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Natural Regeneration of Canopy Trees in a Tropical Dry Forest in Bolivia

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1 NATURAL REGENERATION OF CANOPY TREES IN A TROPICAL DRY FOREST IN BOLIVIA By BONIFACIO MOSTACEDO A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL OF THE UNIVERSITY OF FLOR IDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY UNIVERSITY OF FLORIDA 2007

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2 2007 Bonifacio Mostacedo

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3 To my parents for raising me with love and maki ng me what I am today. To my wife, who brings great joy into my life. To my children, for the inspiration and st rength they gave me throughout the long process of comple ting this dissertation.

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4 ACKNOWLEDGMENTS So many people contributed to this work that I cannot acknowledge them all by name but I do want to single out a few. First of all, I would like to thank my mentor and principal supervisor Francis E. “Jack” Putz for asking me difficult questions and pushing me to think and write logically. He helped me adjust to life in Ga inesville and was a good fr iend throughout my career as a graduate student. Particular ly while I was trying to finish my dissertation, he provided a great deal of support and encouragement. I hope th at we can continue to work together in the future. I also want to thank the members of my advisory committee, Kaoru Kitajima, Karen Kainer, Emilio Bruna, and Colin Chapman, for always being available to pr ovide me guidance. For much of the past decade, Todd Fredericks en has been a key person in my life. We spent a great deal of time together in various fo rests in Bolivia conducting what I believe to be exciting research and teaching wh at I hope were useful field c ourses. No matter how bad the conditions, he always had a smile on his f ace and an amusing comment on his lips. He encouraged me to start my Ph.D ., wrote letters of recommendation on my behalf, and helped me secure financial support for my st udies. He continues to provide critical feedback on my work but, overall, I am most grateful to him for being a good friend. Numerous University of Florida faculty a nd staff contributed substantially to my dissertation project. In partic ular, Doug Levey helped me design the predator exclosures and Stephen Mulkey helped me with the irrigation experiment. Am ong the statisticians who guided me through the design and analysis phases of this research, I would like to thank Larry Winner, Andrew Khuri, and Jorge Cassela. I am also very grateful to Ann Wagner, Pamela Williams, and Paula Maurer from the Botany office for their help.

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5 My Ph.D. studies were supported to a larg e extent by a fellowship from USAID through the BOLFOR Project run by Chemonics Intern ational. Thanks go to John Nitler and Ivo Kraljevic for helping me to secure this fundi ng. They, and several other people at Chemonics, helped me to deal with culture shock and to othe rwise adjust to life in the USA. Part of the research was funded by International Foundation fo r Science (IFS) who gave financial support to do experiments about seedling dynamic. The fina l phase of my dissertation research was supported by a fellowship from the Gordon a nd Betty Moore Foundati on through the Tropical Conservation and Development Program at the University of Florida. I want to thank my officemates and friends in Gainesville including Clea Paz, Geoffrey Blate, Joseph Veldman, Morgan Varner, Bil Gr auel, Skya Murphy, Ana Eleuterio, Paulo Brando, Camila Pisano, and Christine Lucas. Skya, An a, Paulo, and Christine provided me helpful comments on my dissertation. I also want to re cognize Claudia Romero for teaching me various things that made my life easier, helping me w ith the dreaded dissertatio n templates, reviewing the sprouting chapter, and convinc ing me that I could finish this dissertation on a time schedule that seemed unattainable. The Instituto Boliviano de Investigacion Fore stal (IBIF), through its director Marielos Pea-Claros, provided me with all the necessa ry logistical support needed to conduct my fieldwork. Marielos also provi ded many helpful comments on my dissertation proposal. Many other people at IBIF were strong supporte rs of my work, including members of the administrative (Emma Nuez, Laly Dominguez, and Karina Munoz) a nd technical staffs (Marisol Toledo, Zulma Villegas, Juan Carlos Li cona, Alfredo Alarcon, Ca rlos Pinto, Claudio Leao, Vincent Vroomans, Betty Flores and Mayr a Maldonado) as well as Lourens Poorter, a research associate. From Lourens I received many helpful suggestions about my research.

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6 My fieldwork would not have been possible wi thout help of my student assistants and “materos.” In particular, Marlene Soriano helped me collect a portion of the field data for the first and second chapters of th is dissertation. Armando Villca and Turian Palacios helped me with the field research for the third chapte r. Alejandra Calderon, Va nessa Sandoval, Monique Grol, Janeth Mendieta, Carla G onzalez, and Joaquin Cordero helped in various ways in and on the way to the field. I want also to thank ma ny other people for fieldw ork assistance including Juan Carlos Alvarez, Israel Me lgar, Daniel Flores, Hugo Justin iano, Rodolfo and Rafael Rivero, Daniel Alvarez, Alberto Chacon, Antonio Jimenez, Juan Pessoa, Dona “Negrita” Pessoa, and Juan Alvarez. I also want to thank Miguel Ange l Chavez, the driver from IBIF, who was always willing to volunteer to travel the long bumpy road to INPA. The INPA PARKET Company was the gracious host of my field research and helped in many different ways in its execution. The company’s owner, Paul Roosenboom, company manager William Pariona, forester Urbano Choque and many other people working for this prestigious and FSC-certified fore st-products firm contributed to the success of this project. Last but certainly not least, I want to thank my wife, Ynes Uslar, and children, Gabriela Ines and Jose Daniel, for the many sacrifices a family makes when a parent takes on a Ph.D. project. The many nights I spent writing instead of being with them, the long stints of field work, and the even longer periods when I was in Gainesv ille and they were back in Bolivia were hard on all of us but they bore the strain without complaint. I shou ld also thank Ynes for counting thousands of seeds and typing in even more data, but these contributions pa le in comparison with the rest of what she has done for me and the rest of our family.

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7 TABLE OF CONTENTS page ACKNOWLEDGMENTS...............................................................................................................4 LIST OF TABLES................................................................................................................. ..........9 LIST OF FIGURES................................................................................................................ .......11 ABSTRACT....................................................................................................................... ............13 CHAPTER 1 FRUIT PRODUCTION OF TROPICAL DRY FOREST TREES IN BOLIVIA..................15 Introduction................................................................................................................... ..........15 Methods........................................................................................................................ ..........16 Study Area and Climate...................................................................................................16 Species Studied................................................................................................................ 17 Experimental Design and Data Collection......................................................................18 Data Analysis.................................................................................................................. .20 Results........................................................................................................................ .............20 Discussion..................................................................................................................... ..........22 Fruiting of Trees in a Tropical Dry Forest......................................................................22 Factors Affecting Fruit Production..................................................................................24 Conclusions.................................................................................................................... .........26 2 BIOTIC AND ABIOTIC FACTORS A FFECTING TREE SEEDLING DYNAMICS IN A DRY TROPICAL FOREST................................................................................................39 Introduction................................................................................................................... ..........39 Methods........................................................................................................................ ..........42 Study Area and Climate...................................................................................................42 Experimental Design and Data Analysis.........................................................................44 Response of regeneration to silv icultural treatment intensity..................................44 Factors affecting seedling establishment and growth...............................................45 Results........................................................................................................................ .............48 Response of Regeneration to Management Intensities....................................................48 Factors Affecting Seedling Gr owth and Establishment..................................................49 Seedling growth........................................................................................................49 Seedling establisment...............................................................................................50 Discussion..................................................................................................................... ..........51 Logging Effects on Seedlings Dynamics........................................................................51 Factors Affecting Seedling Establishment and Growth..................................................55 Conclusions.................................................................................................................... .........57

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8 3 CONTRIBUTION OF ROOT AND STUMP SPROUTS TO NATURAL REGENERATION IN A LOGGED TROPIC AL DRY FOREST IN BOLIVIA..................79 Introduction................................................................................................................... ..........79 Methods........................................................................................................................ ..........80 Study Area..................................................................................................................... ..80 Experimental Design and Data Collection......................................................................81 Stump sprouts...........................................................................................................81 Comparison of different juvenile types in relation to microsites created by logging..................................................................................................................82 Data Analysis.................................................................................................................. .83 Results........................................................................................................................ .............83 Sprout Characterization...................................................................................................83 Juvenile Types and the Effects of Logging.....................................................................86 Growth of Stump Sprouts................................................................................................86 Discussion..................................................................................................................... ..........87 Natural Regeneration and Shade Tolera nce: True Seedlings vs. Sprouts.......................88 Allometric Relationships with Stump Sprouting.............................................................89 Growth of Stem and Root Sprout s Compared with True Seedlings................................90 Conclusions.................................................................................................................... .........91 LIST OF REFERENCES............................................................................................................. 107 BIOGRAPHICAL SKETCH.......................................................................................................117

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9 LIST OF TABLES Table page 1-1. Overview of reproductive characteristics of th e tree species studied in a tropical dry forest in Bolivia.............................................................................................................. ....28 1-2. Results of the best model after back ward regression steps to remove non-significant variables (P > 0.1) sequentially from th e full multiple regression model including tree size (DBH), percent liana infesta tion, crown area, and crown position as independent variables to explain fruit production.............................................................29 1-3. Analysis of covariance to determine th e liana cutting effect in fruit production of Caesalpinia pluviosa ..........................................................................................................30 2-1. Spatial distributions, crown position, ecological group, geogr aphical range, tree densities and basal area of timber species in a Bolivian tropical dry forest......................59 2-2. Means of seedling densitie s in an unharvested control pl ot, a plot subjected to normal timber harvesting, and a plot subjected to more intensive harvesting followed by silvicultural treatments....................................................................................................... 60 2-3. Mean seedling densities (#/m2) for 3 years (2003-2005) of commercial tree species in microsites created by logging in a Chiquitano dry forest..................................................61 2-4. Establishment and mortality rates of seedlings of commercial tree species monitored over a 3 y period in a control plot and pl ots subjected to two harvesting intensities (N=144 subplots/treatment plot)........................................................................................62 2-5. Results of repeated measure analysis of variance for a spl it plot design run for seedling density for all species combined or six timber tree species analyzed separately..................................................................................................................... ......63 2-6. Mean relative height grow th rates ( 1SE) of seedlings of commercial tree species in response to bromeliad cover re moval evaluated in 4 times...............................................66 2-7. Mean relative height grow th rates ( 1SE) of seedlings of commercial tree species in irrigated and droughted plots evaluated 4 times................................................................67 2-8. Mean relative height grow th rates ( 1SE) of seedlings of commercial tree species in response to mammal exclosure ev aluated 4 times during 2006.........................................68 3-1. Frequency of root and stem sprou ting and shade tolerance of commercial and noncommercial canopy tree species in a tr opical dry forest in Bolivia...................................92 Table 3-2. Summary of the best models and th eir significances from the regression analyses between stump diameter or stump volume (i ndependent variables) and the heights of stump and numbers of sprouts per stump (dependent variables).......................................93

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10 3-3. Mean ( 1SE) densities of true seedli ngs, stem sprouts, and root sprouts in 10 x 4 m plots in microsites created during selective logging. ........................................................94 3-4. Mean ( 1SE) of stump sprout height growth rates (cm/yea r) by ecological groups........95 3-5. Means ( 1SE) of stem heights by sp ecies that sprouted from stems or roots compared to the heights of seedlings.................................................................................96 3-6. Mean ( 1SE) heights of trees 2 m tall that were root sprouts, stem sprouts, or seedlings grouped by ecological guild...............................................................................97

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11 LIST OF FIGURES Figure page 1-1. Locations of the study areas in Bolivia..............................................................................31 1-2. Percentage of trees fruiting in 2003 and 2004 (note that none of these species are dioecious). ................................................................................................................... ......32 1-3. Annual variation in fruit pr oduction in three timber species. ...........................................33 1-4. Comparison of the sizes (DBH) of fr uiting and non-fruiting trees in 2003 and 2004 (note that none of these species are dio ecious and all the trees were reproductively mature)........................................................................................................................ .......34 1-5. Simple linear regressions between crow n area, tree size (DBH ) in relation to log transformed fruit production data for three timber species................................................35 1-6. Fruit production of Caesalpinia pluviosa in relation to DBH and crown area for trees with cut or uncut lianas......................................................................................................3 6 1-7. Percentages of trees fruiting in a logged and an unlogged plot.........................................37 1-8. Average fruiting intensities (% of crow n cover) of timber tree species evaluated in areas subjected to normal selective loggi ng and nearby unlogged control areas in a tropical dry forest............................................................................................................ ...38 2-1. Monthly rainfall (A) and soil moistu re tension measured by Watermark soil sensors (B).................................................................................................................... .....69 2-2. Canopy openness in an unlogged control pl ot, an area subjected to normal timber harvesting (4.7 m3/ha harvested), and an area subject ed to intensiv e harvesting (8.2 m3/ha) 8 months (left-hand bar) and 42 mont hs (right hand bars) after logging...............70 2-3. Design of the experiment on the eff ects of ground bromeliads, irrigation, extended drought, and seed and seedling predator exclosures on seedling establishment................71 2-4. Seedling densities of 11 timbe r species (A) and 10 species without Acosmium cardenasii (B), the most dominant species, in a control plot, a plot subjected to normal timber harvesting, and a plot subjected to intensive timber stand management..................................................................................................................... ..72 2-5. Temporal changes in seedling recruitment (#/m2/y) and mortality for commercial tree species in a Chiquitano dry forest in Bolivia.....................................................................73 2-6. Seedling density over time in response to (A) bromeliad cover, (B) irrigation or drought, and (C) mammalian seed predators.....................................................................74

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12 2-7. Mean densities ( 1SE) of seedlings of commercial timber tree species in 2 m2 plots (N=40) with bromeliads (filled dots) a nd without bromeliads (open dots). .....................75 2-8. Mean of seedling densities ( 1SE) in irrigated and droughted experimental plots..........76 2-9. Mean seedling densities (1 SE) in c ontrol plots (closed dots) and in plots from which mammals were excluded (open dots)......................................................................77 2-10. Mean relative growth height growth ra tes ( 1SE) of seedlings of commercial tree species in response to three experimental treatments: (A) irriga tion or drought; (B) mammalian seed predator exclusi on; and, (C) bromeliad cover........................................78 3-1. Proportions of stumps of commercial timber species that resprouted (number of stumps noted in parenthesis)..............................................................................................98 3-2. The proportions of stumps with live sprouts over time since logging...............................99 3-3. Mean ( 1SE) numbers of sprouts/stum p for the most frequent sprouting species.........100 3-4. Probabilities of stump sprouting as a function of stump diameter for the most frequently sprouting commercial tree speci es (curves fit by logistic regression)............101 3-5. Probabilities of stump sprouting as a function of stump height for the commercial tree species that most fre quently sprouted (curves f it by logistic regression).................102 3-6. Mean ( 1SE) densities of juveniles < 2 m tall of commercial tree species that were true seedlings, stem sprouts, and root sprouts..................................................................103 3-7. Percentage of juveniles < 2 m of diff erent origins after logg ing (for each of 15 dry forest tree species) ordered by their light requirements...................................................104 3-8. Relative growth rates (mean ( 1SE) of stump sprouts measured over the first two years after logging for commercial tree specie s in a tropical dry forest in Bolivia arranged by light requirements........................................................................................105 3-9. Mean growth rates of stump sprouts through time for the main commercial tree species as based on measurements of the ta llest sprouts on different stumps 1, 2, and 5 y after creation............................................................................................................. .106

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13 Abstract of Dissertation Pres ented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy NATURAL REGENERATION OF CANOPY TREES IN A TROPICAL DRY FOREST IN BOLIVIA By Bonifacio Mostacedo May 2007 Chair: Francis E. Putz Major Department: Botany Fruit production, seedling estab lishment, and sprouting of ca nopy trees were studied in a lowland tropical dry forest in the Departme nt of Santa Cruz, Bolivia. Fruit production by reproductively mature trees was monitored over a 5 y period to assess variability among species, trees and years. The effects of tree size, crown area, crown position, and lia na infestation on fruit production were also assessed. In a companion study, I assessed the effects of lianas on fruit production by Caesalpinia pluviosa with a liana cutting experime nt. To determine how logging disturbances affect seedling recruitment, I monito red seedlings for 3 y in different microsites in permanent plots in two selectively logged pl ots and an unlogged c ontrol plot. I also experimentally assessed the effects of brome liad cover, drought stress, and seed/seedling predators on seedling recruitment, survival, and growth. Finally, I monitored the emergence, survival, and growth rates of stump and r oot sprouts over a range of microsites. Percentages of trees fruiting and numbers of fruits produced vari ed among species and years. In most species, trees that did and did not fruit did not differ in si ze or crown position, but in a few cases, the likelihood of fruiting increased with crown area. Contrary to my expectation, there no effect of liana cutting on Caesalpinia pluviosa fruit production was de tected 3 yr after

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14 cutting. Overall, the effect of logging on the pr oportion of trees fruiti ng and fruiting intensity varied among species. Seedling densities 5 y after se lective logging were higher in control than logged plots but this finding was greatly influen ced by the most common species, Acosmium cardenasii (43% of seedlings enumerated). At the microsite level, Acosmium was found in highest densities in undisturbed areas while Centrolobium microchaete, another common species, was more common on log extraction paths. Seedling recruitment rate s were higher in the unlogged plot and in the undisturbed portions of the logged fo rest plots, but seedling mortal ity rates were also higher in these areas. Mortality rates of naturally establ ished seedlings varied greatly among species. Seven of 22 species suffered no mortality during the 2-y monitori ng period, whereas relatively high mortality rates were observed for Caesalpinia (26%/y) Sweetia fruiticosa (25%/y), and Machaerium scleroxylon (22%/y). Results of the experimental study on seedli ngs suggest that bromeliad competition and seed/seedling predators greatly affected tree seed ling establishment. Soil moisture availability also affected seedling establishment, but only as an interaction with the bromeliad removal or predator exclosure treatments. The primary effect of the drought treatment was delayed germination. Despite these general trends, species varied substantially in their sensitivity to bromeliads, drought stress, and predators. Root and stump sprouts constituted about 50% of the individuals <2 m tall of the canopy tree species studied, but the pr oportions of sprouts and true seedlings varied among species. Stump sprouting was common, but the probability of sprouting was not cons istently related to stump diameter or height. Sprout growth rates were consistently high, at least initially, and sprouting is obviously important to post-logging regeneration in this dry tropical forest.

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15 CHAPTER 1 FRUIT PRODUCTION OF TROPICAL DRY FOREST TREES IN BOLIVIA Introduction Tropical dry forests, which until very recen tly covered about 30% of Bolivia, are among the most threatened ecosystems in the world (Dinerstein et al. 1995). Bolivian dry forest are under siege; 32% has already been cleared (Camacho et al. 2001, Rojas et al. 2003) and most of the remainder is under intensive pressure for forest products, grazing, and further conversion. This pressure may be somewhat mitigated in the large portion of the dry forest managed for timber under the guidelines of Bolivia’s 1996 Forestry Law (MDSP 1996, Nittler and Nash 1999). Even when these guidelines are followed, logging disturbances are larger in area than those that occur naturally. Given the documented effects of disturbance a nd habitat modification on reproductive success of tropical dry forest trees (Fuchs et al. 2003), the sustainability of managed forests even when the legally required “best management” practices are used, remains uncertain. To assess how forest management influences the reproduction of commercially valuable tree species, I explore the intertree and inter-annual variation in fruit production in a dry forest in lowland Bolivia. A wide range of mechanistic explanations have been proposed for the marked variation in fruit production among years and am ong individuals in a wide vari ety of forested ecosystems (Abrahamson and Layne 2003, Snook et al. 2005). Fo r seasonally dry tropical forests, interannual variation in rainfall, part icularly as influenced by El Ni o events, has often been invoked as the underlying cause of interannual variation in tree phenol ogy and fruit production (Bullock 1995, Wright and Calderon 2006). Rega rding individual tree characteri stics that might influence reproductive output, several studies have repo rted a minimum size threshold for tree reproduction (Aez 2005, Wright et al. 2005). For trees that have attained reproductive maturity, several studies have shown a re lationship between fruit producti on and stem diameter (Zuidema and Boot 2002, Wadt et al. 2005), crown area (Healy et al. 1999, Zuidema 2003), and crown

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16 position (Healy et al. 1999). Cove r by lianas has also been show n to reduce fruit production by Brazil nuts “ Bertholletia excelsa Bonpl.” (Zuidema and Boot 2002, Wadt et al. 2005, Kainer et al. 2006) and other species (Stevens 1987). Anthropogenic and natural disturbances have been reported to influence canopy tree fruit production in a number of forests around the tropic s but the results have been inconsistent. Logging increased fruit production by remnant trees in a subtropical humid forest in Meghalaya, India (Barik et al. 1996), as well as in a tropical montane forest in Costa Rica (Guariguata and Saenz 2002). In contrast, reproductive output of resi dual trees in a logged di pterocarp forest in Indonesia was lower than in an unlogged control area (Curran et al. 1999). Factors responsible for these contrasting results ar e not clear but, given the importa nce of natural regeneration to sustainable forest management, the issue deserves further exploration. In this study in a seasonal lo wland tropical forest in Boliv ia I report on canopy tree fruit production over a 5-year period. I also examine the relationships between fruit production and crown features, tree size, and liana infestation. Using one comm on tree species that is often heavily liana-laden, I assess the effect of liana cutting on fruit productio n. Finally, I examine the effect of selective logging on fruit producti on by several common canopy tree species. Methods Study Area and Climate Research was conducted at INPA Parket, a 30,00 0-ha tract of privately owned seasonally dry tropical forest located 30 km northeast of the town of Concepcin (16 6' 45" S and 61 42' 47" W) and 250 km northeast of the city of Sant a Cruz, Bolivia (Figure 1-1). The altitude is approximately 380 m, mean annual temperatur e is 24.3 C, mean annual precipitation is 1150 mm (range 798-1859 mm/y), and there is a dry s eason that lasts about five months (MayOctober) during which most trees ar e leafless. Extreme annual variat ion in rainfall is in part

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17 related to the occurrence of El Nio events, wh ich are typically wet in the study area. Many tree species in this forest flower at the end of ra iny season with another pe ak in flowering at the beginning of dry season. Fruiting of the majority of species occurs in mid-dry season (see Table 1-1). The forest canopy is 20-25 m tall and dominated by Tabebuia impetiginosa (Mart. Ex DC.) Standl., Anadenanthera macrocarpa (Benth.) Brenan, Astronium urundeuva (Allemo) Engl., and Centrolobium microchaete (Mart. ex Benth.) Lima ex G. P. Lewis (Pariona 2006). The most abundant species are Acosmium cardenasii H. S. Irwin & Arroyo ( 38 trees/ha 20 cm diameter at breast height, DBH) and Anadenanthera (8 trees/ha 20 cm DBH). Currently, 21 tree species, including these dominants, are harveste d for timber used mostly in the production of parquet flooring. Species Studied My studies focus on commercially valuable timber tree species, most of which produce seeds that are wind, gravity, or e xplosively dispersed (Table 1-1). Overall I consider 31 species (Table 1-1), of which 15 were used to determin e the percentage of fru iting trees, 12 to compare the sizes of reproductively mature trees that did or did not fruit during a 2-y observation period, 6 to compare fruiting intensity in logged and unlogged areas, and 3 to determine the relationships between fruit production and DBH, crown area, crown position, and liana infestation. In addition, one species ( Caesalpinia pluviosa DC.) was used to study the effect of liana cutting on fruit production. Most of the selected species have large or medium-sized fruits with smallto intermediatesized seeds. Of the 31 species considered in this study 25 have hermaphroditic flowers and 20 produce wind-dispersed seeds. Ten species have seeds as their disp ersal units (e.g., Amburana cearensis (Allemao) A.C. Sm.) while in the other 21 specie s, entire fruits are the dispersal units (e.g., Machaerium scleroxylon Tul.). Twenty-four of the 31 species monitored for five years produced

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18 at least some fruits annually, but the in tensity of fruiting varied substantially. Cecropia concolor Willd. produced fruits continuously whereas Schinopsis brasiliensis Engl. fruited supra-annually (sensu Newstrom et al. 1994). Peak fruiting for 23 of 31 species coincided with the dry season (May – September) with most seeds dispersed in the late dry season. Spondias mombin L. the only fleshy-fruited species in the study produced ripe fruit during the ra iny season. Most of the species have only 1 seed per fruit, but Ceiba samauma (Mart.) K. Schum. and Tabebuia have more than 90 seeds/fruit. Wind-disperse d seeds or fruits typically have one ( Pterogyne nitens Tul.) or both sides winged (e.g., Platimiscium ulei Harms), whereas Zeyheria tuberculosa (Vell.) Bureau seeds are entirely surrounded by a wing. Cottony fibers on the seeds of the Bombacaceae studied ( Ceiba and Chorisia speciosa A. St.-Hil.) aid in their dispersal by wind. Experimental Design and Data Collection The study was conducted at two different sites at INPA Parket. The first site was within the Long-Term Silvicultural Research Plots (LTSRP) in Block # 1, 56 km south of the IBIF field station (16 18’ 26.8”S, 61 41’ 13.5”W). Four la rge-scales (20 ha) longterm research plots (LTRSP) were established by the Instituto Bolivia no de Investigacion Forest al (IBIF). One of the plots I used is an unlogged control whereas the other was subjected to normal logging during which a mean of 4.3 trees/ha (4.7 m3/ha of commercial timber) were harvested using standard reduced-impact-logging techniques that include road planning, dire ctional felling, and retention of 20% of the harvestable trees as seed trees (M ostacedo et al. 2006). In e ach plot, I censused all individuals 20 cm DBH of the 15 most abundant tr ee species in two 20 x 400 m strips. The second site is a 1000 x 500 m plot th at I established 2 km northeast of the IBIF field station (16 14’ 58.5” S, 61 41’ 47.4” W). In th is plot, I censused individuals 20 cm DBH of Anadenanthera and Caesalpinia in twelve 20 x 500 m strips space d at 100 m intervals. At both

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19 study sites the censused trees were marked, tagge d, and mapped. To secure sufficient trees of Caesalpinia and Zeyheria, I marked additional trees around the LTRSP. In total, I evaluated 440 trees of 15 species, 116 in the control plot and 147 in the logged plot of the LTRSPs. An additional 144 trees were monitored at the second site that, a year after plot establishment, was selectively logged. In 2003, I evaluated only six species from MayNovember including Caesalpinia, Anadenanthera, Mac haerium scleroxylon, Ceiba, and Zeyheria In 2004, every marked tree in the LTSRP plots was phenologically evaluated five times from May to November. I estimated the pe rcentage of the crown bearing fruits, which I refer to as a measure of fruiting intensity; 100% fruiting intensity indicates that every terminal branch bears at least one fruit. I estimated the number of fruits prod uced by each tree of three species, Caesalpinia Anadenanthera, and Zeyheria, for 5 years (2002-2006). All of th ese species produce fruits with valves that are not removed by animals. By c ounting these undispersed fruit parts under fruiting trees, I avoided many of the difficulties associat ed with estimating seed production by species in which entire fruits are removed by animals. At the beginning of the study I installed five 2x2 m permanent plots on the ground below the crown of each tree in which I counted and removed all fruit valves. Subsequent censuses of fallen fruit valves in these plot s over the next 5 y were used as quantitative estimates of fruit production. I al so measured the DBH, crown area (based on two cardinal diameters), crown position (using the 5 categories of Dawkins (1958)), and liana infestation. Liana-infested Caesalpinia trees (N=32) were used in a manipulative experiment on the effects of lianas on seed production. I measured the DBH and percentage of each tree’s crown covered by lianas, paired the trees on the basis of liana infestation and DBH, and cut all the

