until 400 m was reached. After clearing each transect, the metric tape was used to go back and re-measure each transect. This method was used for all transects bordering the plot as well as th e internal transects cleared to facilitate tree identification and location.

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53 APPENDIX B RANDOM SELECTION OF RE GENERATION SUBPLOTS These subplots were selected by dividing each plot into 16 quadrats with onehundred 10 x 10 m subplots. Excel’s random number generator was used to select random numbers from 1 to 16 and then agai n to select random numbers from 1 to100. These random numbers were paired to select the 32 subplots. The selected sub-plots were mapped on graph paper and located in the field.

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66 BIOGRAPHICAL SKETCH Christie Ann Klimas was born on July 23, 1978, in Cleveland, Ohio. She received a B.S. in environmental science with a bi ology concentration at Southampton College, Long Island University. During ni ne months of her undergradu ate studies, she worked as an intern for Brookhaven National Lab at their Free-Air Carbon Dioxide Exchange (FACE) site in North Carolina. The FACE system facilitates CO2 enhancement of entire forest stands. Following graduation, she spent 1.5 years as a resear ch specialist at Columbia University’s Biosphe re 2 Center in Arizona wh ere she again studied plant carbon uptake under CO2 enhancement. In 2001, she was awarded a Rotary Ambassadorial Fellowship to conduct research in Acre, Brazil. This preliminary research prepared her for master’s level study at the Un iversity of Florida in 2003. This thesis was supported by a Tropical Conservation a nd Development Assistantship and an Environmental Protection Agency STAR (S cience to Achieve Results) Fellowship. In 2006, she was awarded a National Science F oundation IGERT (Inte grative Graduate Education and Research Traineeship) fellows hip to continue doctoral studies at the University of Florida.