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20 lianas on one tree of each pair, se lected at random. Fruit produc tion was measured for 3 y after treatment using the fruit-valve cen sus method described above. Data Analysis To determine whether trees of the 12 monitored species that fruited in 2003 and 2004 differed in DBH from those that did not, I conducted Student’s-t te sts. I tested for simple linear relationships between the number of fruits (log -transformed) produced pe r tree and each tree’s DBH, crown area, crown position, an d liana cover separately and then ran multiple regressions for three monitored species ( Anadenanthera Caesalpinia and Zeyheria ) using the backward method to avoid co-linearity. Only independent variables that explaine d significant amounts of variance (P=<0.05) were included in the models. I ran X2 tests to assess differences between logged and control plots in the percentage of trees fruiting. To compare maximum fruiting intensities in the logged and the control plot for the 6 monitored species, I ran Student’s t-tests after first arcsine transforming the data to achi eve normality (Zar 1981). Finally, I compared fruit production on liana-laden and liana-free trees usi ng analyses of covariance with crown area or DBH as covariates. For all analyses, I used SPSS Version 12.0 (Field 2000). Results The percentage of trees fruiti ng in any particular year vari ed among species (Figure 1-2). Of 14 species monitored during 2004, 7 had >50% of trees in fruit and 4 had >80% of trees in fruit. For a few species that I also monitored in 2003, the percentage of fruiting trees was lower in 2003 than in 2004. For example, while only 25% of Caesalpinia trees fruited in 2003, 100% fruited in 2004. The opposite pattern was observed in Zeyheria ; 40% fruited in 2003 and none in 2004. For the 3 species I monitored fo r 5 years, there was a great de al of inter-annual variation in fruit production (Figure 1-3). For example, a high proportion of Caesalpinia trees fruited at 2-

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21 year intervals whereas many Zeyheria trees fruited at 3-year intervals. In contrast, Anadenanthera fruited during each of the first 3 years of the study and not at all in the last 2. Whether or not a tree fruited was generally not related to its DB H (Figure 1-4). The exceptions were Caesalpinia in 2004 when the trees that failed to fruit were larger than the trees that fruited (t=2.18, P=0.03) and Zeyheria in which the fruiting trees were larger in 2004, the only year in which it fr uited (t=4.91, P<0.0001). For the 3 species in which I monitored fruit production, the number of fruits produced did not vary with DBH or crown position but increas ed linearly with crown area (Figure 1-5). The backward multiple regression of fruit production on tree characteristics revealed that crown area explained the most vari ation (Table 1-2). In Anadenanthera crown area explai ned 32% of the variance in fruit production while DBH, crown pos ition, and liana cover together explained only an additional 10%. In the case of Caesalpinia, crown area explained 23% of the variance in fruit production and the other three variab les only an additional 3%. In Zeyheria crown area and crown position together explained only 24% of the varian ce, to which DBH and liana cover added an additional 1%. There was no apparent effect of liana cutting on fruit production by Caesalpinia when the effects of either DBH or crown area are re moved by ANCOVA (Table 1-3, Figure 1-6). The percentage of trees fru iting in the logged and control plot was similar for 6 of 7 species. The only exception was Centrolobium in which only 27% of tr ees fruiting in the logged plot compared to 90% in the control plot ( X2=12.9, P<0.0001; Figure 1-7). The effect of logging on fruit production varied among the 6 tree species studied (Figure 18). Logging apparently stimulated increased fruiting intensity of Anadenanthera (t=3.40, P=<0.0001). In contrast, fruiting intensity was higher in the unlogged plot for both Centrolobium

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22 (t=4.75, P=0.0008) and Copaifera (t=3.11, P=0.007). There was no difference in fruiting intensity between the logged and unlogged plot for Aspidosperma (t=0.68, P=0.5) and Machaerium scleroxylon (t=0.05, P=0.95). Discussion Fruiting of Trees in a Tropical Dry Forest In any year, the proportion of reproductively matu re trees that fruite d generally varied a great deal among the canopy tree species I studied in lowland Bolivia. Only Anadenanthera trees produced fruit crops in both 2003 a nd 2004. In contrast, no trees of Machaerium scleroxylon and M. acutifolium fruited in 2003, but many did in 2004. Most of the trees of some species fruited in at least some years (e.g., Caesalpinia ) while in others the percentage of fruiting trees was always <40%. Perhaps coincidentally, the five species w ith the greatest proportio n of fruiting trees in 2004 were all legumes, which comprised 8 of the 14 species monitored. Few studies report annual variation in the proportions of fruiting tree s but in a similar forest in Lomerio, Bolivia, only 29% and 36% of reproductively mature trees of 17 commercial tree species fruited during two years of monitoring (Justini ano and Fredericksen 2000). At th e same site, there was a great deal of variation within species in the proporti ons of fruiting trees. For example, the proportion of Copaifera trees fruiting was similar to what I observed in INPA whereas none of the M. scleroxylon trees in Lomeria fruited dur ing the two years of monitori ng. In a tropical dry forest in Mexico studied by Bullo ck (1995), only 8-30% of Jacaratia mexicana A. DC. trees and 050% of Cochlospermun vitifolium (Willd.) Spreng. trees fruited in any one year. Similarly, in a study of Swietenia macrophylla King. on the Yucatan Peninsula, the proportion of fruiting trees varied a greatly among years (Snook et al. 2005). The proportion of Hymenaea courbaril L. trees fruiting in Costa Rica reportedly vari ed with water stress (Janzen 1978).

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23 A larger proportion of trees of most speci es fruited in 2004 than in 2003 or 2005. Such community-wide synchrony in fruiting has been obs erved in many forests over the world but has been particularly well studied in the dipterocar p forests of Southeast As ia where “masting” has long been known (Appanah 1993, Wich and Van Schaik 2000). Less pronounced is the interannual variation in fruit production on Barro Co lorado Island, where mast years are reportedly related to El Nio events (Wright and Calderon 2 006). In the forest of this study, 2003 was at the end of an El Nio event, which was not followed by a strong La Nia. Nevertheless, synchronous fruiting in 2004 could have been due to the timing of water stress, as suggested by Bullock (1995). The three species I followed for five years showed great inter-annua l variation in the numbers of fruits produced per tree. The two legumes, Anadenanthera and Caesalpinia followed the same interannual patterns; both had peak years in 2002 and 2004, but produced few fruits in 2003 and 2005. In contrast, the 35 Zeyheria trees I monitored produced no fruits in 2004 and 2005, but fruited in 2002, 2003, and 2006. Such inte rannual variation in fruit production is common in many species. For example, in a moist tropical forest in Panama, Quararibea asterolepis Pittier, Tetragastris panamensis (Engl.) Kuntze and Trichilia tuberculata (Triana & Planch.) C. DC. showed great inter-annual variation in fruit pr oduction (De Steven and Wright 2002, Snook et al. 2005). While I expected that the proportion of fruiti ng trees would increase with tree size, 12 of the 13 species I monitored in 2003 and 2004 did not di splay this pattern. In fact, in the case of Caesalpinia the fruiting trees were signi ficantly smaller than those that failed to fruit in 2004. Only Zeyheria displayed the expected pattern. The general failure to find a positive relationship between tree size and whether or not a tree reproduced runs count er to the results of two other

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24 studies on this topic, one conduc ted on Barro Colorado Island, Panama (Wright et al. 2005) and the other at my study site (Aez 2005). One expl anation for the difference between my study and others in the literature is that whereas they ty pically used either flow ering or fruiting as an indication of reproduction (Aez 2005, Wright et al. 2005), I used only fruiting. Nevertheless, while the probability of fruiti ng increased with tree size in Copaifera Sweetia and Machaerium it decreased in Anadenanthera and Caesalpinia Factors Affecting Fruit Production Crown area was the best predictor of fruit production in many of the species I studied whereas DBH and crown position were not. Crow n area was also a good predictor of fruit production in Betholletia excelsa trees in Amazonian Bolivia (Zuidema and Boot 2002), while DBH was more closely rela ted to fruit production by Swietenia in Mexico (Snook et al. 2005) and Pinus sylvestris L. in Sweden (Karlss on 2000). Although several st udies have shown strong positive correlations between tree size and fru it production (Karlsson 2000, Zuidema and Boot 2002, Wadt et al. 2005), the relati onships revealed in my study were positive but weak. This difference might be related to the difficulty of making accurate DBH estimates of the trees in my study site, many of which have irregular-shaped tr unks. It is also possible that crown position is not good predictor of fruit producti on in forests that ar e already open-canopied; certainly in my study site it was difficult to differentiate between dominant and co-dominant trees, and I doubt that they differ much in terms of light capture. I expected liana cover to substantially impe de fruit production (Stevens 1987), but my results did not support that expe ctation. For the 16 pairs of Caesalpinia trees from which I monitored fruit production for 3 y after cutting their lianas, fru it production was no higher than in the liana-laden control trees. I can offer a few explanations for th is counter-intuitive result. For one thing, 3 y was perhaps not enough time for the trees to respond to the liberation from their

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25 liana loads. Then there is the problem that fruit production is extremely variable among individuals and among years in this species a nd others in my study si te. Finally, I wonder whether the liana leaf phenology at my study site might have something to do with this result insofar as lianas are typically deciduous at the time of fruiting, which might reduce any deleterious effect they have on fruit production. O bviously, none of these explanations is very compelling and further monitoring is warranted. Although I anticipated that by increasing canopy openness and reducing resource competition, logging would increase fruit pr oduction by remnant trees, my results were inconsistent at best. Firstly, th e percentage of fruiting trees in most of species were similar between logged and control plots, except for Centrolobium For some species (e.g., Copaifera Aspidosperma and Sweetia ) in which there seemed to be a trend towards increased reproduction among remnant trees in selectively logged forest statistical significan ce was not forthcoming perhaps due to small sample sizes (N < 20). Secondly, 3 of the 6 species I monitored star ting 1.5 y after selec tive logging showed no apparent effect of the treatment on fruit product ion, 2 of the species fruited less in the logged than in the control plot, and 1 species showed the opposite pattern. One reason for the observed decrease in fruit production might have been th e effect of lowered tr ee density on pollinator effectiveness or offspring quality (Ghazoul and Shaanker 2004, Knight et al. 2005). Other factors that might have reduced fruit production include fruit abortion due to poor ovule fertilization in the extreme temperatures and low humid provoked by forest openness (Stephenson 1981, Aizen and Feinsinger 1994, Dafni and Firmage 2000) or due seed predation during the early stages of fruit formation (Stephenson 1981). For example, 1 tree/ha 40 cm DBH of Copaifera remained after logging of the pre-loggi ng density of 1.7 trees/ha whereas in the unlogged plot, the

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26 population of adult trees was subs tantially higher (2.7 trees/ha). Similarly, in Southeast Asia, fruit production was reduced in a logged dipterocar p forest (Curran et al. 1999). In contrast, the most light-demanding tree sp ecies in my study forest, Anadenanthera produced more fruit in logged than in unlogged areas. Similarly, Ceiba aesculifolia (Kunth) Britten & Baker f. in a dry forest in Mexico (Herre rias-Diego et al. 2006) and Quercus costaricensis in montane humid forest in Costa Rica (Guariguata and Saenz 2002) both increased fruit prod uction in response to logging. Given the importance of this issue to sustainable timber stand management, more research is needed on the reproductive responses of trees to logging dist urbances. These studies should integrate flower production, pollination, a nd fruit production and should be conducted on species representing a variety of densities and breeding systems. Conclusions In the seasonally dry tropical forest I studi ed in Bolivia, there was a lot of interannual variation in fruit production among species. Whether or not a reproductively mature tree fruits is generally not related to it size. The number of fruits produced pe r tree also did not consistently change with stem diameter or liana cover, but did increase with crown size. Base on these findings, I recommend protection of tree crowns during logging to increase fruit production. Even when I experimentally killed the lianas infesting the crowns of 16 Caesalpinia trees, I did not observe the expected increase in fruit produc tion. To look at the effect of lianas cutting I recommend a long-term study to look at crow n area recovery and fruit production. Similarly, after logging, remnant trees of some species increased their fruit production and some species decreased but most did not change. For most of the species, percentages of trees fruiting were similar between the control and the logged plot; only in Centrolobium was there a difference, with more trees fruiting in the control plot. The small and inconsistent effects of logging on fruit production in my study forest may be explained in part by the relatively low intensity of

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27 harvesting. Obviously, given the substantial inter-t ree, inter-specific, an d inter-annual variation in the fruiting of dry forest tree species, st udies of more than a few years are needed.

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28Table 1-1. Overview of reproductive char acteristics of the tree species studied in a tropi cal dry forest in Bolivia. Species Family Abbreviation Breeding System Dispersal Type Dispersal Unit Fruiting* Frequency Fruiting Seed Dispersal # Seeds / Fruit Acosmium cardenasii Fabaceae ACCA Hermaphrodite Wi nd Fruit Continuous Mar-Aug, OctFeb May-Sep, Feb 1.2 Amburana cearensis Fabaceae AMCE Hermaphrodite Wi nd Seed Annual Jun-Aug Sep 1 Anadenanthera macrocarpa Mimosaceae ANMA Hermaphrodite Gravity Seed Annual Mar-Sep Jul-Oct 10.9 Aspidosperma rigidum Apocynaceae ASRI Hermaphrodite Wi nd Seed Annual Mar-Sep Sep-Nov Astronium urundeuva Anacardiaceae ASUR Dioecious Wind Fruit Annual Aug-Oct Oct 1 Attalea phalerata Arecaceae ATPH Monoecious Animal Fruit Continuous Oct-Apr Feb-May Caesalpinia pluviosa Mimosaceae CAPL Hermaphrodite Explosiv e Fruit Annual Mar-Sep Ago-Nov 4.4 Capparis prisca Capparaceae CAPR Hermaphrodite Animal Fruit Annual Cariniana ianeirensis Lecythidaceae CAIA Hermaphrodite Wind Seed Annual May-Sep Oct Cecropia concolor Cecropiaceae CECO Monoecious Animal Fruit Continuous Oct-Mar Feb-Mar Cedrela fissilis Meliaceae CEFI Hermaphrodite Wind Seed Annual Jul-Aug† Aug-Sep Ceiba samauma Bombacaceae CESA Hermaphrodite Wi nd Seed Annual Jun-Aug Ago 90.4 Centrolobium microchaete Fabaceae CEMI Hermaphrodite Wind Fruit Annual Apr-Sep Jul-Sep 1-2 Chorisia speciosa Bombacaceae CHSP Hermaphrodite Wind Seed Annual Jul-Aug Sep Copaifera chodatiana Caesalpinaceae COCH Hermaphrodite Gr avity Seed Annual Mar-Sep Jul-Oct 1 Cordia alliodora Boraginaceae COAL Hermaphrodite Wind Fruit Annual Jul-Ago Ago 1 Gallesia integrifolia Phytolacaceae GAIN Hermaphrodite Wi nd Fruit Annual May-Sep Oct-Nov 1 Genipa americana Rubiaceae GEAM Hermaphrodite Animal Fruit Annual Dec-Feb Feb-Mar Hymenaea courbaril Caesalpinaceae HYCO Hermaphrodite Grav ity Fruit Annual Jul-Oct Oct-Nov 3.3 Machaerium cf. acutifolium Fabaceae MAAC Hermaphrodite Wind Fruit Supra-annual Apr-May May 1 Machaerium scleroxylon Fabaceae MASC Hermaphrodite Wind Fruit Annual Apr-Sep Aug-Sep 1 Myrcianthes spp. Myrtaceae MYSP Hermaphrodite An imal Fruit Annual Jan-Feb Feb Phyllostylon rhamnoides Rhamnaceae PHRH Hermaphrodite Wind Fruit Supra-annual Oct Oct 1 Platimiscium ulei Fabaceae PLUL Hermaphrodite Wind Fruit Annual Jul-Aug Aug-Sep 1 Pterogyne nitens Caesalpinaceae PTNI Hermaphrodite Wind Fruit Annual Apr-May May-Aug 1 Schinopsis brasiliensis Anacardiaceae SCBR Dioecious Wind Fruit Supra-annual Ago-Sep Sep 1 Spondias mombin Anacardiaceae SPMO Hermaphrodite Animal Fruit Annual Jan-Feb Feb 1-2 Sweetia fruticosa Fabaceae SWFR Hermaphrodite Wi nd Fruit Annual Oct Oct-Nov 1 Syagrus sancona Arecaceae SYSA Monoecious Zoo Fr uit Annual Nov-Dec† Dec-Jan 1 Tabebuia impetiginosa Bignoniaceae TAIM Herm aphrodite Wind Seed Annual Jul Ago 90.7 Tabebuia serratifolia Bignoniaceae TASE Hermaphrodite Wind Seed Annual Ago Sep Zeyheria tuberculosa Bignoniaceae ZETU Hermaphrodite Wind Seed Supra-annual Jun-Jul Jul 30 (*) Clasification made by Newstron et al. (1994). (†) Data extracted from Justiniano and Fredericksen (2000).

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29 Table 1-2. Results of the best model after b ackward regression steps to remove non-significant variables (P > 0.1) sequentially from the full multiple regression model including tree size (DBH), percent liana infestation, cr own area, and crown position as independent variables to explain fruit production. Number of trees (N ), standardized regression coefficients ( ), Student-t test values (t), signifi cance levels (P), and coefficients of multiple determination (R2) are noted. Variables N tP R2 Anadenanthera macrocarpa Crown Area 30 0.5603.590.001 0.32*** Caesalpinia pluviosa Crown Area 50 0.4803.81<0.0001 0.23** Zeyheria tuberculosa Crown Area 35 0.4502.890.007 0.24* Crown Position 35 0.2821.810.080 P 0.05; ** P 0.01; *** P 0.001.

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30 Table 1-3. Analysis of covariance to determine the liana cutting effect in fruit production of Caesalpinia pluviosa The co-variables considered were crown area and DBH. Source Crown Area DBH Mean Square FP Mean Square F P Co-variable 42438.5 2.350.14 22090.01.18 0.29 Liana cutting 9135.0 0.570.48 9673.00.52 0.48 Error 18020.7 18747.0

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31 Figure 1-1. Locations of the study areas in Boli via. Study Area 1 was where I conducted fruit production research (Chapter 1); Study Ar ea 2 was used for seedling recruitment study (Chapter 2); and, Study Area 3 was the s ite for the sprouting studies (Chapter 3). The map is georeferenced using a Universal Transverse Mecator (UTM) System, Zone 20.

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32 Species CAPLANMAACCACOCHMASCCESAGAINCEMIASRISWFRCAIACAPRMAACZETU % of Trees Fruiting 0 20 40 60 80 100 Year 2003 Year2004 (124)(58)(373) (82) (82) (18) (27) (5) (15) (4) (15) (180) (48) (18) (20) (10) (23) (40) (111) (39)** * * ** Figure 1-2. Percentage of trees fruiting in 2003 and 2004 (note th at none of these species are dioecious). Asterisks indicate years in which a species that was not evaluated. Abbreviations of species are given in Tabl e 1-1. Numbers in parenthesis indicate the number of trees evaluated for each species and year.

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33 Caesalpinia pluviosa 20022003200420052006 Fruit Production / Tree 0 200 400 600 800 1000 Zeyheria tuberculosaYear 20022003200420052006 0 20 40 60 80 Anadenanthera macrocarpa 20022003200420052006 0 100 200 300 400 500 600 Figure 1-3. Annual variation in fr uit production in three timber specie s. Vertical lines show one standard error. Note the diffe rences in the y-axis scales.

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34 Anadenanthera Year 2003 Year 2003 Year 2004 Year 2004 Caesalpinia 0 20 40 60 80 100 120 Centrolobium Y Data Acosmiun 0 20 40 60 80 100 120 Gallesia M. scleroxylon DBH (cm) 0 20 40 60 80 100 120 Swettia Fruiting Trees Aspidosperma Ceiba M. acutifolium Zeyheria Copaifera 0 20 40 60 80 100 120 NO YES NO YES NOYES NO YES NO YES NO YES NO YES NO YESNO YES NO YES NO YES NO YES NO YES NO YES NO YES*** **12 68114 258760 8 5 9924 3458 7 144 3628 19 3 135 8 218 5 18 11 6 69 43 Figure 1-4. Comparison of the sizes (DBH) of fruiting and n on-fruiting trees in 2003 and 2004 (note that none of these species are dio ecious and all the trees were reproductively mature). Numbers indicate the sample size for each year and each species. Asterisks indicate significant size differences be tween fruiting and nonfruiting trees as determined with Student-t test. P-value: 0.05, ** 0.001, *** 0.001.

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35 Zeyheria tuberculosa Crown Area 050100150200250 1 10 100 1020304050607080 Anadenanthera macrocarpa 050100150200250 1 10 100 1000 20304050607080 Caesalpinia pluviosa 050100150200250 Fruit Production (Log10# / tree) 1 10 100 1000 DBH 20304050607080 Log (Y) =1.86+0.01(X), r 2 =0.05, F=1.49, P=0.23 Log(Y) = 1.47+0.007(X), r 2 =0.32, F=12.92, P=0.001 Log (Y) = 1.71+0.013 (X), r 2 =0.06, F=3.19, P=0.08 Log (Y) = 1.64+0.004 (X), r 2 =0.23, F=14.55, P=<0.0001 Log (Y) = 0.59+0.16 (X), r 2 =0.08, F=3.07, P=0.08 Log (Y) = 0.91+0.004 (X), r 2 =0.17, F=6.53, P=0.01 Crown Position Spearman's r=-0.25, P=0.18 Spearman's r=-0.135, P=0.35 Spearman's r=0.012, P=0.95Dom Codom Dom Codom Interm Dom Codom Interm Figure 1-5. Simple linear regressions between cr own area, tree size (DBH ) in relation to log transformed fruit production data for thr ee timber species. Spearman correlations were run to determine the relationships be tween crown position (o rdinal variable) and fruit production. Crown position numbers refer to dominant (Dom), co-dominant (Codom), and intermediate exposure (Interm) trees.

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36 Crown Area (m 2 ) 50100150200250300 Fruit Production (Log10 # / tree) 1 10 100 1000 10000 DBH (cm) 203040506070 1 10 100 1000 10000 Lianas Cut Control Figure 1-6. Fruit production of Caesalpinia pluviosa in relation to DBH and crown area for trees with cut or uncut lianas.

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37 Species CentrolobiumCopaiferaCaesalpiniaAnadenantheraMachaeriumAspidospermaSweetia Percentage of Fruiting 0 20 40 60 80 100 Control Plot Logged Plot *** ns ns nsns ns ns(10) (37) (9) (7) (21) (34)(34) (39) (20) (5)(4) (12) (7) (12) Figure 1-7. Percentages of trees fruiting in a logged and an unlogged plot. Number of trees evaluated is noted in parenthe sis. Differences between loggi ng intensities were tested using X2 test at 95% of confidence. *** = <0.0001; ns = non significant.

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38 Anadenanthera macrocarpa 0 20 40 60 80 100 Aspidosperma rigidum 0 20 40 60 80 100 Normal Logging Control Caesalpinia pluviosa Fruiting Intensity (% Crown) 0 20 40 60 80 100 Machaerium scleroxylonMonths 0 20 40 60 80 100 Copaifera chodatiana 0 20 40 60 80 100 Centrolobium microchaete 0 20 40 60 80 100 May04Jul04 Sep04 Oct04Nov04 May04Jul04 Sep04 Oct04Nov04 May04Jul04 Sep04 Oct04Nov04May04Jul04 Sep04 Oct04Nov04 May04Jul04 Sep04 Oct04Nov04 May04Jul04 Sep04 Oct04Nov04 Figure 1-8. Average fruiting inte nsities (% of crown cover) of tim ber tree species evaluated in areas subjected to normal selective loggi ng and nearby unlogged control areas in a tropical dry forest. Vertical li nes indicate standard errors.

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39 CHAPTER 2 BIOTIC AND ABIOTIC FACTORS AFFECT ING TREE SEEDLING DYNAMICS IN A DRY TROPICAL FOREST Introduction Among the many factors that affect tree seedli ng establishment in tropical forests, seed predation, pathogen effects on seedlings and lig ht availability figure prominently (Janzen 1971, Augspurger 1984, Hammond 1995, Huante et al. 1998, Kobe 1999). Given that this view is mostly supported by studies conducted in moist and wet forests, the relative importances of these factors might differ in seasonally dry forest s (Gerhardt 1994, Holbrook et al. 1995). Given the widespread mismanagement and destruction of these forests (Steininger et al. 2001, Pacheco 2006), it is increasingly important to increase our knowledge of ecological processes, such as regeneration, that might lead to their improved management To further this knowledge, I monitored seedling populations a nd conducted experimental studies across a gradient of forest management intensities in a seasonally dry lowland tropical forest in Bolivia In the experiments, I planted seeds of canopy tree speci es and manipulated moisture av ailability, seed and seedling predation, and competition with an abundant understory bromeliad, Pseudananas sagenarius (Arruda) Camargo). Seasonally dry tropical forest s are characterized by low tota l rainfall (typically < 1500 mm) and annual periods of drought (i.e., mont hs during which evapotranspiration exceeds precipitation usually assumed to be months with <100 mm of precipitation)(Hol dridge 1967). The rainfall regime of Bolivian, and apparently many other tropical dry forests (Gerhardt 1996a), is also characterized by huge in terannual variation in precipita tion, dry season duration, temporal continuity of rainfall during the rainy and dry seasons, and the starting and ending dates of the seasons. In addition, paleoecological and archaeol ogical data predict that the dry seasons in tropical dry forests in South America will be extended in duration and drier in the future (Mayle

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40 et al. 2007). These sources of environmental va riation are critical because, although the tree species characteristic of tropical dry forests are presumably adapted to seasonal drought, mortality is reportedly concen trated during dry seasons and dry years (Khurana and Singh 2001, McLaren and McDonald 2003). Within species and age cohorts, the probability of mortality reportedly decreases with increasing seedling size, presumably because larg er individuals have greater access to soil water (Khurana and Singh 2001). Seedling growth rates t ypically increase with increasing illumination but, due to high temperatures and soil surface drying, seedlings growing in large canopy gaps may suffer high risks of mortality, even if they gr ow larger than their sh aded counterparts in the understory. In other words, in a forest with a 6-month rainy season, a newly germinated seedling has 6 mo to grow large enough to survive the subsequent dry season. Based on this idea, I conducted an experiment in which I manipulated the duration and intensity of the dry season and monitored the survival and growth of seedlings of canopy tree species that germinated from sown seeds. In recognition of the importance of inter-specific competition and seed/seedling predators to seedling dynamics, I also manipulat ed these factors in a factorial experiment. In addition to tolerating drought tropical tree seedlings mu st compete for light, water, and nutrients with plants of various growth forms including lianas (Gerwing 2001, Schnitzer and Bongers 2002), ferns (George and Bazzaz 1999), a nd ground bromeliads (Fredericksen et al. 1999, George and Bazzaz 1999). In the seasonally dr y forests of the Chiquitania of lowland Bolivia, competition with a clonal ground bromeliad, Pseudananas sagenarius seems particularly intense. This bromeliad covers 2530% of the ground surface in my study area where it presumably competes for light and soil resources Furthermore, its leaves intercept both rain and falling seeds (Fredericksen et al. 1999), wh ich should affect tree seedling recruitment,

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41 growth, and survival. Finally, by providing cover to small mamma ls and other animals avoiding predators, bromeliads may also affect rates of seed and seedling predation. To investigate these effects, I included a bromeliad removal treatment in my experimental study on the survival and growth of canopy tree seedlings. Predators often reduce seed and seedling survival in trop ical forests (Janzen 1971, Hulme 1996, Asquith et al. 1997). Given that seed pred ation rates, at least by mammals, typically increase with seed size (Mol es and Westoby 2003), and given th at most tree species in dry tropical forests have small, wind-dispersed seeds, rates of seed predati on may be lower in dry than in humid forests. On the other hand, given the seasonal scarcities of f ood in dry forests, seed and seedling predation might be pa rticularly intense, especially for seedlings that remain leafed out and succulent during the early dry season. To investigate the importan ce of seed and seedling predation, as influenced by seasonality, I experi mentally manipulated both soil moisture and accessibility to predators using the pl anted seeds of canopy tree species. In this chapter I present the results of a study in which I monitored the survival and growth of tree seeds planted in plots in which I experimentally manipulated soil moisture, seed and seedling predator access, and competition wi th ground bromeliads. The chapter also includes the results of 3 y of monitoring of seedling establishment, growth, and survival in a large control plot and two otherwise similar pl ots subjected to different intens ities of timber harvesting 7 mo prior to commencement of my study. I hypothesi zed that seedling survival and growth are reduced by bromeliad cover, drought stress, a nd predators. I also hypothesized that the establishment, survival, and gr owth of naturally regenerate d seedlings differs among plots subjected to different intensitie s of disturbance resulting from fo rest management activities.

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42 Methods Study Area and Climate This study was conducted during 2003-2006 in th e 30,000 ha seasonally dry lowland forest (Holdridge 1967) owned by INPA PARKET, about 30 km from Concepcin, Bolivia (16 6' 45"S, 61 42' 47"; Figure 1-1). The study area is located in the transition zone between the forests of the Amazon Basin and those of the Gran Chaco and is locally referred to as Chiquitano dry forest (Killeen et al. 1998, Ibisch and Mrida 2003). Based on 30 years of data collected in Concepcin, the annual average temperature in the area is 24.3 C w ith a minimum average temperature of 12.9 C, generally in July, and maximum average temperature of 31C, generally in November. Over the 30-year monitoring peri od, the annual average of rainfall was 1100 mm but ranged from 700-2000 mm/year, with wet year s corresponding with El Nio events (Coelho et al. 2002) (Figure 2-1). Both 2005 and 2006 we re dry years, with only 980 and 1050 mm of precipitation, respectively. The ra iny season typically runs from October to April, but in 2005, the rains started in September and rain fell at regular intervals until March of 2006. The 2006 rainy season, in contrast, started in October but then no rain fell in November, with more regular rains commencing again in December. The set of Long-Term Silvicultural Research Pl ot (LTSRPs) establishe d at this site by the Instituto Boliviano de Investig aciones Forestales (IBIF) was used for a portion of this study. The LTSRP at INPA includes four 20-ha plots that vary in manageme nt intensity: unlogged control; normal logging; improved management; and, intens ive management (Mostacedo et al. 2006). For this study, the improved management plot was not used. In the “normal logging” plot, the logging company harvested a mean of 4.3 trees/ha (4.7 m3/ha) using their standard reducedimpact-logging techniques that include road plan ning, directional felling, and the retention of 20% of the harvestable trees as seed trees. The mean density of trees 10 cm DBH before

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43 logging was 427 individuals/ ha, of which 40% were Acosmium cardenasii (after first mention of a species I will refer to it by its generic name; for a complete list of scientific names see Table 21). The company used these same logging techniqu es in the “intensive management” plot to harvest a mean of 8.1 trees/ha (8.2 m3/ha). While harvesting the “int ensive management plot,” the skidder drivers mechanically scarified the soil su rface in an average of 0.6 felling gaps/ha (mean area = 50 m2/ha) where there was no existing regene ration of commercial timber species. After logging of the intensive management plot future crop trees (i.e., well-formed trees of commercial species 10-40 cm DBH) were liberated from liana cover (by slashing the lianas with a machete; 21 trees/ha) and liberated from co mpetition from nearby noncommercial trees (by poison girdling; 1.7 trees/ha). These last two treatments for enhanc ing the growth of future crop trees were not applied in the norma l logging plot. In the unlogged fo rest plot, the mean density of trees > 10 cm DBH was 432 indivi duals/ha, of which 38% were Acosmium (Table 2-1). The total basal area of trees >10 cm DBH in the unlogged plot averaged only 19.6 m2/ha and 38% of the trees were liana infested (18% severely so). Canopy openness in the early dry season (May 2003), as measured with a spherical densiometer (Lemmon 1957) held 1 m above th e ground at 140 equally spaced points in the 10 ha permanent plots, averaged 8% in the 20 ha co ntrol (unlogged) plot, 13 % in the plot subjected to normal selective logging in November 2002, and 14% in the plot that was intensively harvested and silviculturally treated also in November 2002 (Figure 2-2). I remeasured canopy openness at the same points in March 2006 and f ound increases in canopy cover in both of the treated plots, but not in the control plot (Figure 2-2).

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44 Experimental Design and Data Analysis Response of regeneration to s ilvicultural treatment intensity Plants 5-100 cm tall of 22 species of sub canopy, canopy, and emergent tree species (Table 2-2) that originated from seeds or sprouts were monitored in subplots in three of the 20-ha (400 x 500 m) LTSRP plots. In the centr al 10 ha (400 x 250 m) of each pl ot, I located 144 pairs of 2 x 1 m subplots separated by 2 m at 25 m intervals (data from the pair ed plots were subsequently combined). Each seedling was marked, mapped, and measured for height at regular intervals of one year for 3 y. The site of each plot pair wa s categorized in one of the following microsites: undisturbed forest (includes na tural canopy gaps and high statur e forest); logging road; and, logging gap. Undisturbed forests we re categorized as t hose sites in which th e structure of the forest did not change during loggi ng. I initially separated skid tra ils from primary and secondary logging roads, but because of small sample sizes, these sites are combined into a category referred to as log extraction paths. Logging ga ps were sites where the canopy was opened during tree felling and log extraction. Based on th e literature (Whitmore 1998, Mostacedo and Fredericksen 1999, Pinard et al. 1999) and field observations, each tree species was placed in one of the following ecological groups: light-demandi ng pioneer; long-lived pioneer; partially shade tolerant; or, shad e tolerant. I compared seedling abundances among the harv esting treatment plots and microsites using repeated measures ANOVA (Scheiner and Gurevitc h 2001). Measurement dates were considered to be a within-subject factor, while harvesting treatment and microsites were considered as between-subject factors. Seedling recruitment rates ( R ) were calculated using the compound interest equation (McCune and Cottam 1985):

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45 1 ) 1 (/ 1 xBx R (2-1) where Bx is the birth rate in the period x calculated for each year and for each m2 of ground area. As recruitment rate I calculated the number of new seedlings of each species in each harvesting treatment m2/year. Mortality rate ( M ) expressed as percentage per year wa s calculated as (Primack et al. 1985, Sheil et al. 1995) tN N N M/ 1 0 1 0] / ) ( 1 [ 1 (2-2) where N0 is the number of living seedlings at time 0, N1 is the number of living seedlings at time N1, and t is the time period between N0 and N1. Mortality rates were calculated by species and harvesting treatment. I calculated the relative growth rate (RGR) as 0 1 0 1)) ( ) ( ( t t H Ln H Ln RGR (2-3) where, H0 is the seedling at the initial measurement, H1 is the height at the second measurement, t0 is the initial time, and t1 is the time of the second measurement (Hoffmann and Poorter 2002). I compared RGR of harvesting trea tments, microsites and species using repeated measures ANOVA. For all analyses I used SPSS Version 12.0. For each factor, I also conducted Bonferoni or LSD post hoc comparisons at the 95% confidence level. Factors affecting seedling establishment and growth Tree species used in the following experimental studies were selected on the basis of seed availability in May-September 2005 when I collec ted seeds from 3-10 trees per species. Of the six species used in the experiment, 4 are wind-dispersed, 2 are dispersed by animals, 2 are

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46 characterized as light-demanding, and 4 are c onsidered shade tolerant (Table 2-1). Brief descriptions of the species follow: Amburana cearensis (Allemo) A. C. Sm. usually produces one 0.49 g dry weight winged seed per dehiscent legume. Aspidosperma rigidum Rusby produces elliptical seeds that weigh 0.24 g and are surrounded by a wing. Ceiba samauma (Mart.) K. Schum. produces 0.17 g seeds that are covered by fine, cotton-like fi bers that aid in wind dispersal. Pterogyne nitens Tul. produces single-seeded wind-disper sed samaras with 0.16 g seeds. Copaifera chodatiana Hassl. seeds are red, weigh 0.33 g, are covered with a white aril, and are animal-dispersed. Hymenaea courbaril L. produces round, large (3.9 g) and animal-dispersed seed s in indehiscent legumes. The experiment used a split-plot design (M ontgomery 2001) in which ground bromeliad cover, water availability, and seed and seedli ng predation were experimentally manipulated (Figure 2-3). The 2 m2 experimental plots were comprised of the two 1 m2 subplots which were replicated 40 times in 10 blocks separated by at least 40 m. The experiment commenced in July 2005 and was monitored until December 2006. The terrestrial bromeliad manipulated in this study is Pseudananas sagenarius locally known as “garabata.” Rosettes of this cl onal species are about 1 m high and cover 1m2 each. Their colonies typically cover 15-40% of the ground in Chiquita no dry forest in clumps up to 2000 m2. This species is more common in mature th an in young stands (Kennard et al. 2002) and its abundance is reduced by fi re (Fredericksen et al. 1999). The water availability treatment involved either irrigating or withholding water from plots by adding water or shieldi ng plants from rainfall during what is typically the transition between the dry and rainy seasons (late Septembe r). Rainfall inputs directly into the drought treatment plots were prevented by covering them with 2.0 x 1.5 m roofs of transparent plastic.

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47 The rain shelters were left for the first 3 mo of what is normally the dry season. The irrigated plots received natural rainfall plus 10 liters of water every 15 days (=20 mm/mo) starting on 21 September 2005 and finishing in at the end of Nove mber when rainfall were more frequent. Soil water tensions at 0-10 cm depth in one droughted and one irrigated plot in each of four blocks were monitored hourly with Watermark soil moistu re sensors attached to data loggers; rainfall inputs were monitored adjacent to the four irrigated plots (Figure 2-1). Droughted plots simulated years with prolonged dry seasons wherea s the irrigated plots simulated wet years with short dry seasons. To exclude seed and seedling predators, half of the plots were surrounded by 60 cm high, 1-mm mesh wire netting fastened to the ground. The exclosures were erected before the seeds were sown and were maintained for the duration (15 mo) of the experiment In each treatment plot, I randomly sowed seeds of all tree species described above (Table 2-1). Each seedling establishing from a planted seed was marked, mapped, and its height measured five times between September 2005 and December 2006. Treatment effects on seedling abundance were tested using repeated measures ANOVA for a split plot design (Scheiner and Gurevitch 2001) with bromeliads (present or absent) as the main factor, and irrigation, predator exclosure, and time as subplot factors. Seedling heights (RGR) for each time period were compared between treatments for each factor using t-test with sequential Bonferroni corrections (Sankoh et al. 1997). Due to seedling mortality over the study, the sample sizes varied so I could not run repeated meas ures ANOVA on the RGR data. I ran the analysis for all 6 species combined and for each species separately.

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48 Results Response of Regeneration to Management Intensities Three years after logging and silvicultural treatment, seedling densities varied with management intensities and am ong microsites. Although 22 species are included in Table 2-1, only data from the most abundant 11 species are presented. Densit ies were higher in the control than in the harvested areas, but did not differ between the normal harvesting and the intensive management plots (Figure 2-4). Ho wever, seedling densities without Acosmium was higher in normal harvesting and lower in the control plot (Figure 2-4). This pattern was maintained five years after logging for all species combined, but for nine of the 11 species evaluated, there were treatment differences and temporal trends (Table 2-2). For example, Copaifera seedling density increased with increasing manage ment intensity. In contrast, Machaerium acutifolium Vogel seedling densities did not differ between the two l ogged plots but were sign ificantly higher in the normal logging plot than in the co ntrol plot. In contrast, seedli ng densities were lower in the intensive management plot than in the normal logging or control plot for four Acosmium Phyllostylon rhamnoides (J. Poiss.) Taub., and Machaerium scleroxylon Tul. Seedling densities varied mark edly among microsites. Overall, undisturbed sites had the highest seedling density (Table 2-3). This pattern was obser ved at the species level for Acosmium Caesalpinia pluviosa M. scleroxylon and Phyllostylon. In contrast, Centrolobium microchaete seedling densities were highe r on logging roads than in ot her microsites. The other nine species had sta tistically similar seedling densities among microsites (Table 2-3). Overall, seedling mortality rates were higher in undisturbed areas than in harvested areas and the mortality rate was higher in 2004 than in 2005 (Figure 2-5). In 2004, 47% and 66% of the seedlings died in the normal and intensive management plot, respectively. In 2005, mortality was still higher in undisturbed control plots, bu t the ratio was reduced (Table 2-4). At species

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49 level, mortality rates were generally higher in un disturbed control plots th an in either of the logged plots. On average, the highest (22-26%/y) mortality rates were for Sweetia fruticosa Spreng Caesalpinia, and M. scleroxylon At the other end of the spectrum, Gallesia integrifolia (Spreng.) Harms and Piptadenia viridifolia (Kunth) Benth. suffered no mortality (Table 2-4). Over all species, the recruitment of new seedlings was 0.8 seedlings /m2 /y. Recruitment was three times higher in undisturbed areas than harvested areas (Figure 2-5) and marginally higher (0.91 seedlings /m2 /y) in 2004 than in 2005 (0.63 seedlings /m2 /y). Acosmium had the highest recruitment rate (0.26 seedlings/m2/y), followed by Phyllostylon (0.13 seedlings/m2/y) and M. acutifolium (0.08 seedlings/m2/y). Acosmium and M. acutifolium recruited mostly in undisturbed forest, Phyllostylon recruits were common in the normally logged plot, and Pterogyne and Aspidosperma recruited most new seedlings in the intensive management plot (Table 2-4). Factors Affecting Seedling Growth and Establishment Seedling growth Relative growth rates of seedlings differed among the three logging microsites either in 2004 (F=4.19, P=0.01) or in 2005 (F=10.76, P=<0 .0001). In both years, RGRs were higher on loging roads and logging gaps than in undisturbed micros ites. RGRs were higher in 2004 than in 2005 (F=5.64, P=0.018) and were higher for long-liv ed pioneer species RGRs than partial shade tolerant and shade tolerant species (F=15.63, P=<0.0001). RGRs differed among years, species, and mi crosites following patterns that were not consistent. For example, the RGRs of Acosmium Aspidosperma Copaifera, and Sweetia were higher in 2004 than in 2005 (F=7.74, P=0.005). For Aspidosperma and Caesalpinia, RGRs did not differ among undisturbed sites, logging roads, and logging ga ps. Among microsites, RGRs for M. acutifolium were higher in logging roads and loggi ng gaps than in undisturbed microsites

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50 in 2004 but not in 2005. RGRs of Acosmium and Sweetia were similar among microsites in 2004 whereas in 2005, seedlings grew faster in loggi ng roads and logging gaps than in undisturbed microsites. Monthly RGRs of seedlings in the experiment al plots were similar in bromeliad-free and bromeliad-infested plot s at different times (t1=0.14, P1=0.89; t2=2.01, P2=0.05; t3=0.48, P3=0.63; t4=0.04, P4=0.39). Similarly, RGRs were did not differ in the droughted and irrigated plots (t1=0.89, P1=0.37; t2=0.87, P2=0.39; t3=0.93, P3=0.35; t4=0.88, P4=0.38) or between the exclosure and non-exclosure plots (t1=0.01, P1=0.99; t2=0.12, P2=0.90; t3=0.40, P3=0.69; t4=0.36, P4=0.72; Figure 2-10). At the species level, RGRs were also similar in bromeliad-free and bromeliadinfested plots, droughted and irrigated plots, a nd exclosure and non-exclosure plots (Table 2-6, 2-7, 2-8). Seedling establisment In general, tree seedling establishment was suppressed by bromeliads, but responses to bromeliad cover varied among species (Figure 2-6) Based on the six spec ies for which there are sufficient data, four had similar numbers of s eedlings in bromeliad patches and bromeliad-free areas. For the other two species ( Hymenaea and Pterogyne ), seedling abundances were lower in bromeliad covered plots (T able 2-5, Figure 2-7). The effects of the drought and irrigation trea tments varied over time (Figure 2-6). Seedlings were initially about 150% more abundant in the irrigated than in the droughted plots (Figure 2-8), but this difference diminished over time. This pattern was consistent among species except for Aspidosperma which had more seedlings in the dr oughted than in the irrigated plots (Table 2-5). Excluding seed and seedling predators genera lly resulted in higher seedling densities (Figure 2-6), with some varia tion among species (Figure 2-9). Amburana Copaifera, and

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51 Hymenaea seedlings were more common within the excloures than in the open-access plots (Table 2-5). In contrast, the exclosure trea tment had no effect on seedling densities of Aspidosperma Ceiba, and Pterogyne. There were some significant interactions am ong factors both overall and at the species level. In particular, irrigated plots with bromeliads from whic h predators were excluded had higher seedling densities than plot s that were irrigated and open to seed and seedling predators (Table 2-5). At the species level there were significant interactions among treatments only for Ceiba and Copaifera For Ceiba in the bromeliad covered plots, irrigation and pred ator exclosure increased seedling densities relative to the droughted and open-access plots. For Copaifera in the bromeliad-infested plots, irriga tion and predator exclosure resulte d in higher seedling densities. In the bromeliad-free plots, drought coupled with predator exclosure re sulted in higher seedling densities than drought alone. Discussion Logging Effects on Seedlings Dynamics Logging usually affects seedling densities due to direct effects and cr eation of microsites that differ in environmental conditions, especi ally light intensities (Beaudet and Messier 2002, van Rheenen et al. 2004). In most tropical forest s, seedling densities of light-demanding species increase on open microsites. In this study, how ever, overall seedling densities were higher in undisturbed forest than in selec tively logged areas (Figure 2-2). Si milarly, total seedling densities were higher on undisturbed microsites compared to microsites created by l ogging. In contrast, at a site only 40 km from INPA, with a similar fo rest type and rainfall re gime Fredericksen and Mostacedo (2000) reported that seedlings of comm ercial timber species increased in abundance on log landings, roads, and logging gaps.

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52 At the species level, the majority of speci es (64%) had similar seedling densities among microsites; only three shade-tolerant specie s had higher seedling densities in undisturbed microsites. One light-demanding species, Centrolobium had higher seedling densities on skid trails and log landings compar ed to other microsites. There are several potential explanations for obs erved differences betwee n the results of this study and those of Fredericksen and Mostacedo ( 2000). The forest canopy of Las Trancas where the latter conducte d their work is dominated by Anadenanthera a light-demanding species, whereas INPA Parket is dominated by Acosmium which is more shade tolerant. Results without Acosmium show that harvested plots have higher seed ling densities than control plot (Figure 24). The simple difference in dominants might account for some of the overall differences observed between the sites. Another diffe rence between the two studies is that Anadenanthera seedlings emerging from seeds dispersed immedi ately before logging were extremely abundant at Las Trancas ( 100-200 individuals/m2; personal observation). In contrast, seedlings of Acosmium were never as abundant at INPA and thos e that were present were apparently the result of several years of recruitment. Open microsites created by l ogging may be disadvantageous for seedling recruitment in INPA Parket because such sites are extremely hot and dry, especially during the dry season. On the other hand, it is possible th at even though there were distur bances associated with fallen trees, roads, and skid trails, th e canopy could still be too closed at INPA to promote abundant seedling recruitment (Jackson et al. 2002). In add ition, the densities of understory plants and other competitors might remain high in gaps, thus suppressing seedling recruitment in disturbed areas, especially of shade-intolera nt species (Beckage et al. 2000, Schnitzer et al. 2000). On the log extraction paths, soil compaction might negati vely affect seedling es tablishment and growth.

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53 Finally, several species in this study are shade-tolerant, including the dominant ( Acosmium ), and do not require disturbed ar eas for regeneration. Seedling recruitment rates were 70% lower in the logged plots than in the unlogged control plot and recruitment decreased with management intensity. B ecause of increases in light availability associated with l ogging, I expected mo re recruitment in logged plots than in unlogged plots. One explanation for this result could be that logging decreased seed-tree densities, which limited seed inputs (Forget et al. 2001, Makana and Thomas 2004, Grogan and Galvao 2006). Similarly, in French Guiana, Forget et al. (2001) reporte d that for all but one species, recruitment was not favor ed by logging. The importance of retention of seed trees was shown by Grogan and Galvao (2006), who reported that mahogany ( Swietenia macrophylla ) seed production near gaps was important for recruitment. In addition, even if seed production is not limiting, logging may create microsites that ar e not appropriate for seed germination and seedling growth due to soil compaction (Pinard et al. 1996) or other fact ors. Site preparation (e.g., soil scarification) in logging gaps and sk id trails can enhance regeneration, as shown by Pinard et al. (1996) in a diptero carp forest in Malaysia, but such treatments do not always have beneficial effects (He uberger et al. 2002). Seedling mortality rates were nearly 50% lowe r in the logged plots than in the unlogged plot, which was the opposite of what I expected. Several studies report that seedling mortality usually increases as harvesting intensity in creases (Chapman and Chapman 1997, Saenz and Guariguata 2001), but the l ogging intensities in my plots were al l low. Nevertheless, mortality of some plant species decreases as light intensity incr eases. For example, in a tropical rain forest in Costa Rica, Kobe (1999) found that mortality rates of three of four canopy species monitored decreased as light levels in creased in the understory.

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54 There was considerable interspecific varia tion in both recruitment and mortality rates. For example, for three tree species ( Acosmium Aspidosperma and Copaifera ) that are in the same shade tolerant guild the percentage of recruitm ent ranged from 7.5-15.5 %/y. The mortality rates for the same species ranged from 0.019-0.255 individuals/m2/y. Acosmium, because of its dominance, drives the overall patter ns of recruitment and mortality. Apparently, while recruitment rates are redu ced by disturbance, the seedlings that do establish grow sufficiently duri ng the rainy season to survive the subsequent dry season. In contrast, in the relatively closed conditions of the cont rol plot, few of the abundant new recruits survive the drought stress of thei r first dry season. Similarly, seedlings of a light-demanding species ( Anadenanthera ) established abundantly, but died rapidly in the control plot. Temporal changes in recruitment and mortalit y are related to varia tion in seed production and rainfall regime (Kitajima and Fenner 2000, Khurana and Singh 2001). In this study, seed production, seedling recruitment, and seedling mo rtality rates were all higher during 2003-2004, a dry year, than during 2004-2005, a relativel y wet year with a short dry season. Within the logged plots, seedling RGRs were t ypically higher in disturbed microsites, but there was some variation among species. Over th e 2 y monitoring period, seedlings grew faster on log extraction paths than undisturbed microsites In a similar study in the same area but using large transects, seedlings in skid trails and logging gaps grew faster than seedlings in undisturbed forest (van Andel 2005). Cont rary to my expectations, Acosmium a shade-tolerant species, had the highest RGR on log extrac tion paths while RGRs of Anadenanthera, a light-demanding species, not vary among microsites. Overall it appears th at disturbed microsites are favorable for seedlings, but this pattern seems to be driven by Acosmium, the most abundant species. For most of the species it seems that, in contrast to wett er tropical forests, RGRs of tropical dry forest

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55 seedlings are not governed primarily by light ava ilability and have a di fferent relationship to disturbance than wet and moist fore st tree species (K hurana and Singh 2001). Factors Affecting Seedling Establishment and Growth Soil moisture availability is a key factor influencing seedling gr owth in dry forests (Khurana and Singh 2001). In my study, seedlings th at lost their leaves during the dry season (e.g., Amburana and Aspidosperma ) grew slowly, while the evergreen species (e.g., Copaifera and Pterogyne ) grew faster. I also found that brome liad cover significantly reduced seedling growth rates. Ground bromeliads interfered with the establishment and growth of tree seedlings in this study, as has been described for another forest in Bolivia (Fredericksen et al. 1999), as well as on Barro Colorado Island in Panama (Brokaw 1983). As in those other two studies, I also observed substantial interspecific variati on in seedling responses to brom eliads. The observation that the irrigation treatment eliminated the generally nega tive effect of bromeliads on seedling survival and growth suggests that the rain -trapping effect of bromeliad ro settes, coupled with any water they draw from the soil, is the cau se of their deleterious effect. Pe rhaps the species that were not sensitive to the pres ence of bromeliads ( Amburana Ceiba Copaifera and Pterogyne ) are more drought-tolerant than the other species. I expected that experimentally shortening the dry season by irrigation would increase the establishment and growth of tree seedlings, wh ile exacerbating dry season drought would have the opposite response. Instead, what I found that the principal imp act of the drought treatments was the timing of seed germination, but not a ll species were affected Seeds in the droughted plots did not germinate until after I removed th e roofs and allowed rain to fall on the soil; seedlings then reached the same densities as in the irrigated plots. A similar study in mahogany

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56 in a dry forest in Guanacaste, Costa Rica report ed the same effect of water availability on seedling density (Gerhardt 1996b). Similarly, RGR were not affected by drought stress. The tendency for species overall (except Aspidosperma ) was higher densities in the irrigated than in the droughted plots at the tim e of germination. This tendency suggests that longer rainy seasons, such as those occurring duri ng El Nio years, can ha ve dramatic effects on seedling establishment (Gilbert et al. 2001, Engelb recht and Kursar 2003). The same pattern of increased mortality during dry years was revealed by the mortality of naturally established seedlings during this study (Figure 2-5). Due to a variety of morphological, physio logical, and phenological traits, tree species differ in the susceptibility of th eir seedlings to drought stress (T able 2-4; (Reader et al. 1993, Khurana and Singh 2001, McLaren and McDonald 2003). For example, Amburana seedlings reduce their moisture requirements by be ing dry season deciduous, leaf areas of Aspidosperma seedlings decline substantially during the dry season, and Acosmium is noteworthy for its deep root system (Wright 1991, Engelbrecht and Kursar 2003). Seed or seedling predation seems to be a very important factor for seedling establishment in tropical dry forests (Janzen 1971, Hulme 1996). I found that plots with exclosures had higher seedlings densities than open plots. Exclosures prevented seedling herbivory, but not all types of seed predation because insects and other small se ed predators were not excluded (Grol 2005). In tropical dry forests, herbivory has the most impact on tree re generation when seedlings are emerging, not after roots and st ems are lignified (Kitajima an d Augspurger 1989, Lucas et al. 2000). Once browsed, established seedlings usua lly resprout, but only lignified seedlings survive. In some cases, herbivory damage could be superficial and not dramatically impact the growth and survival of seedlings (personal observation).

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57 There were some species in which exclosur e treatments had an effect on established seedlings, while for others there were similar. For example, Hymenaea a species with a large seed and seedlings larger than Ceiba had a higher established s eedling caused by the exclosure plots. Several studies su pport the hypothesis that la rge seeds are usually most predated (Blate et al. 1998, Dalling and Hubbell 2002), but s ee Moles et al. (2003). Likewise, Aspidosperma and Pterogyne have seeds that probably were predated mostly by small insects (i.e., bruchid beetles) which were not deterred by the exclosures. Conclusions Five years after selective l ogging and silvicultural treatments, overall tree seedling densities were higher in the contro l plot than in the two logged plot s. It is important to point out that this result was heavily infl uenced by the reduced seedling dens ities in the logged plots of the dominant species, the shade-tolerant Acosmium Other species, which were less common everywhere (e.g., C entrolobium Copaifera, and Machaerium cf. acutifolium ), actually increased in response to forest management activities, a nd many other species were not affected. Within the logged plots, undisturbed microsites ha d higher seedling de nsities except for Centrolobium a light-demanding species that sprouts readily fr om damaged lateral roots, which was more abundant on log extraction paths. Seedling recruitment rates were higher in the unlogged control plot and on undisturbed microsites in the logged plots, but seedling in thes e areas also suffered higher mortality rates than seedlings in disturbed areas. The higher tur nover rates of seedlings under closed canopy conditions coupled with their observed lower grow th rates suggests that rapid growth under open conditions during the rainy season is critical for seedling surviv al during the subsequent dry season. This result is somewhat surprising give n that, even undisturbed fo rest in my study area, the canopy is 30-40% open after leaf fall.

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58 Mortality rates of naturally established seed lings varied greatly among species. Seven of 22 species suffered no mortality during the 2-y monitoring peri od, whereas relatively high mortality rates were observed for Caesalpinia (26%/y) Sweetia (25%/y), and Machaerium scleroxylon (22%/y). Mortality rates of these species in the control and logged plots followed similar patterns. Bromeliad competition and seed/seedling predators greatly reduced seedling recruitment in this tropical dry forest. Experimentally augm ented soil moisture also increased seedling establishment, but only in interaction with bromeliad removal or predator exclosure. Experimental droughting delayed germination but did not influence seed ling densities once the treatment terminated and rainfall was allowed into the plots. Despite these general trends, species varied in their sensitivities to brom eliads, drought stress, and predators. Finally, seedling growth was not promoted by e xperimentally lengthening the rainy season. Nevertheless, the larger seedli ngs that developed during the ex tended rainy season were more likely to survive the subsequent dry season. Given that global change models consistently predict that the study region will receiv e less rainfall and suffer longer dry seasons in the future (e.g., Mayle et al. 2007), regeneration failures ar e likely to become more common.

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59Table 2-1. Spatial distributions, crow n position, ecological group, geographical ra nge, tree densities and basal area of timbe r species in a Bolivian tropical dry forest. Scientific Name Family Spatial Distribution Crown Position Ecological Group Deciduous Range Stem Density (#/ha >10 cm DBH) Basal Area (m2/ha) Acosmium cardenasii Fabaceae Homogeneous SC TS None R 159.25 6.89 Amburana cearensis* Fabaceae Random CA PS Ma y-Aug W 0. 005 0.006 Anadenanthera macrocarpa Mimosaceae Homogeneous EM L Jun-Oct W 13.67 2.00 Aspidosperma rigidum.* Apocynaceae Clumped SC PS Jul-Sep I 12.92 0.40 Astronium urundeuva Anacardiaceae Clumped CA L Jul-Sep W 0.08 0.07 Caesalpinia pluviosa Caesalpinaceae Homogeneous CA PS Aug-Sep I 15.17 1.18 Ceiba samauma* Bombacaeae Random CA L Jul-Aug W 0.67 0.07 Centrolobium microchaete Fabaceae Clumped CA L Jun-Oct I 11.42 0.55 Chorisia speciosa Bombacaceae Random SC L Jul-Aug W 16.67 0.82 Copaifera chodatiana* Caesalpinaceae Random EM TS None R 5.33 0.51 Cordia alliodora Boraginaceae Random SC L Jun-Aug W 2.83 0.05 Hymenaea courbaril* Caesalpinaceae Random CA PS None W 0.17 0.06 Machaerium acutifolium Fabaceae Homogeneous SC LL None I 27.17 0.34 Machaerium scleroxylon Fabaceae Random EM PS Jul-Sep I 4.25 0.68 Phyllostyllon rhamnoides Rhamnaceae Clumpled SC TS None I 4.42 0.63 Pterogyne nitens* Caesalpinaceae Homogeneou s CA LL None W 0.17 0.01 Schinopsis brasiliensis Anacardiaceae Random EM LL Aug-Sep I 0.25 0.144 Sweetia fruticosa Fabaceae Random CA PS Jul-Sep I 12.8 0.22 Tabebuia impetiginosa Bignoniaceae Homogeneous CA LL Jul-Sep W 1.83 0.27 Zeyheria tuberculosa Bignoniaceae Random CA PS Oct-Nov I 2.75 0.04 Others 142.90 4.67 Total 434.00 19.60 Crown position: (EM) = emergent, (CA) = canopy, (SC) = sub-canopy. Ecological group: (L) = light-deman ding pioneer, (LL) = Long-lived pioneer, (PS) = partially shade tolerant, (TS) = shade toler ant. Geographical range: (R) = re stricted, (I) = intermediate, (W) = widespread. Re stricted species are those that are found only in one type of forest and are limited to the Chiquitano dry forest in Bolivi a. Intermediate species are thos e that are found in two or thre e forest types but are either rest ricted to Bolivia or are only rarely found in neighboring countri es. Widespread species are fou nd in several forest types and in other countries. Species marked with asterisks were used in the seedling survival experiment.

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60 Table 2-2. Means of seedling densities in an unharvested control plot, a plot subj ected to normal timber harvesting, and a plo t subjected to more intensive ha rvesting followed by silvicultural treatments. S ubplots within the main treatment plots are treated as replicates. Differen t letters indicate stat istical differences (P<0.05) among harvesting treatments as indicated by repeated measures ANOVAs and LSD post hoc tests. Scientific Name UnharvestedNormal Logging Intensive Management Mean SquareFP Acosmium cardenasii 1.42(0.09)c1.03(0.09)b0.63(0.09)a64.8918.900.0001 Anadenanthera macrocarpa 0.09(0.01)b0.06(0.01)b0.01(0.01)a0.678.150.0001 Aspidosperma rigidum 0.01(0.02)b0.06(0.02)a0.11(0.02)a1.115.330.005 Caesapinia pluviosa 0.12(0.02)b0.09(0.02)b0.02(0.02)a1.1711.500.001 Centrolobium microchaete 0.02(0.01)a0.01(0.01)a0.031.890.170 Chorisia speciosa 0.005(0.003)a0.01(0.003)a0.002(0.003)a0.012.310.100 Copaifera chodatiana 0.10(0.04)b0.05(0.04)b0.32(0.04)a8.4012.330.0001 Machaerium acutifolium 0.04(0.13)c1.31(0.14)b0.72(0.13)a166.7522.170.0001 Machaerium scleroxylon 0.12(0.01)b0.004(0.01)a0.001(0.01)a1.8548.460.0001 Phyllostylon rhamnoides 0.31(0.02)b0.00(0.02)a0.01(0.02)a12.6864.740.0001 Sweetia fruticosa 0.03(0.01)a0.02(0.01)b0.05(0.01)a0.072.530.080

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61Table 2-3. Mean seedling densities (#/m2) for 3 years (2003-2005) of commercial tree species in microsites created by logging in a Chiquitano dry forest. Sample size for each mi crosite type is noted within brackets; standard errors of the means are noted in parenthesis. Seedling densities in the different microsites were compared with repeated measures ANOVA with microsites as the between-subject factor and time as the within-subject factor Different letters indicate different significance between treatments using pairwise Least Significant Difference (LSD) tests. Scientific Name Logging Gap [100] Log Extraction Path [54] Undisturbed [337] Mean Square F P Acosmium cardenasii 0.701 (0.103)b 0.389 (0.140)b 0.945 (0.056)a 24.92 7.88 <0.0001 Anadenanthera macrocarpa 0.018 (0.015) 0.049 (0.021) 0.050 (0.008) 0.119 1.71 0.18 Aspidosperma rigidum 0.044 (0.024) 0.026 (0.033) 0.056 (0.013) 0.07 0.39 0.68 Caesalpinia pluviosa 0.024 (0.015)b 0.037 (0.020)b 0.073 (0.008)a 0.32 4.71 0.009 Ceiba samauma 0.000 (0.001) 0.000 (0.001) 0.001 (0.000) 0.0001 0.68 0.50 Centrolobium microchaete 0.008 (0.005)b 0.022 (0.007)a 0.005 (0.003)b 0.02 2.86 0.05 Chorisia speciosa 0.004 (0.003) 0.002 (0.004) 0.004 (0.002) 0.001 0.23 0.79 Copaifera chodatiana 0.092 (0.045) 0.086 (0.061) 0.146 (0.024) 0.51 0.85 0.43 Gallesia integrifolia 0.000 (0.003)b 0.000 (0.004)b 0.007 (0.002)a 0.009 3.69 0.03 Machaerium acutifolium 0.609 (0.133) 0.668 (0.181) 0.519 (0.072) 2.13 0.40 0.67 Machaerium scleroxylon 0.008 (0.011)b 0.000 (0.015)b 0.048 (0.006)a 0.29 7.89 <0.0001 Phyllostylon rhamnoides 0.007 (0.025) 0.000 (0.035) 0.122 (0.014) 2.203 11.34 <0.0001 Pterogyne nitens 0.000 (0.004) 0.006 (0.005) 0.005 (0.002) 0.003 0.91 0.41 Sweetia fruticosa 0.015 (0.009) 0.011 (0.012) 0.031 (0.005) 0.05 2.24 0.11 Overall Species 1.543 (0.180)b 1.302 (0.245)b 2.019 (0.098)a 52.87 5.44 0.005

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62Table 2-4. Establishment and mortality rates of seedlings of commercial tree species monitored over a 3 y period in a control plot and plots subjected to two harvesting intens ities (N=144 subplots/treatment plot). Scientific Name Mortality Rate (% of Seedlings / y) Number of New Seedlings (#/m2 / y) Undisturbed Normal Logging Intensive Management Overall Undisturbed Normal Logging Intensive Management Overall Acosmium cardenasii 17.0 11.7 9.8 12.8 0.489 0.176 0.101 0.255 Anadenanthera macrocarpa 21.6 5.4 18.8 15.2 0.106 0.079 0.003 0.062 Aspidosperma rigidum 14.5 2.5 5.5 7.5 0.007 0.013 0.037 0.019 Caesalpinia pluviosa 35.8 23.2 18.8 25.9 0.117 0.064 0.011 0.064 Ceiba samauma 0.0 0.0 0.0 0.0 0.001 0.001 0.001 0.001 Centrolobium microchaete 0.0 10.7 43.8 18.2 0.000 0.006 0.000 0.002 Chorisia speciosa 0.0 0.0 0.0 0.0 0.004 0.004 0.000 0.003 Copaifera chodatiana 24.1 8.2 14.3 15.5 0.125 0.015 0.050 0.063 Gallesia integrifolia 0.0 0.0 0.0 0.0 0.034 0.000 0.000 0.011 Machaerium cf. acutifolium 16.1 12.4 12.6 13.7 0.003 0.138 0.106 0.082 Machaerium scleroxylon 34.7 30.8 0.0 21.8 0.157 0.012 0.001 0.056 Phyllostylon rhamnoides 19.5 10.0 0.0 9.8 0.381 0.004 0.011 0.132 Pterogyne nitens 43.8 0.0 0.0 14.6 0.000 0.001 0.022 0.008 Sweetia fruticosa 41.4 24.8 8.3 24.8 0.020 0.010 0.004 0.011 Overall 22.0 12.4 11.2 15.2 1.446 0.523 0.354 0.775

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63 Table 2-5. Results of repeated measure analys is of variance for a sp lit plot design run for seedling density for all species combined or six timber tree species analyzed separately. Double asterisks indicate between subject factors while the unmarked variables are within subject factors. Source Type III Sum of Squares Degrees of Freedom Mean Square F P-value All species Time (Ti) 44.01 4 11.00 25.88 <0.0001 Bromeliad (Br)(**) 163.20 1 163.20 9.99 0.005 Time*Bromeliad (Br) 4.95 4 1.24 2.90 0.03 Irrigation (Ir) 27.83 1 27.83 2.74 0.11 Irrigation*Bromeliad 7.16 1 7.16 0.70 0.41 Exclosure (Ex) 79.66 1 79.66 13.01 0.002 Ex*Br 18.71 1 18.71 3.06 0.09 Ti*Ir 13.99 4 3.49 10.81 <0.0001 Ti*Ir*Br 1.92 4 0.48 1.48 0.22 Ti*Ex 7.75 4 1.94 5.76 <0.0001 Ti*Ex*Br 1.43 4 0.36 1.06 0.38 Ir*Ex 11.73 1 11.73 1.31 0.27 Ir*Ex*Br 16.60 1 16.60 1.86 0.19 Ti*Ir*Ex 0.58 4 0.14 0.39 0.81 Ti*Ir*Ex*Br 0.35 4 0.09 0.24 0.92 Amburana cearensis Time (Ti) 1.74 4 0.44 7.72 <0.0001 Bromeliad (Br)(**) 0.83 1 0.83 0.43 0.53 Time*Bromeliad (Br) 0.13 4 0.03 0.58 0.68 Irrigation (Ir) 3.01 1 3.01 1.87 0.20 Irrigation*Bromeliad 1.09 1 1.09 0.68 0.43 Exclosure (Ex) 4.50 1 4.50 2.59 0.14 Ex*Br 2.52 1 2.52 1.45 0.26 Ti*Ir 1.35 4 0.34 6.03 0.001 Ti*Ir*Br 0.08 4 0.02 0.36 0.84 Ti*Ex 0.55 4 0.14 2.39 0.07 Ti*Ex*Br 0.22 4 0.05 0.95 0.44 Ir*Ex 8.87 1 8.87 3.10 0.10 Ir*Ex*Br 0.09 1 0.09 0.03 0.86 Ti*Ir*Ex 1.03 4 0.26 3.65 0.01 Ti*Ir*Ex*Br 0.08 4 0.02 0.29 0.88 Aspidosperma rigidum Time (Ti) 1.16 4 0.29 8.59 <0.0001 Bromeliad (Br)(**) 0.59 1 0.59 0.25 0.63 Time*Bromeliad (Br) 0.04 4 0.009 0.28 0.89 Irrigation (Ir) 5.37 1 5.37 1.78 0.23 Irrigation*Bromeliad 1.31 1 1.31 0.43 0.53 Exclosure (Ex) 0.19 1 0.19 0.10 0.76 Ex*Br 2.6 1 2.6 1.38 0.28

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64 Source Type III Sum of Squares Degrees of Freedom Mean Square F P-value Ti*Ir 0.32 4 0.08 1.95 0.13 Ti*Ir*Br 0.08 4 0.02 0.47 0.76 Ti*Ex 0.12 4 0.03 1.18 0.34 Ti*Ex*Br 0.13 4 0.03 1.24 0.32 Ir*Ex 1.50 1 1.50 0.87 0.39 Ir*Ex*Br 1.15 1 1.15 0.67 0.44 Ti*Ir*Ex 0.08 4 0.02 0.76 0.56 Ti*Ir*Ex*Br 0.08 4 0.02 0.76 0.56 Ceiba samauma Time (Ti) 0.53 4 0.13 2.02 0.12 Bromeliad(Br)(**) 7.75 1 7.75 4.65 0.07 Time*Bromeliad (Br) 0.05 4 0.01 0.21 0.93 Irrigation (Ir) 1.00 1 1.00 0.72 0.43 Irrigation*Bromeliad 0.53 1 0.53 0.38 0.56 Exclosure (Ex) 0.001 1 0.001 0.004 0.95 Ex*Br 0.06 1 0.06 0.54 0.49 Ti*Ir 0.30 4 0.08 1.06 0.39 Ti*Ir*Br 0.03 4 0.007 0.10 0.98 Ti*Ex 0.12 4 0.03 0.91 0.47 Ti*Ex*Br 0.05 4 0.01 0.42 0.79 Ir*Ex 1.69 1 1.69 9.28 0.023 Ir*Ex*Br 0.79 1 0.79 4.35 0.082 Ti*Ir*Ex 0.13 4 0.03 0.94 0.46 Ti*Ir*Ex*Br 0.09 4 0.02 0.67 0.62 Copaifera chodatiana Time (Ti) 4.1 4 1.03 15.09 <0.0001 Bromeliad (Br)(**) 2.96 1 2.96 1.49 0.24 Time*Bromeliad (Br) 0.16 4 0.04 0.60 0.66 Irrigation (Ir) 1.88 1 1.88 3.64 0.07 Irrigation*Bromeliad 0.06 1 0.06 0.12 0.73 Exclosure (Ex) 3.53 1 3.53 3.18 0.09 Ex*Br 0.03 1 0.03 0.03 0.87 Ti*Ir 0.73 4 0.18 3.59 0.01 Ti*Ir*Br 0.41 4 0.10 2.03 0.10 Ti*Ex 0.53 4 0.13 2.25 0.07 Ti*Ex*Br 0.13 4 0.03 0.56 0.69 Ir*Ex 0.56 1 0.56 0.76 0.39 Ir*Ex*Br 5.13 1 5.13 6.96 0.02 Ti*Ir*Ex 0.12 4 0.03 0.59 0.67 Ti*Ir*Ex*Br 0.31 4 0.08 1.46 0.22 Hymenaea courbaril Time (Ti) 1.85 4 0.46 16.09 <0.0001 Bromeliad (Br)(**) 1.63 1 1.63 3.16 0.09 Time*Bromeliad (Br) 0.09 4 0.02 0.83 0.51 Table 2-5. Continued.

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65 Source Type III Sum of Squares Degrees of Freedom Mean Square F P-value Irrigation (Ir) 0.37 1 0.37 1.33 0.27 Irrigation*Bromeliad 1.13 1 1.13 0.47 0.51 Exclosure (Ex) 5.28 1 5.28 13.23 0.003 Ex*Br 0.26 1 0.26 0.65 0.44 Ti*Ir 0.30 4 0.07 1.96 0.11 Ti*Ir*Br 0.14 4 0.03 0.90 0.47 Ti*Ex 1.4 4 0.35 8.53 <0.0001 Ti*Ex*Br 0.08 4 0.02 0.49 0.74 Ir*Ex 0.08 1 0.08 0.45 0.52 Ir*Ex*Br 0.28 1 0.28 1.58 0.23 Ti*Ir*Ex 0.42 4 0.10 2.95 0.03 Ti*Ir*Ex*Br 0.18 4 0.05 1.29 0.28 Pterogyne nitens Time (Ti) 1.09 4 0.27 6.68 <0.0001 Bromeliad (Br)(**) 2.12 1 2.12 7.74 0.02 Time*Bromeliad (Br) 0.39 4 0.09 2.36 0.07 Irrigation (Ir) 0.26 1 0.26 1.40 0.25 Irrigation*Bromeliad 0.71 1 0.71 4.03 0.07 Exclosure (Ex) 0.05 1 0.05 0.21 0.65 Ex*Br 0.18 1 0.18 0.75 0.41 Ti*Ir 0.62 4 0.16 4.65 0.003 Ti*Ir*Br 0.08 4 0.02 0.63 0.64 Ti*Ex 0.09 4 0.02 0.62 0.65 Ti*Ex*Br 0.14 4 0.03 0.98 0.43 Ir*Ex 0.13 1 0.13 0.71 0.42 Ir*Ex*Br 0.01 1 0.01 0.06 0.81 Ti*Ir*Ex 0.11 4 0.02 0.45 0.77 Ti*Ir*Ex*Br 0.06 4 0.02 0.45 0.51 Table 2-5. Continued.

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66 Table 2-6. Mean relative height growth rates ( 1SE) of seedlings of commercial tree species in response to bromeliad cover removal eval uated in 4 times. Significant differences between treatments were evaluated using t-tests at 95% of confidence level with sequential Bonferroni corrections (PB = 0.0125). Species Date BromeliadNo Bromeliad NMean (SE)NMean (SE) t-testP Amburana cearensis Jan06 110.032 (0.019)18 -0.025 (0.026) 1.570.13 March 06 100.046 (0.029)220.063 (0.021) 0.430.67 May 06 8 -0.022 (0.012)21 -0.015 (0.010) 0.370.72 Dec 06 60.026 (0.009)170.024 (0.008) 0.140.89 Aspidosperma rigidum Jan06 8 -0.009 (0.021)190.020 (0.019) 0.890.38 March 06 9 -0.0004 (0.018)210.096 (0.027) 2.220.03 May 06 90.0145 (0.02)21 -0.003 (0.004) 1.570.13 Dec 06 80.027 (0.006)190.014 (0.003) 1.840.08 Ceiba samauma Jan06 20.170 (0.026)120.080 (0.035) 1.020.33 March 06 30.090 (0.046)150.103 (0.028) 0.200.84 May 06 4 -0.038 (0.015)16 -0.022 (0.008) 0.930.37 Dec 06 30.022 (0.008)7 -0.010 (0.011) 1.650.14 Copaifera chodatiana Jan06 200.038 (0.015)340.029 (0.015) 0.360.72 March 06 250.033 (0.010)390.056 (0.017) 0.990.33 May 06 18 -0.014 (0.009)380.005 (0.007) 1.440.15 Dec 06 80.009 (0.007)260.018 (0.006) 0.760.45 Hymenaea courbaril Jan06 10.34657480.141 (0.101) 0.680.52 March 06 90.026 (0.046)170.028 (0.023) 0.050.96 May 06 60.009 (0.016)18 -0.012 (0.019) 0.620.54 Dec 06 0-20.016 (0.010) -Pterogyne nitens Jan06 20.084 (0.007)90.109 (0.058) 0.200.85 March 06 20.230 (0.078)160.107 (0.027) 1.490.15 May 06 30.024 (0.024)130.032 (0.014) 0.530.81 Dec 06 3 -0.009 (0.013)6 -0.036 (0.014) 0.530.25

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67 Table 2-7. Mean relative height growth rates ( 1SE) of seedlings of commercial tree species in irrigated and droughted plots evaluated 4 times. Significant differences between treatments were determined using t-test at 95% of confidence level with sequential Bonferroni corrections (PB = 0.0125). Species Date Irrigated Droughted NMean (SE)NMean (SE) t-testP Amburana cearensis Jan06 25 -0.006 (0.021)40.015 (0.035) 0.400.69 March 06 210.044 (0.018)110.084 (0.035) 1.110.28 May 06 19 -0.024 (0.012)10 -0.002 (0.005) 1.340.19 Dec 06 150.025 (0.008)80.024 (0.013) 0.100.92 Aspidosperma rigidum Jan06 7 -0.006 (0.066)200.017 (0.081) 0.660.51 March 06 70.152 (0.072)230.041 (0.014) 2.390.02 May 06 70.004 (0.017)230.002 (0.004) 0.190.85 Dec 06 60.024 (0.008)210.016 (0.003) 0.930.36 Ceiba samauma Jan06 130.074 (0.026)10.346 2.820.01 March 06 110.086 (0.033)70.125 (0.037) 0.760.46 May 06 14 -0.029 (0.009)6 -0.017 (0.011) 0.750.46 Dec 06 6 -0.013 (0.009)40.019 (0.015) 1.890.09 Copaifera chodatiana Jan06 350.039 (0.011)190.022 (0.024) 0.720.47 March 06 370.040 (0.013)270.056 (0.021) 0.660.51 May 06 310.001 (0.008)25 -0.003 (0.009) 0.300.76 Dec 06 180.008 (0.006)160.024 (0.007) 1.760.09 Hymenaea courbaril Jan06 90.164 (0.092)--March 06 150.020 (0.023)110.037 (0.042) 0.380.71 May 06 12 -0.001 (0.011)12 -0.012 (0.029) 0.110.74 Dec 06 10.02610.005 -Pterogyne nitens Jan06 100.107 (0.052)10.077 0.180.86 March 06 100.085 (0.036)80.165 (0.037) 1.530.15 May 06 80.025 (0.021)80.035 (0.014) 0.400.70 Dec 06 3 -0.036 (0.029)6 -0.023 (0.009) 0.530.61

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68 Table 2-8. Mean relative height growth rates ( 1SE) of seedlings of commercial tree species in response to mammal exclosure evaluated 4 times during 2006. Significant differences between treatments were evaluated using t-test at 95% of confidence level with sequential Bonferroni corrections (PB = 0.0125). Species Date Exclosure No Exclosure NMean (SE)NMean (SE) t-testP Amburana cearensis Jan06 22 -0.011 (0.023)70.021 (0.019) 0.76 0.45 March 06 230.059 (0.018)90.052 (0.041) 0.20 0.84 May 06 21 -0.013 (0.007)8 -0.026 (0.022) 0.72 0.48 Dec 06 180.018 (0.007)50.047 (0.012) 1.79 0.09 Aspidosperma rigidum Jan06 170.015 (0.018)100.004 (0.027) 0.35 0.73 March 06 170.075 (0.034)130.056 (0.021) 0.43 0.67 May 06 17 -0.002 (0.006)130.008 (0.008) 1.05 0.30 Dec 06 170.020 (0.004)100.014 (0.006) 0.99 0.33 Ceiba samauma Jan06 70.094 (0.039)70.092 (0.051) 0.04 0.97 March 06 80.141 (0.032)100.069 (0.034) 1.52 0.15 May 06 9 -0.018 (0.007)11 -0.031 (0.011) 0.88 0.39 Dec 06 30.002 (0.013)7 -0.001 (0.013) 0.17 0.87 Copaifera chodatiana Jan06 310.022 (0.017)230.047 (0.011) 1.13 0.26 March 06 390.058 (0.012)250.029 (0.022) 1.28 0.21 May 06 350.002 (0.007)21 -0.006 (0.011) 0.69 0.49 Dec 06 210.014 (0.006)130.018 (0.007) 0.39 0.70 Hymenaea courbaril Jan06 80.178 (0.103)10.053 0.40 0.70 March 06 240.028 (0.023)20.026 (0.026) 0.02 0.98 May 06 21 -0.005 (0.017)3 -0.013 (0.025) 0.16 0.88 Dec 06 10.00510.026 -Pterogyne nitens Jan06 70.142 (0.060)40.039 (0.073) 1.05 0.32 March 06 90.082 (0.032)90.159 (0.041) 1.51 0.15 May 06 80.028 (0.017)80.032 (0.018) 0.14 0.89 Dec 06 4 -0.025 (0.017)5 -0.029 (0.015) 0.15 0.89

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69 Months AugSepOctNovDecJanFebMarAprMayJunJul Rainfall (mm/month) Soil Moisture Tension (centibars) -300 -200 -100 0 100 200 300 Rainfall 2005-2006 Water added Soil moisture droughted Soil moisture irrigated Rainfall Mean 30 years (A) (B) Figure 2-1. Monthly rainfall (A) and soil mo isture tension measured by Watermark soil sensors (B). Also shown in A is the water a dded to the irrigated soil plots. Vertical lines indicate 1 standard (N=4).

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70 Canopy Openness (%) 0-1010-2020-3030-4040-5050-60> 60 Frequency 0 20 40 60 80 100 Control Normal Logging Intensive Management Figure 2-2. Canopy openness in an unlogged control plot, an area subjected to normal timber harvesting (4.7 m3/ha harvested), and an area subject ed to intensive harvesting (8.2 m3/ha) 8 months (left-hand bar) and 42 mont hs (right hand bars) after logging. Canopy openness measures were made with a sp herical densitometer at the end of the rainy season at 1 m above ground at 144 equa lly spaced points in each 10 ha plots.

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71 Block 1 Block 2 Block 3 Block 10 BromeliadsNo Bromeliads Bromeliads Bromeliads Bromeliads No Bromeliads No Bromeliads No Bromeliads R1 R1 R1 R1 R1 R1 R1 R1R2 R2 R2 R2 R2 R2 R2 R2 " R = Replications NW-NE WA-NE WA-EX NW-EX SP1SP2SP3SP4WA = WATER NW = NO WATER EX = EXCLOSURE NE = NO EXCLOSURE SP = Species SP5SP6 Figure 2-3. Design of the experiment on the e ffects of ground bromelia ds, irrigation, extended drought, and seed and seedling predator exclosures on seedling establishment.

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72 Time (Years) 0.0 0.5 1.0 1.5 2.0 Number of Seedlings (#/m 2 ) 0.0 1.0 2.0 3.0 4.0 5.0 Undisturbed Normal Harvesting Intensive Management a b b a b c (A) All Species (B) Without Acosmium 2003 2004 2005 Figure 2-4. Seedling densities of 11 tim ber species (A) and 10 species without Acosmium cardenasii (B), the most dominant species, in a control plot, a plot subjected to normal timber harvesting, and a plot subjec ted to intensive timber stand management. The forestry treatments were carried out in 2001, 19 mo prior to the first census. Data are from 4 m2 subplots distributed regularly in a grid with 25 m spacing through each 10 ha treatment plot. Vertical lines indicate standard errors Different letters indicate different significance between treatment s using pairwise LSD tests at 95% confidence.

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73 Time (years) 20042005 Seedling Recruitment (#/m 2 /yr) 0.0 0.2 0.4 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 Mortality Rate (% of seedlings/yr) 0 5 10 15 20 25 30 35 Control Normal Harvesting Intensive Harvesting Figure 2-5. Temporal change s in seedling recruitment (#/m2/y) and mortality for commercial tree species in a Chiquitano dry forest in Bolivia.

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74 0 1 2 3 4 Bromeliad No Bromeliad Seedling Density (# / m2) 0 1 2 3 4 Droughted Irrigated Time 0 1 2 3 4 Exclosure No Exclosure Sep5 Nov5 Jan6 Mar6May6(A) (B) (C)Dec6 Figure 2-6. Seedling density over time in respons e to (A) bromeliad cover, (B) irrigation or drought, and (C) mammalian seed predators. Arrow indicates the time when I stopped the irrigation and shielding plants from rainfa ll. Vertical lines show standard errors of the means (N=40).

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75 Amburana cearensis (7) 0.0 0.2 0.4 0.6 0.8 Bromeliad No Bromeliad Pterogyne nitens (8) 0.0 0.2 0.4 0.6 0.8 Aspidosperma rigidum (10) 0.0 0.2 0.4 0.6 0.8 Ceiba samauma (8) Seedling Density (# / m2) 0.0 0.2 0.4 0.6 0.8 Copaifera chod atiana (8) 0.0 0.2 0.4 0.6 0.8 Hymenaea courbaril (3) 0.0 0.2 0.4 0.6 0.8 Sep5 Nov5 Jan6 Mar6 May6 Dec6 Sep5 Nov5 Jan6 Mar6 May6 Dec6 Figure 2-7. Mean densities ( 1SE) of seedlings of commercial timber tree species in 2 m2 plots (N=40) with bromeliads (filled dots) a nd without bromeliads (open dots). The number of seeds sown in each 1 m2 plot is indicated af ter each species name.

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76 Amburana cearensis 0.0 0.2 0.4 0.6 0.8 1.0 Irrigation No irrigation Aspidosperma rigidum 0.0 0.2 0.4 0.6 0.8 1.0 Ceiba samauma Seedling Density (# / m2) 0.0 0.2 0.4 0.6 0.8 1.0 Copaifera chodatiana 0.0 0.2 0.4 0.6 0.8 1.0 Hymenaea courbaril 0.0 0.2 0.4 0.6 0.8 1.0 Pterogyne nitens 0.0 0.2 0.4 0.6 0.8 1.0 Sep5Nov5 Jan6 Mar6 May6 Dec6 Sep5Nov5 Jan6 Mar6 May6 Dec6 Figure 2-8. Mean of seedling densities ( 1SE) in irrigated and droughted experimental plots. Note differences in y-axis scales. Arro ws indicate the time when I stopped the irrigation and shielding plants from rainfall.

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77 Amburana cearensis 0.0 0.2 0.4 0.6 0.8 1.0 Aspidosperma rigidum 0.0 0.2 0.4 0.6 0.8 1.0 Exclosure No exclosure Ceiba samauma Seedling Density (# / m2) 0.0 0.2 0.4 0.6 0.8 1.0 Copaifera chodatiana 0.0 0.2 0.4 0.6 0.8 1.0 Hymenaea courbaril 0.0 0.2 0.4 0.6 0.8 1.0 Pterogyne nitens 0.0 0.2 0.4 0.6 0.8 1.0 Sep5 Nov5 Jan6 Mar6 May6Dec6 Sep5 Nov5 Jan6 Mar6 May6Dec6 Figure 2-9. Mean seedling densit ies (1 SE) in control plots (c losed dots) and in plots from which mammals were excluded (open dots).

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78 Jan6 Mar6 May6 (A) 0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 Droughted Irrigated (B) Ln Relative Height Gr owth Rate (cm/month) 0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 No Exclosure Exclosure (C) Time 0.00 0.02 0.04 0.06 0.08 0.10 0.12 0.14 No Bromeliad Bromeliad [99] (45) [101] (87) [91] (84) [49] (54) [92] (52) [120] (68) [111] (64) [64] (41) [44] (100) [58] (130) [48] (127) [28] (77)Dec6 Figure 2-10. Mean relative growth height growth rates ( 1SE) of seedlings of commercial tree species in response to three experimental treatments: (A) irriga tion or drought; (B) mammalian seed predator exclusion; and, (C) bromeliad cover. Numbers indicate the number of seedlings used for each evaluati on; numbers in brackets are the seedlings in irrigated, exclosure, or bromeliad-cove r plots while numbers in parenthesis are for droughted, non-exclosure, and no-bromelia d plots. There were no significant differences between treatments for each time calculated with t-test at the 95% confidence level with seque ntial Bonferroni correctio ns (repeated-measures ANOVA was not suitable because sample sizes decl ined over the study period due to seedling mortality).

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79 CHAPTER 3 CONTRIBUTION OF ROOT A ND STUMP SPROUTS TO NATURAL REGENERATION IN A LOGGED TROPICAL DRY FOREST IN BOLIVIA Introduction Securing sufficient natural regeneration of co mmercial tree species after logging is critical for sustainable forest management. Most stud ies of tropical forest regeneration focus on tree recruitment from seeds and, consequently, rege neration is often viewed as depending on seed production, seed dispersal, seed viability, and the environmental requirements for seed germination and seedling establishment (Hol l 1999, Dalling and Hubbell 2002, De Steven and Wright 2002). In tropical dry fo rests, many tree species produc e abundant and well-dispersed seeds with high viability, but due to seed pred ation (Janzen 1971), wate r stress (Gerhardt 1994), and a multitude of other factors, successful recrui tment from seed is rare for many species in these forests (Mostacedo and Fredericksen 1999). Fu rthermore, in forests in general and in dry forests in particular, tree seedlings that do b ecome established often gr ow more slowly than sprouts (Miller and Kauffman 1998, Khurana and Singh 2001). Given the limitations on seed dispersal and germination, as well as on seedling establishment and survival, succe ssful dry forest regeneration af ter logging, severe windstorms, or fires may depend greatly on contributions from stump and root sprouts (Kruger et al. 1997, Kammesheidt 1998, Miller and Kauffman 1998, Gould et al. 2002, Homma et al. 2003). The general capacity of dry forest tree species to sprout may be an adaptive response to a history of fire (e.g., Bond and Midgley 1995) or other dist urbances. Whatever the ultimate cause, in a variety of seasonal tropical forests, logging repo rtedly stimulates abundant stump sprouting of felled and broken trees, and root sprouting from superficial roots damaged by heavy equipment (Kauffman 1991, Kammesheidt 1998, Miller and Kauffman 1998, Kammesheidt 1999, Bell

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80 2001). In the Chiquitano dry forest in Bolivia although sprouting has been reported few times following logging (Fredericksen et al. 2000) and fires (Gould et al. 2002, Kennard and Putz 2005), little is known about the overall contributions of sprout-origin plants to forest recovery. Whereas interspecific comparisons of sprou ting ability are numerous for Mediterranean ecosystems (Bellingham 2000, Pausas 2001), how this ability varies with light requirements and other ecological attributes is le ss clear for tropical dry forest speci es (but see Paciorek et al. 2000 for resprouting across the spectrum of shade tolerance of trees on Barro Colorado Island). Sprouting is of interest to forest managers a nd ecologists because sprout s often have more rapid grow rates than true seedlings (Danie l et al. 1979, Clark and Hallgren 2003). The purpose of this study was to examine the contribution of sprouts to the natural regeneration of a tropical dry forest following lo gging. More specifically, I (1) characterized stump and root sprouting features of the comm ercial canopy tree species. I (2) quantified the effect of logging on relative abundances and growth rates of stump sprouts, root sprouts, and true seedlings. I (3) related the species-specific prob abilities of stump sprouting as a function of stump diameter and stump height; and I (4) expl ored how sprouting varies with the ecological requirements of canopy tree species. Methods Study Area This study was conducted on the property of INPA Parket (hereafter INPA), a 30,000-ha tract of privately-owned seasonall y dry tropical forest 30 km NE of the town of Concepcin (16 6' 45"S, 61 42' 47"), 250 km northeast of the city of Santa Cruz de la Sierra, Bolivia (Figure 11). The study area is flat to gent ly sloping, with an altitude of approximately 380 m, mean annual temperature of 24.3 C, and mean annual precip itation of 1100 mm. During the 5-mo dry season (May-October), most trees are deciduous and ma ny tree species flower an d fruit following rain

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81 events in the midto late-dry season. The forest canopy is 20-25 m tall with common species including Acosmium cardenasii Tabebuia impetiginosa Anadenanthera macrocarpa Astronium urundeuva and Centrolobium microchaete (Pinard et al. 1999); after first mention, species will be referred to by their generic na mes (for full names see Table 3-1). Currently, 21 tree species, including those mentioned above, ar e harvested for timber processed mostly into parquet flooring. During the rainy season, canopy openness, as measured 1 m above the ground with a spherical densiometer, was 8% and 14% in contro l and logged areas, resp ectively. During the dry season, canopy openness triples because many tree species are deciduous. The understory is dense, partially due to the abunda nce of lianas, and typically 30 -40% of the ground is covered by the bromeliad, Pseudananas sagenarius Experimental Design and Data Collection Stump sprouts I sampled sprouts from the stumps of harvested trees in three areas that varied in time since logging. The first area (50 ha) was se lectively logged by INPA Parket ( 4 trees/ha and 4 m3/ha, and 10-12 species harvested) 1 y before I be gan my study. Here I mapped, marked, and measured the diameters and heights of the stum ps and all stump sprouts of the five most commonly harvested tress ( Anadenanthera Centrolobium Copaifera chodatiana Tabebuia and Zeyheria tuberculosa ; tree densities reported in Table 2-1) and monitored sprout survival and height growth for one year. The second study area (40 ha) is located in the 20-ha permanent plots maintained by the Instituto Boliviano de Investigacin Forestal (IBIF) for monitoring forest dynamics after logging. Two years prior to my study, 4-8 trees/ha (5.3-6.4 m3/ha, 14 species) were logged from these plots. I monitored spro uting as described above The third area (30 ha) was logged (2-3 trees/ha, 3 m3/ha, and 5-7 species harvested) 5 y before my study. I measured

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82 the frequency and height of stum p sprouts of six tree species ( Anadenanthera Caesalpinia Centrolobium Copaifera M. scleroxylon and Tabebuia ) in this area, using a 7 m telescoping measuring rod. In 2003 I checked for sprouts on the stumps of trees harvested in 1998 (6 species), 2001(10 species), and 2002 (6 species). The 498 stum ps evaluated in the three areas were from trees 40 cm DBH that were felled with chai nsaws 10-90 cm height above ground. Each species was placed in one of the following four ecological guilds based on field observations and the literature (Whitmore 1998, Mo stacedo and Fredericksen 1999, Poorter et al. 2006): lightdemanding pioneers have light requirements and are short-lived; long-lived pioneers also are light-demanding but are longer lived; somewhat shad e tolerant species establish in the shade but only mature under moderate to high light intensitie s; and, shade-tolerant species can establish and survive in the shade. I count ed all sprouts and measured the heights of the two tallest on each stump (from the point of origin) as well as the he ight and diameter of each stump dating from the 2001 and 2002 harvests. Comparison of different juvenile types in relation to microsites created by logging In two of IBIF’s 20-ha experi ment plots, I compared the densities and sizes of seedlings and sprouts < 2 m tall in the following microsites created by an episode of selective logging that occurred 1.5 years prior to my study: logging gaps (280-330 m2, N = 16); logging roads (N = 16); log landings (N = 8); primar y skid trails (N = 16); and, se condary skid trails (N = 16). Secondary skid trails were those used to extract a single log, while primary skid trails were those where skidder had extracted 2 logs. In each microsite, all plants < 2 m tall of 16 canopy tree species in 10 x 4 m plots were classified as havi ng developed directly from a germinated seed or sprouted from a root or stem; determination of plant origin often invol ved excavation, but was generally unambiguous.

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83 Data Analysis I used logistic regression to determine the prob ability of a stump sprouting in relation to its diameter and height for each the five species that sprouted frequently ( Anadenanthera, Centrolobium, Copaifera, Tabebuia, and Zeyheria ). Nagelkerke R-square values were used to determine the percentage of variance explained by each regression and a Hosmer and Lemeshow X2 goodness-of-fit tests was used to determine the significance of each relationship (Field 2000). To determine whether there are relationshi ps between stump diameter and height (independent variables) with the number and maximum height s of sprouts (response variables), I used linear regressions or nonlin ear regression analyses based on linear, quadratic, cubic, and inverse models. For each species, the simplest (i.e., fewest parameters) model with a high R2 value was selected in which each parameter had a reasonable biological explanation. Analyses of variance (ANOVAs) followed by Tukey’s post-hoc comparisons were used to compare densities of seedlings and root or stem sprouts among logged microsites. Absolute relative annual height growth rates of stump spr outs were calculated for 10 species based on their height 2 y after logging, and hei ghts after 1, 2, and 5 y after lo gging for 6 species. Stump sprout heights were compared among ecological guilds using ANOVAs and Tukey’s post-hoc tests. For each species and ecological guild, an ANOVA and then Tukey’s post-hoc comparison were used to compare mean growth among origin types (i.e., stump sprout, root sprout, or true seedling). All analyses were carried out with SPSS 12.0 for Windows. Results Sprout Characterization. Stump sprouting was common after logging in th e dry forest studied; 27 of the 31 species monitored at least occasionally reprouted from stumps; 62% did so frequently (Table 3-1). Centrolobium Zeyheria and Tabebuia were the most frequent st ump sprouters (Figure 3-1).

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84 Among the six commercial tree species monitored, the proportion of stumps with living sprouts decreased with time since logging (Figure 3-2). Overall, for stum ps censused 1, 2, and 5 years after logging the proportion of st umps with live sprouts was 55%, 43%, and 38%, respectively, but the rate of stump sprout mortality va ried by species. In pa rticular, 80% of the Caesalpinia stumps and 73% of the Centrolobium stumps had live sprouts 5 y after the trees were felled. Whereas high proportions of Anadenanthera and Copaifera stumps initially sprouted (23 and 13%, respectively), neither specie s had living stump sprouts in the plot logged 5 y prior to my study. Root sprouting was also common after logging in the tropical dry fore st of INPA. Of the 31 tree species monitored (Table 3-1), 16 sprout ed from lateral roots, 7 species at high frequencies. Acosmium cardenasii Centrolobium and Casearia gossypiosperma were the most frequent root sprouters. Most of the 27 species that frequently sprout ed from roots or stumps were shade tolerant (9) or at least partially shade-tolerant (8 ). Light-demanding pioneer species sprouted infrequently, if at all (e.g., Astronium Piptadenia viridifolia Acacia bonariensis, and Schinopsis brasiliensis ). The most frequent sprouters we re the long-lived pioneer species, Centrolobium and Tabebuia and the partial shade-tolerant species, Zeyheria (Table 3-1). Caesalpinia pluviosa had the higher (20.04.4) number of sprouts per stump, followed by Centrolobium (15.41.47) and Zeyheria (14.21.5). The other f our of the seven species monitored had < 5 sprouts/stump (SE=0.5), with the absolute lowest numbers observed in Copaifera and Anadenanthera (Figure 3-3) Considering all sprouted stumps, there was a significant negative linear relationship between the number of sprouts per stump and stump diameter (T able 3-2). In contrast, when

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85 species were considered separa tely, the only species that sh owed a significant (quadratic) relationship between number of sprouts and stump diameter was Copaifera, and that relationship was positive (i.e., opposite from the overall trend). For all species considered together, there was a negative linear relationship between the num ber of sprouts and stump height, but the relationship varied among species. The number of Caesalpinia sprouts increased with stump height whereas in Centrolobium and Tabebuia the relationship was negative (Table 3-2). Sprout height growth, based on measures of th e tallest stump sprouts 1-5 y after the trees were cut, generally decreased with stump diameter, but species varied in this relationship. Only 3 of the 7 species for which I have suffi cient data showed significant trends: Caesalpinia showed a positive cubic relationship; in Machaerium the relationship was inverse positive; and, Copaifera had a quadratic and positive relationship (Table 32). Sprout height growth rates decreased with stump height when all of the species were consider ed together. In contrast, at the species level, only Caesalpinia and Machaerium showed significant relationships between sprout growth rates and stump height, but in the former the relatio nship was negative and qu adratic and the latter, positive and inverse (Table 3-2). The probability of stump sprouting as relate d to stump diameter varied among species (Figure 3-3). In Copaifera Anadenanthera Tabebuia and Centrolobium, the proportions of sprouted stumps were approximately 0.17, 0.24, 0.57, and 0.97, respectively, and did not vary with stump diameter. In Zeyheria sprouting reached 98% of the stumps 38-40 cm diameter but decreased to only 40% among stumps 90 cm in diameter (Figure 3-4). The probability of sprouting in relation to stum p height also varied among species Figure 3-5). The proportions of sprouted stumps of Copaifera Anandenanthera, and Centrolobium did not vary with stump diameter. In contrast, a Tabebuia stump 10 cm tall was almost certain to

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86 sprout (0.98) whereas this probability declined to 0.13 for a 72 cm tall stump. In Zeyheria the probability of sprouting was high (0.91) and did not vary with stump height (Figure 3-5). Juvenile Types and the Effects of Logging In the plots censused 1.5 years after logging, 45% of juveniles < 2 m tall of canopy tree species were root and stem sprouts, not true s eedlings (Figure 3-6). At the species level there was great variation in the proporti ons of true seedlings (Figure 37). All 15 species evaluated were represented by some sprouts and sprouted at least occasionally fr om roots whereas only 9 species were represented by stem sprouts. Thr ee species sprouted predominantly from roots whereas stem sprouting was the predominant mode of regeneration in only one species. Lightdemanding species tended to regenerate more from seeds and root sprouts than from stem sprouts (F=12.10, P=<0.0001), while partia lly shade tolerant and shade to lerant regenerated more from seeds (F=4.46, P=0.01; F=8.01, P=0.0004; respectively). Densities of plants < 2 m tall of canopy tree species did not vary among the logging microsites (F = 1.37, P = 0.24), but microsites di ffered in the relative contributions of true seedlings and sprouts (Table 3-3). True seedli ngs were twice as abunda nt as root and stem sprouts combined in logging gaps (F = 9.91, P = 0.0001). In contrast, there was no difference in plant density by origin in logging roads (F = 0. 38, P = 0.68). Densities of plants from root sprouts and true seedlings were similar in log landings (F = 1.95, P = 0.18). On primary skid trails most plants <2 m tall were true seedlings with fewer root sprouts, and almost no stem sprouts (F = 13.57, P < 0.0001). On secondary sk id trails, true seedlings were much more common than plants of either sprout type (F = 4.6, P = 0.01). Growth of Stump Sprouts Based on measures of the tallest sprout per stump, the growth rates of stump sprouts varied among species by more than an order of magnitude (Figure 3-8). Anadenanthera (197 cm/y),

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87 Centrolobium (195 cm/y) and Zeyheria (185 cm/y) had the highest growth rates, while Aspidosperma (3.5 cm/y) and Copaifera (25 cm/y) had the lowest Stump sprouts of lightdemanding pioneer species grew fast er than those of shade-tolerant species, but there was a great deal of within species variation, especially in the growth rates of the la tter (Table 3-4). Stump sprouts of long-lived pioneer spec ies grew at about the same rate s as shade tolerant species. Among individuals of canopy tree species < 2 m tall, root and stem sprouts both grew faster than seedlings and in 5 of 12 species monito red, root sprouts grew fa ster than stem sprouts (Table 3-5). Centrolobium and Chorisia speciosa had the highest root sp rout growth rates. I found no Anadenanthera Aspidosperma Casearia arborea M. scleroxylon Phyllostylon rhamnoides, or Piptadenia stem sprouts (Table 3-5). Among th e root sprouters, light-demanding and long-lived pioneer species gr ew faster than partially shad e tolerant and shade tolerant species. Among the stem sprouters and true seedli ngs, shade-tolerant species grew slower than belonging to other light-syndrom e classes (Table 3-6). Stump sprout heights varied over time and am ong the six species censused 1, 2, and 5 y after logging (Figure 3-9). Appa rent absolute growth rates (c m/y) increased through the second year and then height increments stopped except in Centrolobium which continued to grow at a rapid rate through the fifth year. Anadenanthera sprouts were appare ntly growing rapidly through the second year, but I co uld find no live stumps in the plot logged 5 y prior to my census. Discussion Of the 31 canopy tree species studied in a dry tropical forest in Bolivia, 27 (87%) have some capacity to sprout from either roots or st umps. Sprouting is apparently characteristic of many tropical dry forest tree species and helps them persist in an environment where stress is severe and disturbances are frequent (Bellingham 2000, Bond and Midgley 2001). As observed

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88 in the USA (Jones and Raynal 1988) and Mexico (Dickinson 1998), root sprouting was promoted by logging damage to roots in my study species. The high proportion of sp ecies that sprouted from broken and cut stems in INPA may be relate d to the high frequency with which stems lose their terminal buds due to herb ivore browsing during th e dry season when other browse is scarce as well as to the direct effect s of drought stress (Bossard and Rejmanek 1994, Del Tredici 2001, Groll 2005). Natural Regeneration and Shade Tol erance: True Seedlings vs. Sprouts Scarcity of natural regeneration from seeds is common for most tree sp ecies in tropical dry forest in Bolivia (Mostacedo and Fredericksen 1999). The main reasons for this scarcity appear to be high seed predation, low seed viability, and high seedling mortalit y during the dry season. Sprouting from broken and cut stems, along with root sprouting, appears to be a very important regeneration mechanisms in tr opical dry forests in Brazil (Castellani and Stubblebine 1993), Jamaica (Bellingham et al. 1994), and Venezuela and Paraguay (Kammesh eidt 1999), including the forest I studied in Bolivia where 45% of the regeneration of canopy trees originated from root or stem sprouts. Sprouting is a common mode of tree regeneration in forest s around the world. For example in oak forests in the USA (Clark and Hallgren 2 003, Nyland et al. 2006) a nd in boreal forest in Russia (Homma et al. 2003), trees reportedly regenera te mainly from sprouts. Sprouting seems to represent the predominant mode of regeneration in forests frequen tly subjected to logging, wind damage, and fire (Bond and Midgley 2001). Natural regeneration by sprouting from latera l roots was common in some commercial species in my study site. In particular, Centrolobium Tabebuia Aspidosperma and M. scleroxylon regenerated mostly from root sprouts. Centrolobium was previously reported as a root sprouting species (Frederickse n et al. 2000), but the importance of this mode of regeneration

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89 in the other species has apparently been overlooked. It remains to be seen whether root sprouts mature into sound trees, and there ar e reasons to suspect that they wi ll not. First of all, given that most new stems emerge from damaged stems or r oots, sprouts of all sorts seem particularly prone to butt and root rots. Second, I observed that several Centrolobium root sprouts that were 5-8 cm DBH 5 y after sprouting still had not de veloped their own root systems. Those that I excavated emerged from large diameter roots running about 5 cm below the soil surface but had developed almost no roots of their own and we re thus mechanically unstable when pushed perpendicular to the orientati on of the source root. Given the general importance of root sprouting after fires, logging, a nd other severe disturbances, such as found in tropical dry forest in Paraguay and moist semi-deciduous forest in Venezuela (Kammesheidt 1999), root sprout longevity is an issue that deserves more attention from researchers. Partially shade-tolerant and shade-tolerant species were more likely to sprout than lightdemanding species. Most of the part ially shade-tolerant species in INPA sprouted from either roots or stems; similar findings were reported for a moist but seasonal tropical forest in Panama (Paciorek et al. 2000). In contrast in a moist tropical forest but after slash-and-burn agriculture of eastern Paraguay, light-demanding species co ntributed more sprouts than shade-tolerant species (Kammesheidt 1998). Although some light -demanding species in INPA did not sprout (13%), others stump sprouted frequen tly, such as species in the Bombacaceae and Flacourtiaceae. Furthermore, the light-demanding pioneer species th at did sprout grew faster than sprouts from other ecological groups. Allometric Relationships with Stump Sprouting The weak and inconsistent trends in the re lationship between either stump diameter or stump height and the growth rates or number of sprouts per stump means that I have little basis on which to make firm recommendations for sprout management. Consistent patterns in sprout

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90 responses are also not appa rent in the literature. Whereas several studies mention that in the first years after tree cutting there is a positive relationship between stum p diameter and sprout growth rates (Jobidon 1997), other studie s report the opposite (Trani et al. 2005), and generally the relationship does not remain signifi cant after a few years. These resu lts suggest that factors other than stump size controls sprouting (McConna ughay et al. 1996, Ma sri et al. 1998). The probability of sprouting varied substantia lly among species but I observed no effect of stump diameter on the probability of sprouting in four of the five species studied. For example, Centrolobium had the highest probability of stump sprouting (97%), while Copaifera had the lowest (17%). In contrast, Zeyheria showed a decreasing probabil ity of stump sprouting with increasing stump diameter, a pattern also observed in a wetter but still seasonal tropical lowland forest in Panama (Putz and Brokaw 1989) and in an oak forest in southern Indiana (Weigel and Peng 2002). The probability of stump sprouting did not vary either with stump height except in Tabebuia in which the probability decreased with stump height. These results suggest that harvesting trees of any tree size will promote same probability of sprouting, except for Zeyheria in which it is better to cut smaller trees and in Tabebuia in which low stumps are preferred if sprouting is to be encouraged. Growth of Stem and Root Sprouts Compared with True Seedlings One advantage of natural regeneration via spro uting is that sprouts typically grow more rapidly than true seedlings, at least initially (Gould et al. 2002, Kennard et al. 2002), which was confirmed by this study. I also observed that sma ll plants of sprout origin typically appeared less affected by drought than true seed lings (Personal Observation). Ne vertheless, in my study forest as well as in Australia (Enright and Goldblum 1999) and South Africa (Kruge r et al. 1997), true seedlings of some species grew just as fast as sprouts. In several sp ecies, especially light-

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91 demanding pioneers (e.g., Cordia Casearia ), height growth rates were similar between true seedlings and root or stem sprouts. Conclusions Due to the high costs and frequent failure s of seed and seedling planting, natural regeneration is critical for th e sustainable management of trop ical dry forest tree species in Bolivia. Given that so many tree species sprout prolifically from stumps of all sizes or from lateral roots, especially after m echanical damage, sprouts need to be considered as a source of regeneration. The abundance of sp routs and their typica lly rapid growth rates, when compared with those of true seedlings, adds to the poten tial value of sprouts for forest management. In forests not designated for timber stand management sprouts deserve at least as much attention from researchers as seeds and true seedlings. That said, future studies should consider the longterm fates of sprouts. The observation in this study that the stump sprouts of most species essentially stopped growing after 2 y needs to be verified and otherwis e explored, as do the factors that cause high rates of mo rtality of the sprouted stumps of some species. In the case of root sprouts, which were also abundant in my study area, long-term monitoring is needed to determine whether they ever grow up to be sound, canopy trees.

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92Table 3-1. Frequency of root and stem sprouting and shade tolerance of commerc ial and non-commercial ca nopy tree species in a tropical dry forest in Bolivia. Shade to lerance is base on Pinard et al. (199 9), and Mostacedo and Fredericksen (1999): L=Light-demanding pioneer, LL=Long-liv ed pioneer, PS=Partially shade to lerant, ST=Shade tolerant. Resprout Type/ Frequency Species Abbreviation Family ROOT STEM Shade Tolerance Commercial Timber Species Amburana cearensis (Allemo) A.C. Sm. AMCE Caesalpinace ae No Yes/Low PS Anadenanthera macrocarpa (Benth.) Brenan ANMA Mimosaceae No Yes/Low L Aspidosperma rigidum Rusby ASRI Apocynaceae Yes/Low Yes/High ST Astronium urundeuva (Allemo) Engl. ASUR Anacardiaceae No No L Caesalpinia pluviosa DC. CAPL Caesalpiniacea e Yes/Low Yes/High PS Cariniana ianeirensis R. Knuth CAIA Lecythidaceae Yes/Low Yes/High LL Cedrela fissilis Vell CEFI Meliaceae Yes/Low Yes/Low L Centrolobium microchaete (Mart. ex Benth.) Lima ex G. P. Lewis CEMI Fabaceae Yes/High Yes/High L Copaifera chodatiana Hassl. COCH Caesalpiniaceae Yes/Low Yes/High ST Cordia alliodora (Ruiz & Pav.) Oken COAL Boraginaceae Yes/Low Yes/Low L Hymenaea courbaril L. HYCO Caesalpiniaceae Yes/Low Yes/Low PS Machaerium scleroxylon Tul. MASC Fabaceae Yes/Low Yes/High PS Phyllostylon rhamnoides (J. Poiss.) Taub. PHRH Rhamnaceae No Yes/High ST Platymiscium ulei Harms PLUL Fabaceae No Yes/High L Schinopsis brasiliensis Engl. SCBR Anacardiaceae No No L Sweetia fruticosa Spreng. SWFR Fabaceae No Yes/Low PS Tabebuia impetiginosa (Mart. ex DC.) Standl. TAIM Bignoniaceae Yes/Low Yes/High LL Tabebuia serratifolia (Vahl) G. Nicholson TASE Bignoniaceae Yes/High Yes/Low LL Zeyheria tuberculosa (Vell.) Bureau ZETU Bignoniaceae No Yes/High PS Non-commercial Species Acacia bonariensis Gillies ex Hook. & Arn. ACBO Mimosaceae No No L Acosmium cardenasii H.S. Irwin & Arroyo ACCA Fabaceae Yes/High Yes/High ST Aspidosperma cylindrocarpon Mll. Arg. ASCY Apocynaceae Yes/High Yes/High LL Capparis prisca J.F. Macbr. CAPR Capparaceae No Yes/High ST Casearia gossypiosperma Briq. CAGO Flacourtiaceae Yes/High Yes/High L Ceiba samauma (Mart.) K. Schum. CESA Bombacaceae No Yes/Low L Chorisia speciosa A. St.-Hil. CHSP Bombacaceae Yes/High Yes/High L Eriotheca roseorum (Cuatrec.) A. Robyns ERRO Bombacaceae No Yes/Low L Gallesia integrifolia (Spreng.) Harms GAIN Phytolacaceae No Yes/High LL Machaerium acutifolium Vogel MAAC Fabaceae Yes/High Yes/Low PS Piptadenia viridiflora (Kunth) Benth. PIVI Mimosaceae No No L Spondias mombin L. SPMO Anacardiaceae No Yes/Low PS

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93 Table 3-2. Summary of the best models and their significances from the regres sion analyses between stump diameter or stump volume (independent variables) and the he ights of stump and numbers of sprouts per stump (dependent variables). Tests were made for species that were sample d with > 8 individuals at alpha = 0.05. Models: L=Linear, Q=Quadratic, C=Cubic, I=Inverse. Signs mean positive (+) or negative (-) relationship between two variables. # of sprouts Height of sprouts Scientific Name Model R2 DF F P Model R2 F P Stump Diameter Anadenanthera macrocarpa L 0.015 36 0.56 0.461 L 0.034 1.27 0.2680 Caesalpinia pluviosa Q(+) 0.267 18 3.27 0.061 C(+) 0.406 6.16 0.0090 Centrolobium microchaete L 0.005 90 0.41 0.522 L 0.012 1.09 0.3000 Copaifera chodatiana Q(-) 0.390 21 6.71 0.006 Q(-) 0.536 12.11 <0.0001 Machaerium scleroxylon Q 0.365 5 1.44 0.321 I(+) 0.630 10.24 0.0190 Tabebuia impetiginosa L 0.026 54 1.45 0.234 L 0.010 0.54 0.4660 Zeyheria tuberculosa L 0.001 62 0.05 0.819 I 0.036 2.33 0.1320 All species L(-) 0.020 301 5.61 0.020 L(-) 0.040 3.95 0.0400 Stump Height Anadenanthera macrocarpa L 0.01 36 0.49 0.49 L 0.001 0.03 0.87 Caesalpinia pluviosa L(+) 0.4 19 12.90 0.002 Q(-) 0.32 4.24 0.03 Centrolobium microchaete I(-) 0.36 90 49.96 <0.0001 I 0.02 2.06 0.16 Copaifera chodatiana L 0.03 22 0.66 0.42 L 0.02 0.42 0.52 Machaerium scleroxylon L 0.006 6 0.04 0.85 I(+) 0.62 10.03 0.02 Tabebuia impetiginosa I(-) 0.34 59 29.82 <0.0001 L 0.02 0.90 0.35 Zeyheria tuberculosa I 0.03 63 2.06 0.16 L 0.008 0.53 0.47 All Species L(-) 0.03 311 8.91 0.003 I(-) 0.16 56.6 <0.0001

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94Table 3-3. Mean ( 1SE) densities of tr ue seedlings, stem sprouts, and root sprout s in 10 x 4 m plots in microsites created du ring selective logging. Different letters indicate differences between microsites in the densities of plants of different origins using Tukey post hoc comparisons with 95% of confidence. Microsites SeedlingStem SproutRoot SproutMean of SquareFP Logging Gaps 78.75 (10.47)a21.87 (5.18)b23.44 (8.82)b113419.910.0001 Logging Roads 28.43 (11.45)a8.33 (3.33)a26.09 (6.80)a5130.380.68 Landings 34.58 (15.84)a0.62 (0.62)a24.64 (7.83)a14111.950.18 Primary Skid Trail 50.15 (9.47)a4.21 (1.87)c24.06 (4.89)b849313.570.0001 Secondary Skid Trail 59.84 (11.49)a31.41 (10.62)b19.53 (5.17)b68664.640.01

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95 Table 3-4. Mean ( 1SE) of stump sprout he ight growth rates (cm/ year) by ecological groups. Different letters indicate significant differences between ecological groups using Tukey post hoc comparisons with 95% confidence. Ecological Group N M ean Mean Square F P Light-demanding Pioneer 47 194.7 (17.4)a 57005.7 5.85 0.001 Long-lived Pioneer 6 84.8 (17.8)c Partially Shade Tolerant 24 159.5 (12.3)b Shade Tolerant 4 14.5 (8.8)c

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96 Table 3-5. Means ( 1SE) of stem height s by species that sprouted from stems or r oots compared to the heights of seedlings. Di fferent letters indicate significant differences in plant origins within species using Tukey post hoc comparisons at the 95% confidence level. Species SeedlingStem ResproutRoot SproutMean Square FP Acosmium cardenasii 18.3(2.3)b42.4(4.0)a46.2(3.6)a2862327.350.0001 Anadenanthera macrocarpa 25.1(1.5)b-72.6(10.6)a1104219.60.0001 Aspidosperma rigidum 30.0-55.0(5.68)5681.760.22 Caesalpinia pluviosa 54.7(16.5)100.5(29.8)112.5(29.8)67701.900.18 Casearia arborea 125.0(52.0)-105.0(90.1)3000.0370.86 Casearia gossypiosperma 102.7(10.9)100.0(19.5)78.8(13.8)18440.970.39 Centrolobium microchaete 67.8(26.6)b138.7(39.9)a154.2(4.8)a328505.150.006 Chorisia speciosa 141.2(15.7)-125.0(22.2)3520.200.67 Copaifera chodatiana 31.7(16.6)61.4(12.9)33.0(28.8)9611.150.37 Cordia alliodora 135.0(39.1)155.0(67.6)55.0(67.6)30400.660.60 Machaerium acutifolium 33.6(2.5)b59.2(5.3)a60.7(4.1)a2329121.090.0001 Machaerium scleroxylon --70.0(5.0)--Phyllostylon rhamnoides 14.0(2.27)-15.00.80.030.86 Piptadenia viridifolia 69.0(6.2)b-140.0(19.1)a5464312.510.001 Sweetia fruticosa 58.0(17.2)110.0120.025521.230.36 Tabebuia impetiginosa 44.5(36.8)90.0(52.0)116.9(11.4)100011.850.17

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97 Table 3-6. Mean ( 1SE) heights of trees 2 m tall that were root sprouts, stem sp routs, or seedlings grouped by ecological guild. Different letters indicate si gnificant differences between ecological groups using Tukey post hoc comparisons at the 95% confidence level. Origin types Light-demanding pioneer Long-lived pioneer Partially shade tolerant Shade tolerantMean Square FP Root sprout 149.1 (3.9)a116.9 (10.7)b64.6 (8.2)c46.7 (7.3)c32399766.8< 0.0001 Stem sprout 86.4 (13.0)ab90.0 (34.5)a64.0 (7.3)ab43.7 (5.7)b95934.10.009 Seedling 44.7 (2.0)a44.5 (20.1)a35.9 (2.8)a18.5 (2.9)b3029818.6< 0.0001

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98 Species CEMICAPLMASCZETUTAIMANMACOCH Stump Sprouting (%) 0 20 40 60 80 100 (N=49) (N=18) (N=8) (N=16)(N=38) (N=38) (N=22) Figure 3-1. Proportions of stum ps of commercial timber species that resprouted (number of stumps noted in parenthesis). CEMI = Centrolobium microchaete CAPL = Caesalpinia pluviosa MASC = Machaerium scleroxylon TAIM = Tabebuia impetiginosa ANMA = Anadenanthera macrocarpa COCH = Copaifera chodatiana

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99 Species CentrolobiumCaesalpiniaM. scleroxylonTabebuiaAnadenantheraCopaifera Resprouting Frequency (%) 0 20 40 60 80 100 Year 1 (Site 1) Year 2 (Site 2) Year 5 (Site 3) Figure 3-2. The proportions of stumps with li ve sprouts over time since logging. For complete species names see Table 3-1.

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100 Species CAPLCEMIZETUTAIMMASCANMACOCH # Sprouts / Stump 0 5 10 15 20 25 30 Figure 3-3. Mean ( 1SE) numbers of sprouts/ stump for the most frequent sprouting species. CEMI = Centrolobium microchaete CAPL = Caesalpinia pluviosa MASC = Machaerium scleroxylon TAIM = Tabebuia impetiginosa ANMA = Anadenanthera macrocarpa COCH = Copaifera chodatiana

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101 Overall Species 406080100 0.0 0.2 0.4 0.6 0.8 1.0 Presence (1) /Absence (0) Probability Centrolobium microchaete 406080100 Probability of St ump Resprouting 0.0 0.2 0.4 0.6 0.8 1.0 Copaifera chodatiana 406080100 0.0 0.2 0.4 0.6 0.8 1.0 Tabebuia impetiginosaStump Diameter (cm) 406080100 0.0 0.2 0.4 0.6 0.8 1.0 Zeyheria tuberculosa 406080100 0.0 0.2 0.4 0.6 0.8 1.0 Anadenanthera macrocarpa 406080100 0.0 0.2 0.4 0.6 0.8 1.0 r2 = 0.04, X2 = 6.26, P = 0.62 r2 = 0.02, X2 = 3.44, P = 0.84 r2 = 0.05, X2 = 8.43, P = 0.39 r2 = 0.06, X2 = 11.81, P = 0.16 r2 = 0.005, X2 = 3.17, P = 0.92 r2 = 0.14, X2= 4.49, P = 0.81 Figure 3-4. Probabiliti es of stump sprouting as a function of stump diameter for the most frequently sprouting commercial tree speci es (curves fit by logistic regression).

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102 Overall Species 020406080 0.0 0.2 0.4 0.6 0.8 1.0 Presence (1) / Absence (0) Probability Anadenanthera macrocarpa 020406080 0.0 0.2 0.4 0.6 0.8 1.0 Centrolobium microchaete 020406080 Probability of Stump Resprouting 0.0 0.2 0.4 0.6 0.8 1.0 Copaifera chodatiana 020406080 0.0 0.2 0.4 0.6 0.8 1.0 Tabebuia impetiginosa Stump Height (cm) 020406080 0.0 0.2 0.4 0.6 0.8 1.0 Zeyheria tuberculosa 020406080 0.0 0.2 0.4 0.6 0.8 1.0 r2 = 0.001, X2 = 6.66, P = 0.46 r2 = 0.001, X2 = 6.62, P = 0.46 r2 = 0.01, X2 = 5.98, P = 0.54 r2 = 0.08, X2 = 8.09, P = 0.32 r2 = 0.23, X2 = 8.09, P = 0.32 r2 = 0.07, X2= 8.78, P = 0.12 Figure 3-5. Probabilities of st ump sprouting as a function of st ump height for the commercial tree species that most frequently sprout ed (curves fit by logistic regression).

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103 Juvenile Types SeedlingStem SproutRoot Sprout Stem Density (#/100 m 2 ) 0 10 20 30 40 50 60 A B B Figure 3-6. Mean ( 1SE) densities of juvenile s < 2 m tall of commercial tree species that were true seedlings, stem sprouts, and root sp routs. Different letter s indicate significant differences between origin types determined with Tukey post hoc comparisons with 95% confidence.

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104 Light DemandingSpecies PHRHACCACOCHMASCMAACASRICAPLSWFRTAIMCEMIANMACAGOCHSPPIVICOAL % of Individuals 0 20 40 60 80 100 120 140 Root Sprout Stem Resprout Seedling (-) (+) Figure 3-7. Percentage of juve niles < 2 m of different origins after logging (for each of 15 dry forest tree species) ordered by their light requirements.

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105 Light DemandingSpecies COCHMASCMAACASRISCBRZETUCAPLTAIMCEMIANMA Relative Growth Rate (cm/year) 0 50 100 150 200 250 300 (-) (+)(2)(6) (1) (2) (1) (6) (11) (6)(41)(5) Figure 3-8. Relative growth rate s (mean ( 1SE) of stump spr outs measured over the first two years after logging for commercial tree specie s in a tropical dry forest in Bolivia arranged by light requirements. Abbreviations of species are shown in Table 3-1.

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106 Time (Years) 0123456 Maximum Stem Length (cm) 0 200 400 600 800 1000 Anadenanthera macrocarpa Caesalpinia pluviosa Centrolobium microchaete Copaifera chodatiana Machaerium scleroxylon Tabebuia impetiginosa Figure 3-9. Mean growth rates of stump sp routs through time for the main commercial tree species as based on measurements of the tall est sprouts on different stumps 1, 2, and 5 y after creation.

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107 LIST OF REFERENCES Abrahamson, W. G., and J. N. Layne. 2003. Longterm patterns of acorn production for five oak species in xeric Florida uplands. Ecology 84:2476-2492. Aizen, M. A., and P. Feinsinger. 1994. Forest fragmentation, pollinati on, and plant reproduction in a Chaco dry forest, Argentina. Ecology 75:330-351. Aez, M. 2005. Analisis del comportamiento re productivo de 35 especies arboreas en un bosque seco chiquitano en el departamento de Santa Cruz, Bolivia. Bachelor's thesis. Universidad Autonoma Gabriel Re ne Moreno, Santa Cruz, Bolivia. Appanah, S. 1993. Mass flowering of dipterocarp forests in the aseasonal tropics. Journal of Biosciences 18:457-474. Asquith, N. M., S. J. Wright, and M. J. Cl auss. 1997. Does mammal community composition control recruitment in neotropical fo rests? Evidence from Panama. Ecology 78:941-946. Augspurger, C. K. 1984. Light requirements of neotropical tree seedli ngs: A comparative study of growth and survival. Journal of Ecology 72:777-795. Barik, S. K., R. S. Tripathi, H. N. Pandey, a nd P. Rao. 1996. Tree regeneration in a subtropical humid forest: Effect of cultural distur bance on seed production, dispersal and germination. Journal of Applied Ecology 33:1551-1560. Beaudet, M., and C. Messier. 2002. Variati on in canopy openness and light transmission following selection cutting in northern ha rdwood stands: an assessment based on hemispherical photographs. Agri cultural and Forest Meteorology 110:217-228. Beckage, B., J. S. Clark, B. D. Clinton, a nd B. L. Haines. 2000. A long-term study of tree seedling recruitment in southern Appalachia n forests: the effects of canopy gaps and shrub understories. Canadian Journal of Forest Research-Revue Canadienne de Recherche Forestiere 30:1617-1631. Bell, D. T. 2001. Ecological response syndromes in the flora of southwestern Western Australia: Fire resprouters versus reseeders. Botanical Review 67:417-440. Bellingham, P. J. 2000. Resprouting as a life hi story strategy in woody plant communities. Oikos 89:409-416. Bellingham, P. J., E. V. J. Tanner, and J. R. Healey. 1994. Sprouting of trees in Jamaican montane forests, after a hu rricane. Journal of Ecology 82:747-758. Blate, G. M., D. R. Peart, and M. Leighton. 1998. Post-dispersal predation on isolated seeds: a comparative study of 40 tree species in a Southeast Asian rainforest. Oikos 82:522-538. Bond, W. J., and J. J. Midgley. 2001. Ecology of sprouting in woody plants: the persistence niche. Trends in Ecology & Evolution 16:45-51.

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117 BIOGRAPHICAL SKETCH Bonifacio Mostacedo was born in Sucre, Boliv ia, and spent the first years of his life helping his father on their small family farm For Bonifacio to furt her his studies beyond secondary school required a great deal of sacrifice and hard work for both him and his family. He studied agriculture at the Au tonomous University Gabriel Rene Moreno (UAGRM) in Santa Cruz, Bolivia. While taking classes he became very interested in plant taxonomy and started working, first as a volunteer, at the Noel Kempff Mercado Natura l History Museum. During that period of his life he had the opportunity to work with Robin Foster, from the Field Museum of Chicago and the late Alwin Gentry, from the Missouri Botanical Garden. Both of these worldrenowned scientists and natural hi storians inspired him to keep working in botany and ecology. While associated with the Museum, he had many opportunities to visit remo te parts of his native country and to become personally familiar w ith many of its ecological communities. He graduated from UAGRM as an Engineer in Agriculture in 1995. After his undergraduate training, he received a scholar ship from BOLFOR Project to do his master’s degree at the Autonomous National University of Mexico (U NAM), Mexico, from which he graduated in 1997 with a M.Sc. degree in ecology and envi ronmental sciences. After graduating from UNAM, he worked for three years as Research Co-Coordinator for the BOLFOR Project and was appointed as a guest professor at UAGRM. Th e next stage in Bonifaci o’s educational career began in 2001 when he received a scholarship from BOLFOR to do his Ph.D. in the Department of Botany at the University of Florida. After taking classes in Gainesville he returned to Bolivia to simultaneously conduct his field research and work with the newly formed Instituto Boliviano de Investigacin Forestal (IBIF), first as Resear ch Coordinator and then as Executive Director. Upon graduation, he plans to return to Bolivia and to continue working with IBIF and UAGRM in Santa Cruz.


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Title: Natural Regeneration of Canopy Trees in a Tropical Dry Forest in Bolivia
Physical Description: Mixed Material
Copyright Date: 2008

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NATURAL REGENERATION OF CANOPY TREES IN A TROPICAL DRY FOREST IN
B OLIVIA




















By

BONIFACIO MOSTACEDO


A DISSERTATION PRESENTED TO THE GRADUATE SCHOOL
OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT
OF THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY

UNIVERSITY OF FLORIDA

2007




































O 2007 Bonifacio Mostacedo


































To my parents for raising me with love and making me what I am today. To my wife, who brings
great joy into my life. To my children, for the inspiration and strength they gave me throughout
the long process of completing this dissertation.









ACKNOWLEDGMENTS

So many people contributed to this work that I cannot acknowledge them all by name but I

do want to single out a few. First of all, I would like to thank my mentor and principal supervisor

Francis E. "Jack" Putz for asking me difficult questions and pushing me to think and write

logically. He helped me adjust to life in Gainesville and was a good friend throughout my career

as a graduate student. Particularly while I was trying to finish my dissertation, he provided a

great deal of support and encouragement. I hope that we can continue to work together in the

future .

I also want to thank the members of my advisory committee, Kaoru Kitajima, Karen

Kainer, Emilio Bruna, and Colin Chapman, for always being available to provide me guidance.

For much of the past decade, Todd Fredericksen has been a key person in my life. We

spent a great deal of time together in various forests in Bolivia conducting what I believe to be

exciting research and teaching what I hope were useful field courses. No matter how bad the

conditions, he always had a smile on his face and an amusing comment on his lips. He

encouraged me to start my Ph.D., wrote letters of recommendation on my behalf, and helped me

secure financial support for my studies. He continues to provide critical feedback on my work

but, overall, I am most grateful to him for being a good friend.

Numerous University of Florida faculty and staff contributed substantially to my

dissertation proj ect. In particular, Doug Levey helped me design the predator exclosures and

Stephen Mulkey helped me with the irrigation experiment. Among the statisticians who guided

me through the design and analysis phases of this research, I would like to thank Larry Winner,

Andrew Khuri, and Jorge Cassela. I am also very grateful to Ann Wagner, Pamela Williams, and

Paula Maurer from the Botany office for their help.










My Ph.D. studies were supported to a large extent by a fellowship from USAID through

the BOLFOR Proj ect run by Chemonics Intemnational. Thanks go to John Nitler and Ivo

Kralj evic for helping me to secure this funding. They, and several other people at Chemonics,

helped me to deal with culture shock and to otherwise adjust to life in the USA. Part of the

research was funded by Intemnational Foundation for Science (IFS) who gave financial support to

do experiments about seedling dynamic. The final phase of my dissertation research was

supported by a fellowship from the Gordon and Betty Moore Foundation through the Tropical

Conservation and Development Program at the University of Florida.

I want to thank my officemates and friends in Gainesville including Clea Paz, Geoffrey

Blate, Joseph Veldman, Morgan Vamner, Bil Grauel, Skya Murphy, Ana Eleuterio, Paulo Brando,

Camila Pisano, and Christine Lucas. Skya, Ana, Paulo, and Christine provided me helpful

comments on my dissertation. I also want to recognize Claudia Romero for teaching me various

things that made my life easier, helping me with the dreaded dissertation templates, reviewing

the sprouting chapter, and convincing me that I could finish this dissertation on a time schedule

that seemed unattainable.

The Instituto Boliviano de Investigacion Forestal (IBIF), through its director Marielos

Pefia-Claros, provided me with all the necessary logistical support needed to conduct my

fieldwork. Marielos also provided many helpful comments on my dissertation proposal. Many

other people at IBIF were strong supporters of my work, including members of the

administrative (Emma Nufiez, Laly Dominguez, and Karina Munoz) and technical staffs

(Marisol Toledo, Zulma Villegas, Juan Carlos Licona, Alfredo Alarcon, Carlos Pinto, Claudio

Leatio, Vincent Vroomans, Betty Flores and Mayra Maldonado) as well as Lourens Poorter, a

research associate. From Lourens I received many helpful suggestions about my research.










My Hieldwork would not have been possible without help of my student assistants and

"materos." In particular, Marlene Soriano helped me collect a portion of the Hield data for the

first and second chapters of this dissertation. Armando Villca and Turian Palacios helped me

with the Hield research for the third chapter. Alej andra Calderon, Vanessa Sandoval, Monique

Grol, Janeth Mendieta, Carla Gonzalez, and Joaquin Cordero helped in various ways in and on

the way to the field. I want also to thank many other people for Hieldwork assistance including

Juan Carlos Alvarez, Israel Melgar, Daniel Flores, Hugo Justiniano, Rodolfo and Rafael Rivero,

Daniel Alvarez, Alberto Chacon, Antonio Jimenez, Juan Pessoa, Dona "Negrita" Pessoa, and

Juan Alvarez. I also want to thank Miguel Angel Chavez, the driver from IBIF, who was always

willing to volunteer to travel the long bumpy road to INPA.

The INPA PARKET Company was the gracious host of my Hield research and helped in

many different ways in its execution. The company's owner, Paul Roosenboom, company

manager William Pariona, forester Urbano Choque, and many other people working for this

prestigious and FSC-certified forest-products firm contributed to the success of this proj ect.

Last but certainly not least, I want to thank my wife, Ynes Uslar, and children, Gabriela

Ines and Jose Daniel, for the many sacrifices a family makes when a parent takes on a Ph.D.

proj ect. The many nights I spent writing instead of being with them, the long stints of field work,

and the even longer periods when I was in Gainesville and they were back in Bolivia were hard

on all of us but they bore the strain without complaint. I should also thank Ynes for counting

thousands of seeds and typing in even more data, but these contributions pale in comparison with

the rest of what she has done for me and the rest of our family.












TABLE OF CONTENTS


page

ACKNOWLEDGMENTS .............. ...............4.....


LIST OF TABLES ................ ...............9............ ....


LIST OF FIGURES ................. ...............11................


AB S TRAC T ............._. .......... ..............._ 13...


CHAPTER


1 FRUIT PRODUCTION OF TROPICAL DRY FOREST TREES IN BOLIVIA ..................1 5


Introducti on ................. ...............15.................
Methods .............. .. ...... .... .............1

Study Area and Climate............... ...............16
Species Studied...................... .....................1
Experimental Design and Data Collection ................. ...............18........... ...
Data Analysis............... ...............20
Re sults ................ ...............20.................
D discussion ............... ... .......... ... .. ...............22..
Fruiting of Trees in a Tropical Dry Forest .............. ...............22....
Factors Affecting Fruit Production............... ...............2
Conclusions............... ..............2


2 BIOTIC AND ABIOTIC FACTORS AFFECTING TREE SEEDLING DYNAMICS IN
A DRY TROPICAL FOREST............... ...............39.


Introducti on ................. ...............39.................
Methods .............. .. ...... .... .............4

Study Area and Climate............... .. ...............4
Experimental Design and Data Analysis ................. ........... .... .. ........ .......4
Response of regeneration to silvicultural treatment intensity ................. ...............44
Factors affecting seedling establishment and growth............... ...............45.
Re sults................... ............ .............. ........ .......4

Response of Regeneration to Management Intensities ................. ................. ....._48
Factors Affecting Seedling Growth and Establishment .............. .....................4
Seedling growth............... ...............49.
Seedling establishment ................. ...............50.................
D discussion ................ ........... .... ....... .............5

Logging Effects on Seedlings Dynamics .............. ...... ...............51.
Factors Affecting Seedling Establishment and Growth .............. .....................5
Conclusions............... ..............5












3 CONTRIBUTION OF ROOT AND STUMP SPROUTS TO NATURAL
REGENERATION INT A LOGGED TROPICAL DRY FOREST INT BOLIVIA ..................79


Introducti on ................. ...............79.................
M ethods .............. ...............80....

Study A rea ................ .. ........ ....... ...... ...... .............8
Experimental Design and Data Collection .............. ...............81....
Stump sprouts ................. ........... ... .......... .. ... ............8
Comparison of different juvenile types in relation to microsites created by
logging ................. ...............82.......... .....
Data Analysis............... ...............83
R e sults................ .. ......... ...............83.......

Sprout Characterization. .............. ... ...............83.
Juvenile Types and the Effects of Logging ................. ...............86........... ..
Growth of Stump Sprouts ................. ...............86................
Discussion ............... .... .. ........... .. ........... .... .................8
Natural Regeneration and Shade Tolerance: True Seedlings vs. Sprouts .......................88
Allometric Relationships with Stump Sprouting ................. ............ .........._._. ...89
Growth of Stem and Root Sprouts Compared with True Seedlings............... ...............9
Conclusions............... ..............9


LIST OF REFERENCES ................. ...............107...___ ......


BIOGRAPHICAL SKETCH ............ ............ ...............117...










LIST OF TABLES


Table page

1-1. Overview of reproductive characteristics of the tree species studied in a tropical dry
forest in Bolivia............... ...............28

1-2. Results of the best model after backward regression steps to remove non-significant
variables (P > 0.1) sequentially from the full multiple regression model including
tree size (DBH), percent liana infestation, crown area, and crown position as
independent variables to explain fruit production. ............. ...............29.....

1-3. Analysis of covariance to determine the liana cutting effect in fruit production of
Caesalpinia pluviosa.................. ...............30.................

2-1. Spatial distributions, crown position, ecological group, geographical range, tree
densities and basal area of timber species in a Bolivian tropical dry forest. ................... ..59

2-2. Means of seedling densities in an unharvested control plot, a plot subj ected to normal
timber harvesting, and a plot subj ected to more intensive harvesting followed by
silvicultural treatments. ........... ........... ...............60.....

2-3. Mean seedling densities (#/m2) for 3 years (2003-2005) of commercial tree species in
microsites created by logging in a Chiquitano dry forest. .......____ ......_ ..............61

2-4. Establishment and mortality rates of seedlings of commercial tree species monitored
over a 3 y period in a control plot and plots subj ected to two harvesting intensities
(N=144 subplots/treatment plot). .............. ...............62....

2-5. Results of repeated measure analysis of variance for a split plot design run for
seedling density for all species combined or six timber tree species analyzed
separately .. ............... ...............63._____......

2-6. Mean relative height growth rates (A 1SE) of seedlings of commercial tree species in
response to bromeliad cover removal evaluated in 4 times.. ............ ......................6

2-7. Mean relative height growth rates (A 1SE) of seedlings of commercial tree species in
irrigated and droughted plots evaluated 4 times. ............. ...............67.....

2-8. Mean relative height growth rates (A 1SE) of seedlings of commercial tree species in
response to mammal exclosure evaluated 4 times during 2006............... ..................6

3-1. Frequency of root and stem sprouting and shade tolerance of commercial and non-
commercial canopy tree species in a tropical dry forest in Bolivia.................. ...............92

Table 3-2. Summary of the best models and their significance from the regression analyses
between stump diameter or stump volume (independent variables) and the heights of
stump and numbers of sprouts per stump (dependent variables). ........._..__.........._.......93










3-3. Mean (a 1SE) densities of true seedlings, stem sprouts, and root sprouts in 10 x 4 m
plots in microsites created during selective logging. ................... ..............9

3-4. Mean (a 1SE) of stump sprout height growth rates (cm/year) by ecological groups........95

3-5. Means (a 1SE) of stem heights by species that sprouted from stems or roots
compared to the heights of seedlings. .............. ...............96....

3-6. Mean (a 1SE) heights of trees I 2 m tall that were root sprouts, stem sprouts, or
seedlings grouped by ecological guild. .............. ...............97....










LIST OF FIGURES


Figure page

1-1. Locations of the study areas in Bolivia ................. ...............31..............

1-2. Percentage of trees fruiting in 2003 and 2004 (note that none of these species are
dioecious). ............ ...............32.....

1-3. Annual variation in fruit production in three timber species. ........... ......................3

1-4. Comparison of the sizes (DBH) of fruiting and non-fruiting trees in 2003 and 2004
(note that none of these species are dioecious and all the trees were reproductively
m ature). ............. ...............3 4....

1-5. Simple linear regressions between crown area, tree size (DBH) in relation to log
transformed fruit production data for three timber species............... ...............35

1-6. Fruit production of Caesalpinia pluviosa in relation to DBH and crown area for trees
with cut or uncut lianas. .............. ...............36....

1-7. Percentages of trees fruiting in a logged and an unlogged plot. ................ ........._.......37

1-8. Average fruiting intensities (% of crown cover) of timber tree species evaluated in
areas subj ected to normal selective logging and nearby unlogged control areas in a
tropical dry forest. .............. ...............38....

2-1. Monthly rainfall (A) and soil moisture tension measured by Watermark@ soil
sensors (B).. ............ ...............69.....

2-2. Canopy openness in an unlogged control plot, an area subj ected to normal timber
harvesting (4.7 m3/ha harvested), and an area subj ected to intensive harvesting (8.2
m3/ha) 8 months (left-hand bar) and 42 months (right hand bars) after logging............_...70

2-3. Design of the experiment on the effects of ground bromeliads, irrigation, extended
drought, and seed and seedling predator exclosures on seedling establishment. ...............71

2-4. Seedling densities of 11 timber species (A) and 10 species without Acosmium
cardena~sii (B), the most dominant species, in a control plot, a plot subj ected to
normal timber harvesting, and a plot subjected to intensive timber stand
management. ............. ...............72.....

2-5. Temporal changes in seedling recruitment (#/m2/y) and mortality for commercial tree
species in a Chiquitano dry forest in Bolivia. .............. ...............73....

2-6. Seedling density over time in response to (A) bromeliad cover, (B) irrigation or
drought, and (C) mammalian seed predators. ............. ...............74.....










2-7. Mean densities (A 1SE) of seedlings of commercial timber tree species in 2 m2 plOts
(N=40) with bromeliads (filled dots) and without bromeliads (open dots). ....................75

2-8. Mean of seedling densities (A 1SE) in irrigated and droughted experimental plots..........76

2-9. Mean seedling densities (11 SE) in control plots (closed dots) and in plots from
which mammals were excluded (open dots). .............. ...............77....

2-10. Mean relative growth height growth rates (A 1SE) of seedlings of commercial tree
species in response to three experimental treatments: (A) irrigation or drought; (B)
mammalian seed predator exclusion; and, (C) bromeliad cover ................. ................. .78

3-1. Proportions of stumps of commercial timber species that resprouted (number of
stumps noted in parenthesis). ..........__.......__ ...............98..

3-2. The proportions of stumps with live sprouts over time since logging ............... ...............99

3-3. Mean (a 1SE) numbers of sprouts/stump for the most frequent sprouting species.........100

3-4. Probabilities of stump sprouting as a function of stump diameter for the most
frequently sprouting commercial tree species (curves fit by logistic regression). ...........101

3-5. Probabilities of stump sprouting as a function of stump height for the commercial
tree species that most frequently sprouted (curves fit by logistic regression). ................102

3-6. Mean (a 1SE) densities of juveniles < 2 m tall of commercial tree species that were
true seedlings, stem sprouts, and root sprouts............... ...............103

3-7. Percentage of juveniles < 2 m of different origins after logging (for each of 15 dry
forest tree species) ordered by their light requirements ................. ................. ...._104

3-8. Relative growth rates (mean (A 1SE) of stump sprouts measured over the first two
years after logging for commercial tree species in a tropical dry forest in Bolivia
arranged by light requirements. ............. ...............105....

3-9. Mean growth rates of stump sprouts through time for the main commercial tree
species as based on measurements of the tallest sprouts on different stumps 1, 2, and
5 y after creation. ............. ...............106....









Abstract of Dissertation Presented to the Graduate School
of the University of Florida in Partial Fulfillment of the
Requirements for the Degree of Doctor of Philosophy

NATURAL REGENERATION OF CANOPY TREES IN A TROPICAL DRY FOREST IN
B OLIVIA


By

Bonifacio Mostacedo

May 2007

Chair: Francis E. Putz
Major Department: Botany

Fruit production, seedling establishment, and sprouting of canopy trees were studied in a

lowland tropical dry forest in the Department of Santa Cruz, Bolivia. Fruit production by

reproductively mature trees was monitored over a 5 y period to assess variability among species,

trees and years. The effects of tree size, crown area, crown position, and liana infestation on fruit

production were also assessed. In a companion study, I assessed the effects of lianas on fruit

production by Caesalpinia pluviosa with a liana cutting experiment. To determine how logging

disturbances affect seedling recruitment, I monitored seedlings for 3 y in different microsites in

permanent plots in two selectively logged plots and an unlogged control plot. I also

experimentally assessed the effects of bromeliad cover, drought stress, and seed/seedling

predators on seedling recruitment, survival, and growth. Finally, I monitored the emergence,

survival, and growth rates of stump and root sprouts over a range of microsites.

Percentages of trees fruiting and numbers of fruits produced varied among species and

years. In most species, trees that did and did not fruit did not differ in size or crown position, but

in a few cases, the likelihood of fruiting increased with crown area. Contrary to my expectation,

there no effect of liana cutting on Caesalpinia pluviosa fruit production was detected 3 yr after









cutting. Overall, the effect of logging on the proportion of trees fruiting and fruiting intensity

varied among species.

Seedling densities 5 y after selective logging were higher in control than logged plots but

this finding was greatly influenced by the most common species, Acosnzium cardena~sii (43% of

seedlings enumerated). At the microsite level, Acosnzium was found in highest densities in

undisturbed areas while Centrolobium naicrochaete, another common species, was more common

on log extraction paths. Seedling recruitment rates were higher in the unlogged plot and in the

undisturbed portions of the logged forest plots, but seedling mortality rates were also higher in

these areas. Mortality rates of naturally established seedlings varied greatly among species.

Seven of 22 species suffered no mortality during the 2-y monitoring period, whereas relatively

high mortality rates were observed for Caesalpinia (26%/y), Sweetia f~uiticosa (25%/y), and

Machaerium scleroxylon (22%/y).

Results of the experimental study on seedlings suggest that bromeliad competition and

seed/seedling predators greatly affected tree seedling establishment. Soil moisture availability

also affected seedling establishment, but only as an interaction with the bromeliad removal or

predator exclosure treatments. The primary effect of the drought treatment was delayed

germination. Despite these general trends, species varied substantially in their sensitivity to

bromeliads, drought stress, and predators.

Root and stump sprouts constituted about 50% of the individuals <2 m tall of the canopy

tree species studied, but the proportions of sprouts and true seedlings varied among species.

Stump sprouting was common, but the probability of sprouting was not consistently related to

stump diameter or height. Sprout growth rates were consistently high, at least initially, and

sprouting is obviously important to post-logging regeneration in this dry tropical forest.









CHAPTER 1
FRUIT PRODUCTION OF TROPICAL DRY FOREST TREES INT BOLIVIA

Introduction

Tropical dry forests, which until very recently covered about 30% of Bolivia, are among

the most threatened ecosystems in the world (Dinerstein et al. 1995). Bolivian dry forest are under

siege; 32% has already been cleared (Camacho et al. 2001, Rojas et al. 2003) and most of the

remainder is under intensive pressure for forest products, grazing, and further conversion. This

pressure may be somewhat mitigated in the large portion of the dry forest managed for timber under

the guidelines of Bolivia's 1996 Forestry Law (MDSP 1996, Nittler and Nash 1999). Even when

these guidelines are followed, logging disturbances are larger in area than those that occur naturally.

Given the documented effects of disturbance and habitat modification on reproductive success of

tropical dry forest trees (Fuchs et al. 2003), the sustainability of managed forests even when the

legally required "best management" practices are used, remains uncertain. To assess how forest

management influences the reproduction of commercially valuable tree species, I explore the inter-

tree and inter-annual variation in fruit production in a dry forest in lowland Bolivia.

A wide range of mechanistic explanations have been proposed for the marked variation in

fruit production among years and among individuals in a wide variety of forested ecosystems

(Abrahamson and Layne 2003, Snook et al. 2005). For seasonally dry tropical forests, inter-

annual variation in rainfall, particularly as influenced by El Nifio events, has often been invoked

as the underlying cause of inter-annual variation in tree phenology and fruit production (Bullock

1995, Wright and Calderon 2006). Regarding individual tree characteristics that might influence

reproductive output, several studies have reported a minimum size threshold for tree

reproduction (Afiez 2005, Wright et al. 2005). For trees that have attained reproductive maturity,

several studies have shown a relationship between fruit production and stem diameter (Zuidema

and Boot 2002, Wadt et al. 2005), crown area (Healy et al. 1999, Zuidema 2003), and crown










position (Healy et al. 1999). Cover by lianas has also been shown to reduce fruit production by

Brazil nuts "Bertholletia excels Bonpl." (Zuidema and Boot 2002, Wadt et al. 2005, Kainer et

al. 2006) and other species (Stevens 1987).

Anthropogenic and natural disturbances have been reported to influence canopy tree fruit

production in a number of forests around the tropics but the results have been inconsistent.

Logging increased fruit production by remnant trees in a subtropical humid forest in Meghalaya,

India (Barik et al. 1996), as well as in a tropical montane forest in Costa Rica (Guariguata and

Saenz 2002). In contrast, reproductive output of residual trees in a logged dipterocarp forest in

Indonesia was lower than in an unlogged control area (Curran et al. 1999). Factors responsible

for these contrasting results are not clear but, given the importance of natural regeneration to

sustainable forest management, the issue deserves further exploration.

In this study in a seasonal lowland tropical forest in Bolivia I report on canopy tree fruit

production over a 5-year period. I also examine the relationships between fruit production and

crown features, tree size, and liana infestation. Using one common tree species that is often

heavily liana-laden, I assess the effect of liana cutting on fruit production. Finally, I examine the

effect of selective logging on fruit production by several common canopy tree species.

Methods

Study Area and Climate

Research was conducted at INPA Parket, a 30,000-ha tract of privately owned seasonally

dry tropical forest located 30 km northeast of the town of Concepci6n (160 6' 45 S and 610 42'

47" W) and 250 km northeast of the city of Santa Cruz, Bolivia (Figure 1-1). The altitude is

approximately 380 m, mean annual temperature is 24.3 OC, mean annual precipitation is 1150

mm (range 798-1859 mm/y), and there is a dry season that lasts about five months (May-

October) during which most trees are leafless. Extreme annual variation in rainfall is in part









related to the occurrence of El Nifio events, which are typically wet in the study area. Many tree

species in this forest flower at the end of rainy season with another peak in flowering at the

beginning of dry season. Fruiting of the maj ority of species occurs in mid-dry season (see Table

1-1). The forest canopy is 20-25 m tall and dominated by Tabebuia impetiginosa (Mart. Ex DC.)

Standl., Anadenanthera macrocarpa (Benth.) Brenan, Astronium urundeuva (Allemio) Engl.,

and Centrolobium microchaete (Mart. ex Benth.) Lima ex G. P. Lewis (Pariona 2006). The most

abundant species are Acosmium cardena~sii H. S. Irwin & Arroyo (- 38 trees/ha > 20 cm

diameter at breast height, DBH) and Anadenanthera (8 trees/ha > 20 cm DBH). Currently, 21

tree species, including these dominants, are harvested for timber used mostly in the production of

parquet flooring.

Species Studied

My studies focus on commercially valuable timber tree species, most of which produce

seeds that are wind, gravity, or explosively dispersed (Table 1-1). Overall I consider 31 species

(Table 1-1), of which 15 were used to determine the percentage of fruiting trees, 12 to compare

the sizes of reproductively mature trees that did or did not fruit during a 2-y observation period, 6

to compare fruiting intensity in logged and unlogged areas, and 3 to determine the relationships

between fruit production and DBH, crown area, crown position, and liana infestation. In

addition, one species (Caesalpinia pluviosa DC.) was used to study the effect of liana cutting on

fruit production. Most of the selected species have large or medium-sized fruits with small- to

intermediate- sized seeds.

Of the 3 1 species considered in this study 25 have hermaphroditic flowers and 20 produce

wind-dispersed seeds. Ten species have seeds as their dispersal units (e.g., Amburana cearensis

(Allemao) A.C. Sm.) while in the other 21 species, entire fruits are the dispersal units (e.g.,

Machaerium scleroxylon Tul.). Twenty-four of the 3 1 species monitored for five years produced









at least some fruits annually, but the intensity of fruiting varied substantially. Cecropia concolor

Willd. produced fruits continuously whereas Schinopsis brasiliensis Engl. fruited supra-annually

(sensu Newstrom et al. 1994). Peak fruiting for 23 of 31 species coincided with the dry season

(May September) with most seeds dispersed in the late dry season. Spondia~s mombin L. the

only fleshy-fruited species in the study produced ripe fruit during the rainy season. Most of the

species have only 1 seed per fruit, but Ceiba samnauma (Mart.) K. Schum. and Tabebuia have

more than 90 seeds/fruit. Wind-dispersed seeds or fruits typically have one (Pterogyne nitens

Tul.) or both sides winged (e.g., Platimiscium ulei Harms), whereas Zeyheria tuberculosa (Vell.)

Bureau seeds are entirely surrounded by a wing. Cottony fibers on the seeds of the

Bombacaceae studied (Ceiba and Chorisia speciosa A. St.-Hil.) aid in their dispersal by wind.

Experimental Design and Data Collection

The study was conducted at two different sites at INPA Parket. The first site was within the

Long-Term Silvicultural Research Plots (LTSRP) in Block # 1, 5-6 km south of the IBIF field

station (160 18' 26.8"S, 610 41' 13.5"W). Four large-scales (20 ha) long-term research plots

(LTRSP) were established by the Instituto Boliviano de Investigacion Forestal (IBIF). One of the

plots I used is an unlogged control whereas the other was subj ected to normal logging during

which a mean of 4.3 trees/ha (4.7 m3/ha of commercial timber) were harvested using standard

reduced-impact-logging techniques that include road planning, directional felling, and retention

of 20% of the harvestable trees as seed trees (Mostacedo et al. 2006). In each plot, I censused all

individuals > 20 cm DBH of the 15 most abundant tree species in two 20 x 400 m strips. The

second site is a 1000 x 500 m plot that I established 2 km northeast of the IBIF field station (160

14' 58.5" S, 610 41' 47.4" W). In this plot, I censused individuals > 20 cm DBH of

Anadenanthera and Caesalpinia in twelve 20 x 500 m strips spaced at 100 m intervals. At both










study sites the censused trees were marked, tagged, and mapped. To secure sufficient trees of

Caesalpinia and Zeyheria, I marked additional trees around the LTRSP.

In total, I evaluated 440 trees of 15 species, 116 in the control plot and 147 in the logged

plot of the LTRSPs. An additional 144 trees were monitored at the second site that, a year after

plot establishment, was selectively logged. In 2003, I evaluated only six species from May-

November including Caesalpinia, Anadenanthera, Machaerium scleroxylon, Ceiba, and

Zeyheria. In 2004, every marked tree in the LTSRP plots was phenologically evaluated Hyve

times from May to November. I estimated the percentage of the crown bearing fruits, which I

refer to as a measure of fruiting intensity; 100% fruiting intensity indicates that every terminal

branch bears at least one fruit.

I estimated the number of fruits produced by each tree of three species, Caesalpinia,

Anadenanthera, and Zeyheria, for 5 years (2002-2006). All of these species produce fruits with

valves that are not removed by animals. By counting these undispersed fruit parts under fruiting

trees, I avoided many of the difficulties associated with estimating seed production by species in

which entire fruits are removed by animals. At the beginning of the study I installed Hyve 2x2 m

permanent plots on the ground below the crown of each tree in which I counted and removed all

fruit valves. Subsequent censuses of fallen fruit valves in these plots over the next 5 y were used

as quantitative estimates of fruit production. I also measured the DBH, crown area (based on two

cardinal diameters), crown position (using the 5 categories of Dawkins (1958)), and liana

infestation.

Liana-infested Caesalpinia trees (N=32) were used in a manipulative experiment on the

effects of lianas on seed production. I measured the DBH and percentage of each tree' s crown

covered by lianas, paired the trees on the basis of liana infestation and DBH, and cut all the









lianas on one tree of each pair, selected at random. Fruit production was measured for 3 y after

treatment using the fruit-valve census method described above.

Data Analysis

To determine whether trees of the 12 monitored species that fruited in 2003 and 2004

differed in DBH from those that did not, I conducted Student's-t tests. I tested for simple linear

relationships between the number of fruits (log-transformed) produced per tree and each tree' s

DBH, crown area, crown position, and liana cover separately and then ran multiple regressions

for three monitored species (Anadenanthera, Caesalpinia, and Zeyheria) using the backward

method to avoid co-linearity. Only independent variables that explained significant amounts of

variance (P=<0.05) were included in the models. I ran X2 tests to assess differences between

logged and control plots in the percentage of trees fruiting. To compare maximum fruiting

intensities in the logged and the control plot for the 6 monitored species, I ran Student' s t-tests

after first arcsine transforming the data to achieve normality (Zar 1981). Finally, I compared fruit

production on liana-laden and liana-free trees using analyses of covariance with crown area or

DBH as covariates. For all analyses, I used SPSS Version 12.0 (Field 2000).

Results

The percentage of trees fruiting in any particular year varied among species (Figure 1-2).

Of 14 species monitored during 2004, 7 had >50% of trees in fruit and 4 had >80% of trees in

fruit. For a few species that I also monitored in 2003, the percentage of fruiting trees was lower

in 2003 than in 2004. For example, while only 25% of Caesalpinia trees fruited in 2003, 100%

fruited in 2004. The opposite pattern was observed in Zeyheria; 40% fruited in 2003 and none in

2004.

For the 3 species I monitored for 5 years, there was a great deal of inter-annual variation in

fruit production (Figure 1-3). For example, a high proportion of Caesalpinia trees fruited at 2-









year intervals whereas many Zeyheria trees fruited at 3-year intervals. In contrast,

Anadenanthera fruited during each of the first 3 years of the study and not at all in the last 2.

Whether or not a tree fruited was generally not related to its DBH (Figure 1-4). The

exceptions were Caesalpinia in 2004 when the trees that failed to fruit were larger than the trees

that fruited (t=2.18, P=0.03) and Zeyheria in which the fruiting trees were larger in 2004, the

only year in which it fruited (t=4.91, P<0.0001).

For the 3 species in which I monitored fruit production, the number of fruits produced did

not vary with DBH or crown position but increased linearly with crown area (Figure 1-5). The

backward multiple regression of fruit production on tree characteristics revealed that crown area

explained the most variation (Table 1-2). In Anadenanthera, crown area explained 32% of the

variance in fruit production while DBH, crown position, and liana cover together explained only

an additional 10%. In the case of Caesalpinia, crown area explained 23% of the variance in fruit

production and the other three variables only an additional 3%. In Zeyheria, crown area and

crown position together explained only 24% of the variance, to which DBH and liana cover

added an additional 1%.

There was no apparent effect of liana cutting on fruit production by Caesalpinia when the

effects of either DBH or crown area are removed by ANCOVA (Table 1-3, Figure 1-6).

The percentage of trees fruiting in the logged and control plot was similar for 6 of 7

species. The only exception was Centrolobium in which only 27% of trees fruiting in the logged

plot compared to 90% in the control plot (X2=12.9, P<0.0001; Figure 1-7).

The effect of logging on fruit production varied among the 6 tree species studied (Figure 1-

8). Logging apparently stimulated increased fruiting intensity ofAnadenanthera (t=3.40,

P=<0.0001). In contrast, fruiting intensity was higher in the unlogged plot for both Centrolobium










(t=4.75, P=0.0008) and Copaifera (t=3.11, P=0.007). There was no difference in fruiting

intensity between the logged and unlogged plot for Aspidospernza (t=0.68, P=0.5) and

Machaerium scleroxylon (t=0.05, P=0.95).

Discussion

Fruiting of Trees in a Tropical Dry Forest

In any year, the proportion of reproductively mature trees that fruited generally varied a

great deal among the canopy tree species I studied in lowland Bolivia. Only Anadenanthera trees

produced fruit crops in both 2003 and 2004. In contrast, no trees of2achaerium scleroxylon and

M~ acutifolium fruited in 2003, but many did in 2004. Most of the trees of some species fruited in

at least some years (e.g., Caesalpinia) while in others the percentage of fruiting trees was always

<40%. Perhaps coincidentally, the five species with the greatest proportion of fruiting trees in

2004 were all legumes, which comprised 8 of the 14 species monitored. Few studies report

annual variation in the proportions of fruiting trees but in a similar forest in Lomerio, Bolivia,

only 29% and 36% of reproductively mature trees of 17 commercial tree species fruited during

two years of monitoring (Justiniano and Fredericksen 2000). At the same site, there was a great

deal of variation within species in the proportions of fruiting trees. For example, the proportion

of Copaifera trees fruiting was similar to what I observed in INPA whereas none of the M~

scleroxylon trees in Lomeria fruited during the two years of monitoring. In a tropical dry forest

in Mexico studied by Bullock (1995), only 8-30% of Jacaratia nzexicana A. DC. trees and 0-

50% of Cochlospernaun vitifolium (Willd.) Spreng. trees fruited in any one year. Similarly, in a

study of Swietenia nzacrophylla King. on the Yucatan Peninsula, the proportion of fruiting trees

varied a greatly among years (Snook et al. 2005). The proportion of Hynzenaea courbaril L. trees

fruiting in Costa Rica reportedly varied with water stress (Janzen 1978).









A larger proportion of trees of most species fruited in 2004 than in 2003 or 2005. Such

community-wide synchrony in fruiting has been observed in many forests over the world but has

been particularly well studied in the dipterocarp forests of Southeast Asia where "masting" has

long been known (Appanah 1993, Wich and Van Schaik 2000). Less pronounced is the inter-

annual variation in fruit production on Barro Colorado Island, where mast years are reportedly

related to El Nifio events (Wright and Calderon 2006). In the forest of this study, 2003 was at the

end of an El Nifio event, which was not followed by a strong La Nifia. Nevertheless,

synchronous fruiting in 2004 could have been due to the timing of water stress, as suggested by

Bullock (1995).

The three species I followed for five years showed great inter-annual variation in the

numbers of fruits produced per tree. The two legumes, Anadenanthera and Caesalpinia,

followed the same interannual patterns; both had peak years in 2002 and 2004, but produced few

fruits in 2003 and 2005. In contrast, the 35 Zeyheria trees I monitored produced no fruits in 2004

and 2005, but fruited in 2002, 2003, and 2006. Such interannual variation in fruit production is

common in many species. For example, in a moist tropical forest in Panama, Quararibea

a~sterolepis Pittier, Tetraga;stris pa~namennsi (Engl.) Kuntze, and Trichilia tuberculata (Triana &

Planch.) C. DC. showed great inter-annual variation in fruit production (De Steven and Wright

2002, Snook et al. 2005).

While I expected that the proportion of fruiting trees would increase with tree size, 12 of

the 13 species I monitored in 2003 and 2004 did not display this pattern. In fact, in the case of

Caesalpinia, the fruiting trees were significantly smaller than those that failed to fruit in 2004.

Only Zeyheria displayed the expected pattern. The general failure to find a positive relationship

between tree size and whether or not a tree reproduced runs counter to the results of two other









studies on this topic, one conducted on Barro Colorado Island, Panama (Wright et al. 2005) and

the other at my study site (Afiez 2005). One explanation for the difference between my study and

others in the literature is that whereas they typically used either flowering or fruiting as an

indication of reproduction (Afiez 2005, Wright et al. 2005), I used only fruiting. Nevertheless,

while the probability of fruiting increased with tree size in Copaifera, Sweetia and Machaerium,

it decreased in Anadenanthera and Caesalpinia.

Factors Affecting Fruit Production

Crown area was the best predictor of fruit production in many of the species I studied

whereas DBH and crown position were not. Crown area was also a good predictor of fruit

production in Betholletia excelsa trees in Amazonian Bolivia (Zuidema and Boot 2002), while

DBH was more closely related to fruit production by Swietenia in Mexico (Snook et al. 2005)

and Pinus sylvestris L. in Sweden (Karlsson 2000). Although several studies have shown strong

positive correlations between tree size and fruit production (Karlsson 2000, Zuidema and Boot

2002, Wadt et al. 2005), the relationships revealed in my study were positive but weak. This

difference might be related to the difficulty of making accurate DBH estimates of the trees in my

study site, many of which have irregular-shaped trunks. It is also possible that crown position is

not good predictor of fruit production in forests that are already open-canopied; certainly in my

study site it was difficult to differentiate between dominant and co-dominant trees, and I doubt

that they differ much in terms of light capture.

I expected liana cover to substantially impede fruit production (Stevens 1987), but my

results did not support that expectation. For the 16 pairs of Caesalpinia trees from which I

monitored fruit production for 3 y after cutting their lianas, fruit production was no higher than

in the liana-laden control trees. I can offer a few explanations for this counter-intuitive result. For

one thing, 3 y was perhaps not enough time for the trees to respond to the liberation from their









liana loads. Then there is the problem that fruit production is extremely variable among

individuals and among years in this species and others in my study site. Finally, I wonder

whether the liana leaf phenology at my study site might have something to do with this result

insofar as lianas are typically deciduous at the time of fruiting, which might reduce any

deleterious effect they have on fruit production. Obviously, none of these explanations is very

compelling and further monitoring is warranted.

Although I anticipated that by increasing canopy openness and reducing resource

competition, logging would increase fruit production by remnant trees, my results were

inconsistent at best. Firstly, the percentage of fruiting trees in most of species were similar

between logged and control plots, except for Centrolobium. For some species (e.g., Copaifera,

Aspidosperma, and Sweetia) in which there seemed to be a trend towards increased reproduction

among remnant trees in selectively logged forest, statistical significance was not forthcoming

perhaps due to small sample sizes (N < 20).

Secondly, 3 of the 6 species I monitored starting 1.5 y after selective logging showed no

apparent effect of the treatment on fruit production, 2 of the species fruited less in the logged

than in the control plot, and 1 species showed the opposite pattern. One reason for the observed

decrease in fruit production might have been the effect of lowered tree density on pollinator

effectiveness or offspring quality (Ghazoul and Shaanker 2004, Knight et al. 2005). Other factors

that might have reduced fruit production include fruit abortion due to poor ovule fertilization in

the extreme temperatures and low humid provoked by forest openness (Stephenson 1981, Aizen

and Feinsinger 1994, Dafni and Firmage 2000) or due seed predation during the early stages of

fruit formation (Stephenson 1981). For example, 1 tree/ha > 40 cm DBH of Copaifera remained

after logging of the pre-logging density of 1.7 trees/ha whereas in the unlogged plot, the










population of adult trees was substantially higher (2.7 trees/ha). Similarly, in Southeast Asia,

fruit production was reduced in a logged dipterocarp forest (Curran et al. 1999). In contrast, the

most light-demanding tree species in my study forest, Anadenanthera, produced more fruit in

logged than in unlogged areas. Similarly, Ceiba aesculifolia (Kunth) Britten & Baker f. in a dry

forest in Mexico (Herrerias-Diego et al. 2006) and Quercus costaricensis in montane humid

forest in Costa Rica (Guariguata and Saenz 2002) both increased fruit production in response to

logging. Given the importance of this issue to sustainable timber stand management, more

research is needed on the reproductive responses of trees to logging disturbances. These studies

should integrate flower production, pollination, and fruit production and should be conducted on

species representing a variety of densities and breeding systems.

Conclusions

In the seasonally dry tropical forest I studied in Bolivia, there was a lot of interannual

variation in fruit production among species. Whether or not a reproductively mature tree fruits is

generally not related to it size. The number of fruits produced per tree also did not consistently

change with stem diameter or liana cover, but did increase with crown size. Base on these

findings, I recommend protection of tree crowns during logging to increase fruit production.

Even when I experimentally killed the lianas infesting the crowns of 16 Caesalpinia trees, I did

not observe the expected increase in fruit production. To look at the effect of lianas cutting I

recommend a long-term study to look at crown area recovery and fruit production. Similarly,

after logging, remnant trees of some species increased their fruit production and some species

decreased but most did not change. For most of the species, percentages of trees fruiting were

similar between the control and the logged plot; only in Centrolobium was there a difference,

with more trees fruiting in the control plot. The small and inconsistent effects of logging on fruit

production in my study forest may be explained in part by the relatively low intensity of










harvesting. Obviously, given the substantial inter-tree, inter-specific, and inter-annual variation

in the fruiting of dry forest tree species, studies of more than a few years are needed.
















Table 1-1. Overview of reproductive characteristics of the tree species studied in a tropical dry forest in Bolivia.


Species

4cosnzinn cardenasii

4nibumana cearensis
4nadenanthern
niacrocarpa
4spidosperyna rigidunt
4stronium urundeuva
4ttalea phalemata
Caesalpinia pluviosa
Capparis prisca
Caniniana laneirensis
Cecropia concolor
Cedrela fissilis
Ceiba saniania
Centrolobium
nzicrochaete
00 Chonisia speciosa
Copaifery chodatiana
Cordia alliodorn
Gallesia integrifolia
Genipa americana
Hvnzenaea courbaril
M2/achaen'unt cf.

M2/achaerium scleroxvlon
AMvrcianthes spp.
Phyllostvlon rhaninoides
Platintiscium ulei
Pterogyne nitens
Schinopsis brasiliensis
Spondias nionbin
Sweetia fruticosa
Syagrus sancona
Tabebuia intpetiginosa
Tabebuia \renu~rit;dia
Zevhen'a tuberculosa


Family

Fabaceae


Abbreviation Breeding
System
ACCA Hermaphrodite


Dispersal
Type
Wind

Wind
Gravity

Wind
Wind
Animal
Explosive
Animal
Wind
Animal
Wind
Wind
Wind

Wind
Gravity
Wind
Wind
Animal
Gravity
Wind

Wind
Animal
Wind
Wind
Wind
Wind
Animal
Wind
Zoo
Wind
Wind
Wind


Dispersal
Unit
Fnuit

Seed
Seed

Seed
Fmuit
Fnuit
Fnuit
Fnuit
Seed
Fnuit
Seed
Seed
Fnuit

Seed
Seed
Fnuit
Fnuit
Fnuit
Fmuit
Fnuit

Fnuit
Fnuit
Fmuit
Fnuit
Fnuit
Fnuit
Fnuit
Fnuit
Fnuit
Seed
Seed
Seed


Fnuiting*
Frequency
Continuous

Annual
Annual

Annual
Annual
Continuous
Annual
Annual
Annual
Continuous
Annual
Annual
Annual

Annual
Annual
Annual
Annual
Annual
Annual
Supra-annual

Annual
Annual
Supra-annual
Annual
Annual
Supra-annual
Annual
Annual
Annual
Annual
Annual
Supra-annual


Fmuiting

Mar-Aug, Oct-
Feb
Jun-Aug
Mar-Sep

Mar-Sep
Aug-Oct
Oct-Apr
Mar-Sep

May-Sep
Oct-Mar
Jul-Aug?
Jun-Aug
Apr-Sep

Jul-Aug
Mar-Sep
Jul-Ago
May-Sep
Dec-Feb
Jul-Oct
Apr-May

Apr-Sep
Jan-Feb
Oct
Jul-Aug
Apr-May
Ago-Sep
Jan-Feb
Oct
Nov-Dect
Jul
Ago
Jun-Jul


Seed
Dispersal
May-Sep,
Feb
Sep
Jul-Oct

Sep-Nov
Oct
Feb-May
Ago-Nov

Oct
Feb-Mar
Aug-Sep
Ago
Jul-Sep

Sep
Jul-Oct
Ago
Oct-Nov
Feb-Mar
Oct-Nov
May

Aug-Sep
Feb
Oct
Aug-Sep
May-Aug
Sep
Feb
Oct-Nov
Dec-Jan
Ago
Sep
Jul


# Seeds /
Fmuit
1.2

1
10.9


Fabaceae AMCE
Mimosaceae ANMA


Hermaphrodite
Hermaphrodite

Hermaphrodite
Dioecious
Monoecious
Hermaphrodite
Hermaphrodite
Hermaphrodite
Monoecious
Hermaphrodite
Hermaphrodite
Hermaphrodite

Hermaphrodite
Hermaphrodite
Hermaphrodite
Hermaphrodite
Hermaphrodite
Hermaphrodite
Hermaphrodite

Hermaphrodite
Hermaphrodite
Hermaphrodite
Hermaphrodite
Hermaphrodite
Dioecious
Hermaphrodite
Hermaphrodite
Monoecious
Hermaphrodite
Hermaphrodite
Hermaphrodite


Apocynaceae
Anacardiaceae
Arecaceae
Mimosaceae
Capparaceae
Lecythidaceae
Cecropiaceae
Meliaceae
Bombacaceae
Fabaceae

Bombacaceae
Caesalpinaceae
Boraginaceae
Phytolacaceae
Rubiaceae
Caesalpinaceae
Fabaceae

Fabaceae
Myrtaceae
Rhamnaceae
Fabaceae
Caesalpinaceae
Anacardiaceae
Anacardiaceae
Fabaceae
Arecaceae
Bignoniaceae
Bignoniaceae
Bignoniaceae


ASRI
ASUR
ATPH
CAPL
CAPR
CAIA
CECO
CEFI
CESA
CEMI

CHSP
COCH
COAL
GAIN
GEAM
HYCO
MAAC

MASC
MYSP
PHRH
PLUL
PTNI
SCBR
SPMO
SWFR
SYSA
TAIM
TASE
ZETU


(*) Clasification made by Newstron et al. (1994). (Jf) Data extracted from Justiniano and Fredericksen (2000).











Table 1-2. Results of the best model after backward regression steps to remove non-significant
variables (P > 0.1) sequentially from the full multiple regression model including tree
size (DBH), percent liana infestation, crown area, and crown position as independent
variables to explain fruit production. Number of trees (N), standardized regression
coefficients (P), Student-t test values (t), significance levels (P), and coefficients of
multiple determination (R2) are noted.


P R2


Variables


Anadenanthera macrocarpa
Crown Area

Caesalpinia pluviosa
Crown Area


30 0.560 3.59


50 0.480 3.81


0.001 0.32***


<0.0001 0.23**


Zeyheria tuberculosa
Crown Area
Crown Position


0.450 2.89
0.282 1.81


0.007
0.080


0.24*


* P 5 0.05; ** P 5 0.01; *** P I 0.001.










Table 1-3. Analysis of covariance to determine the liana cutting effect in fruit production of
Caesalpinia pluviosa. The co-variables considered were crown area and DBH.
Source Crown Area DBH
Mean F P Mean F P
Square Square
Co-variable 42438.5 2.35 0.14 22090.0 1.18 0.29
Liana cutting 9135.0 0.57 0.48 9673.0 0.52 0.48
Error 18020.7 18747.0


















































Legend
INPA,
jrIBIF: Field Station
Main Roads

I::::IStudy Area 1
~T1 Study Area 2
SStuldy Area 3




1 0.5 0 1 2 3
sersaa I e-- nm Kilorneters


\---









-** : = -


I
628000


I
632000


I
636000


I
640000


I
644000


Figure 1-1. Locations of the study areas in Bolivia. Study Area 1 was where I conducted fruit
production research (Chapter 1); Study Area 2 was used for seedling recruitment
study (Chapter 2); and, Study Area 3 was the site for the sprouting studies (Chapter
3). The map is georeferenced using a Universal Transverse Mecator (UTM) System,
Zone 20.


































Species
.Percentage of trees fruiting in 2003 and 2004 (note that none of these species are
dioecious). Asterisks indicate years in which a species that was not evaluated.
Abbreviations of species are given in Table 1-1. Numbers in parenthesis indicate the
number of trees evaluated for each species and year.


-


20 -




Figure 1-2


CAPL ANMA ACCA COCH MASC CESA GAIN CEMI ASRI SWFR CAIA CAPR MAAC ZETU












600


500-

400-

300-

200-

100-



10002002 2003 2004 2005 2006
e Caesalpinia pluviosa

1- 800 -


.2 600-


S 400-


t*' 200-



80 2002 2003 2004 2005 2006
Zeyheria tuberculosa .

60-


40-




20


2002 2003 2004 2005 2006

Year
Figure 1-3. Annual variation in fruit production in three timber species. Vertical lines show one
standard error. Note the differences in the y-axis scales.


















Acosmiun









12 68
NO YES

Caesalpinia









99 24 3 45
NO YES NO YES

Copaifera









3 13
NO YES

M/. scleroxylon*


Aspidosperma









8 5
NO YES

Centrolobium









144 36 28 19
NO YES NO YES

M. acutifolium









21 8
NO YES

Zeyheria .









69 43
NO YES


Anadenanthera
1 Year 2003
IIIIII Year 2003
o Year 2004
M 1Year 2004





114 258 7 60
NO YES NO YES

Ceiba









8 7
NO YES

Gallesia









5 8
NO YES

Swetia


5 18
NO YES


11 6
NO YES


Fruiting Trees

Figure 1-4. Comparison of the sizes (DBH) of fruiting and non-fruiting trees in 2003 and 2004
(note that none of these species are dioecious and all the trees were reproductively
mature). Numbers indicate the sample size for each year and each species. Asterisks
indicate significant size differences between fruiting and non-fruiting trees as
determined with Student-t test. P-value: I 0.05, ** < 0.001, *** I 0.001.

























Log(Y) 1 47+0 007(X),
r =0 32, F=12 92,
P=0 001

) 50 100 150 200 250


Spearman's r=-0 25, P=0 18


Spearman's r=-0 135, P=0 35


0 50 100 150 200 250












Log (Y) = 0 91+0 004 (X),
r2=0 17, F=6 53,
P=0 01


Spearman's r=0 012, P=0 95


Anadenanthera macrocarpa


1000-


100-


10-


20 30 40 50 60 70 80
Caesalpinia pluviosa


Dom Codom


10


Log (Y) = 1 64+0 004 (X),
r2=0 23, F=14 55,
P=<0 0001


30 40 50 60 70 80

Zeyheria tuberculosa


Dom Cod~om Interm


0 50 100 150 200 250


20 30 40 50 60 70 80


Dom Codom Interm


Crown Area DBH Crown Position

Figure 1-5. Simple linear regressions between crown area, tree size (DBH) in relation to log
transformed fruit production data for three timber species. Spearman correlations
were run to determine the relationships between crown position (ordinal variable) and
fruit production. Crown position numbers refer to dominant (Dom), co-dominant
(Codom), and intermediate exposure (Interm) trees.


Log (Y) =1 86+0 01(X),
12=0 05, F=1 49,
P=0 23


Log (Y)= 1 71+0 013 (X),
r2=0 06, F=3 19,
P=0 08


Log (Y) = 0 59+0 16 (X),
r2=0 08, F=3 07,
P=0 08










I


I


I


10000


SLianas Cut
O control

O0 O


ge* o a



O e


1000 -



100 -


10 -



1


10000 -



1000 -



100 -



10 -



1-


O O
O g*O O *

o o







SO


100


150


200


250


300


Crown Area (m2)
Figure 1-6. Fruit production of Caesalpinia pluviosa in relation to DBH and crown area for trees
with cut or uncut lianas.


DBH (cm)













100-





80-





2 60-
L.
O


S40-





20-





0-




Figure 1-7.


(34Y (139 1 (5) 1(1 21 I12)
Caesalpinia Anadenanthera M~achaerium Aspidosperma Sweetia


Centrolobium Copaifera


Species
Percentages of trees fruiting in a logged and an unlogged plot. Number of trees
evaluated is noted in parenthesis. Differences between logging intensities were tested
using X2 test at 95% of confidence. *** <0.0001; ns non significant.













SAnadenanthera macrocarpa l uu


Aspidosperma rigidum


May04 Jul04 Sep04 Oct04 Nov04


100

80

60

40

20

0




S100

S80

60 -

S2-




0

L.


80-

60 _

40 -

20 -

0 -




100

80

60

40

20

0

10-


8-
100

4-
80

0-


-* Normal Logging
-0 Control


May04 Jul04 Sep04 Oct04 Nov04


May04 Jul04 Sep04 Oct04 Nov04


Caesalpinia pluviosa


Centrolobium
microchaete


May04 Jul04 Sep04 Oct04 Nov04


100

80

60

40

20

0


Copaifera chodatiana


Machaerium scleroxylon


May04 Jul04 Sep04 Oct04 Nov04


May04 Jul04 Sep04 Oct04 Nov04


Months

Figure 1-8. Average fruiting intensities (% of crown cover) of timber tree species evaluated in
areas subj ected to normal selective logging and nearby unlogged control areas in a
tropical dry forest. Vertical lines indicate standard errors.









CHAPTER 2
BIOTIC AND ABIOTIC FACTORS AFFECTING TREE SEEDLING DYNAMICS IN A DRY
TROPICAL FOREST

Introduction

Among the many factors that affect tree seedling establishment in tropical forests, seed

predation, pathogen effects on seedlings and light availability figure prominently (Janzen 1971,

Augspurger 1984, Hammond 1995, Huante et al. 1998, Kobe 1999). Given that this view is

mostly supported by studies conducted in moist and wet forests, the relative importance of these

factors might differ in seasonally dry forests (Gerhardt 1994, Holbrook et al. 1995). Given the

widespread mismanagement and destruction of these forests (Steininger et al. 2001, Pacheco

2006), it is increasingly important to increase our knowledge of ecological processes, such as

regeneration, that might lead to their improved management. To further this knowledge, I

monitored seedling populations and conducted experimental studies across a gradient of forest

management intensities in a seasonally dry lowland tropical forest in Bolivia. In the experiments,

I planted seeds of canopy tree species and manipulated moisture availability, seed and seedling

predation, and competition with an abundant understory bromeliad, Pseudananas~ddd~~~ddd~~~dd sagenarius

(Arruda) Camargo).

Seasonally dry tropical forests are characterized by low total rainfall (typically < 1500

mm) and annual periods of drought (i.e., months during which evapotranspiration exceeds

precipitati on usually as sumed to b e months with < 100 mm of precipitati on)(Hol dri dge 1 967).

The rainfall regime of Bolivian, and apparently many other tropical dry forests (Gerhardt 1996a),

is also characterized by huge interannual variation in precipitation, dry season duration, temporal

continuity of rainfall during the rainy and dry seasons, and the starting and ending dates of the

seasons. In addition, paleoecological and archaeological data predict that the dry seasons in

tropical dry forests in South America will be extended in duration and drier in the future (Mayle









et al. 2007). These sources of environmental variation are critical because, although the tree

species characteristic of tropical dry forests are presumably adapted to seasonal drought,

mortality is reportedly concentrated during dry seasons and dry years (Khurana and Singh 2001,

McLaren and McDonald 2003).

Within species and age cohorts, the probability of mortality reportedly decreases with

increasing seedling size, presumably because larger individuals have greater access to soil water

(Khurana and Singh 2001). Seedling growth rates typically increase with increasing illumination

but, due to high temperatures and soil surface drying, seedlings growing in large canopy gaps

may suffer high risks of mortality, even if they grow larger than their shaded counterparts in the

understory. In other words, in a forest with a 6-month rainy season, a newly germinated seedling

has 6 mo to grow large enough to survive the subsequent dry season. Based on this idea, I

conducted an experiment in which I manipulated the duration and intensity of the dry season and

monitored the survival and growth of seedlings of canopy tree species that germinated from

sown seeds. In recognition of the importance of inter-specific competition and seed/seedling

predators to seedling dynamics, I also manipulated these factors in a factorial experiment.

In addition to tolerating drought, tropical tree seedlings must compete for light, water,

and nutrients with plants of various growth forms including lianas (Gerwing 2001, Schnitzer and

Bongers 2002), ferns (George and Bazzaz 1999), and ground bromeliads (Fredericksen et al.

1999, George and Bazzaz 1999). In the seasonally dry forests of the Chiquitania of lowland

Bolivia, competition with a clonal ground bromeliad, Pseudananas~ddd~~~ddd~~~dd sagenarius, seems

particularly intense. This bromeliad covers 25-30% of the ground surface in my study area where

it presumably competes for light and soil resources. Furthermore, its leaves intercept both rain

and falling seeds (Fredericksen et al. 1999), which should affect tree seedling recruitment,









growth, and survival. Finally, by providing cover to small mammals and other animals avoiding

predators, bromeliads may also affect rates of seed and seedling predation. To investigate these

effects, I included a bromeliad removal treatment in my experimental study on the survival and

growth of canopy tree seedlings.

Predators often reduce seed and seedling survival in tropical forests (Janzen 1971, Hulme

1996, Asquith et al. 1997). Given that seed predation rates, at least by mammals, typically

increase with seed size (Moles and Westoby 2003), and given that most tree species in dry

tropical forests have small, wind-dispersed seeds, rates of seed predation may be lower in dry

than in humid forests. On the other hand, given the seasonal scarcities of food in dry forests, seed

and seedling predation might be particularly intense, especially for seedlings that remain leafed

out and succulent during the early dry season. To investigate the importance of seed and seedling

predation, as influenced by seasonality, I experimentally manipulated both soil moisture and

accessibility to predators using the planted seeds of canopy tree species.

In this chapter I present the results of a study in which I monitored the survival and

growth of tree seeds planted in plots in which I experimentally manipulated soil moisture, seed

and seedling predator access, and competition with ground bromeliads. The chapter also includes

the results of 3 y of monitoring of seedling establishment, growth, and survival in a large control

plot and two otherwise similar plots subj ected to different intensities of timber harvesting 7 mo

prior to commencement of my study. I hypothesized that seedling survival and growth are

reduced by bromeliad cover, drought stress, and predators. I also hypothesized that the

establishment, survival, and growth of naturally regenerated seedlings differs among plots

subj ected to different intensities of disturbance resulting from forest management activities.












Study Area and Climate

This study was conducted during 2003-2006 in the 30,000 ha seasonally dry lowland forest

(Holdridge 1967) owned by INPA PARKET, about 30 km from Concepci6n, Bolivia (160 6'

45"S, 610 42' 47"; Figure 1-1). The study area is located in the transition zone between the

forests of the Amazon Basin and those of the Gran Chaco and is locally referred to as Chiquitano

dry forest (Killeen et al. 1998, Ibisch and Merida 2003). Based on 30 years of data collected in

Concepci6n, the annual average temperature in the area is 24.3 OC with a minimum average

temperature of 12.90 C, generally in July, and maximum average temperature of 310C, generally

in November. Over the 30-year monitoring period, the annual average of rainfall was 1 100 mm

but ranged from 700-2000 mm/year, with wet years corresponding with El Nifio events (Coelho

et al. 2002) (Figure 2-1). Both 2005 and 2006 were dry years, with only 980 and 1050 mm of

precipitation, respectively. The rainy season typically runs from October to April, but in 2005,

the rains started in September and rain fell at regular intervals until March of 2006. The 2006

rainy season, in contrast, started in October but then no rain fell in November, with more regular

rains commencing again in December.

The set of Long-Term Silvicultural Research Plot (LTSRPs) established at this site by the

Institute Boliviano de Investigaciones Forestales (IBIF) was used for a portion of this study. The

LTSRP at INPA includes four 20-ha plots that vary in management intensity: unlogged control;

normal logging; improved management; and, intensive management (Mostacedo et al. 2006). For

this study, the improved management plot was not used. In the "normal logging" plot, the

logging company harvested a mean of 4.3 trees/ha (4.7 m3/ha) using their standard reduced-

impact-logging techniques that include road planning, directional felling, and the retention of

20% of the harvestable trees as seed trees. The mean density of trees > 10 cm DBH before


Methods









logging was 427 individuals/ha, of which 40% were Acosnzium cardena~sii (after first mention of

a species I will refer to it by its generic name; for a complete list of scientific names see Table 2-

1). The company used these same logging techniques in the "intensive management" plot to

harvest a mean of 8.1 trees/ha (8.2 m3/ha). While harvesting the "intensive management plot,"

the skidder drivers mechanically scarified the soil surface in an average of 0.6 felling gaps/ha

(mean area = 50 m2/ha) where there was no existing regeneration of commercial timber species.

After logging of the intensive management plot, future crop trees (i.e., well-formed trees of

commercial species 10-40 cm DBH) were liberated from liana cover (by slashing the lianas with

a machete; 21 trees/ha) and liberated from competition from nearby non-commercial trees (by

poison girdling; 1.7 trees/ha). These last two treatments for enhancing the growth of future crop

trees were not applied in the normal logging plot. In the unlogged forest plot, the mean density of

trees > 10 cm DBH was 432 individuals/ha, of which 3 8% were Acosnzium (Table 2-1). The total

basal area of trees >10 cm DBH in the unlogged plot averaged only 19.6 m2/ha and 3 8% of the

trees were liana infested (18% severely so).

Canopy openness in the early dry season (May 2003), as measured with a spherical

densiometer (Lemmon 1957) held 1 m above the ground at 140 equally spaced points in the 10

ha permanent plots, averaged 8% in the 20 ha control (unlogged) plot, 13% in the plot subj ected

to normal selective logging in November 2002, and 14% in the plot that was intensively

harvested and silviculturally treated also in November 2002 (Figure 2-2). I remeasured canopy

openness at the same points in March 2006 and found increases in canopy cover in both of the

treated plots, but not in the control plot (Figure 2-2).









Experimental Design and Data Analysis

Response of regeneration to silvicultural treatment intensity

Plants 5-100 cm tall of 22 species of subcanopy, canopy, and emergent tree species (Table

2-2) that originated from seeds or sprouts were monitored in subplots in three of the 20-ha (400 x

500 m) LTSRP plots. In the central 10 ha (400 x 250 m) of each plot, I located 144 pairs of 2 x 1

m subplots separated by 2 m at 25 m intervals (data from the paired plots were subsequently

combined). Each seedling was marked, mapped, and measured for height at regular intervals of

one year for 3 y. The site of each plot pair was categorized in one of the following microsites:

undisturbed forest (includes natural canopy gaps and high stature forest); logging road; and,

logging gap. Undisturbed forests were categorized as those sites in which the structure of the

forest did not change during logging. I initially separated skid trails from primary and secondary

logging roads, but because of small sample sizes, these sites are combined into a category

referred to as log extraction paths. Logging gaps were sites where the canopy was opened during

tree felling and log extraction. Based on the literature (Whitmore 1998, Mostacedo and

Fredericksen 1999, Pinard et al. 1999) and field observations, each tree species was placed in one

of the following ecological groups: light-demanding pioneer; long-lived pioneer; partially shade

tolerant; or, shade tolerant.

I compared seedling abundances among the harvesting treatment plots and microsites using

repeated measures ANOVA (Scheiner and Gurevitch 2001). Measurement dates were considered

to be a within-subj ect factor, while harvesting treatment and microsites were considered as

between-subj ect factors.

Seedling recruitment rates (R) were calculated using the compound interest equation

(McCune and Cottam 1985):










R = (1 +Bx)'lx 1 (2-1)

where Bx is the birth rate in the period x calculated for each year and for each m2 Of

ground area. As recruitment rate I calculated the number of new seedlings of each species in

each harvesting treatment m2/er

Mortality rate (M)1 expressed as percentage per year was calculated as (Primack et al. 1985,

Sheil et al. 1995)

M~ = 1- [1- (No N,)l /No]"' (2-2)

where No is the number of living seedlings at time 0, N, is the number of living seedlings at

time N1, and t is the time period between No and N Mortality rates were calculated by species

and harvesting treatment.

I calculated the relative growth rate (RGR) as

(Ln(H, )- Ln(Ho))
RGR = (2-3)
t, to

where, Ho is the seedling at the initial measurement, H; is the height at the second

measurement, to is the initial time, and tl is the time of the second measurement (Hoffmann and

Poorter 2002). I compared RGR of harvesting treatments, microsites and species using repeated

measures ANOVA. For all analyses I used SPSS Version 12.0. For each factor, I also conducted

Bonferoni or LSD post hoc comparisons at the 95% confidence level.

Factors affecting seedling establishment and growth

Tree species used in the following experimental studies were selected on the basis of seed

availability in May-September 2005 when I collected seeds from 3-10 trees per species. Of the

six species used in the experiment, 4 are wind-dispersed, 2 are dispersed by animals, 2 are









characterized as light-demanding, and 4 are considered shade tolerant (Table 2-1). Brief

descriptions of the species follow:

Amburana cearensis (Allemio) A. C. Sm. usually produces one 0.49 g dry weight winged

seed per dehiscent legume. Aspidosperma rigidum Rusby produces elliptical seeds that weigh

0.24 g and are surrounded by a wing. Ceiba samnauma (Mart.) K. Schum. produces 0.17 g seeds

that are covered by fine, cotton-like fibers that aid in wind dispersal. Pterogyne nitens Tul.

produces single-seeded wind-dispersed samaras with 0.16 g seeds. Copaifera chodatiana Hassl.

seeds are red, weigh 0.33 g, are covered with a white aril, and are animal-dispersed. Hymenaea

courbaril L. produces round, large (3.9 g), and animal-dispersed seeds in indehiscent legumes.

The experiment used a split-plot design (Montgomery 2001) in which ground bromeliad

cover, water availability, and seed and seedling predation were experimentally manipulated

(Figure 2-3). The 2 m2 experimental plots were comprised of the two 1 m2 Subplots which were

replicated 40 times in 10 blocks separated by at least 40 m. The experiment commenced in July

2005 and was monitored until December 2006.

The terrestrial bromeliad manipulated in this study is Pseudananas~ddd~~~ddd~~~dd sagenarius, locally

known as "garabata." Rosettes of this clonal species are about 1 m high and cover 1m2 each.

Their colonies typically cover 15-40% of the ground in Chiquitano dry forest in clumps up to

2000 m2. This species is more common in mature than in young stands (Kennard et al. 2002) and

its abundance is reduced by fire (Fredericksen et al. 1999).

The water availability treatment involved either irrigating or withholding water from

plots by adding water or shielding plants from rainfall during what is typically the transition

between the dry and rainy seasons (late September). Rainfall inputs directly into the drought

treatment plots were prevented by covering them with 2.0 x 1.5 m roofs of transparent plastic.









The rain shelters were left for the first 3 mo of what is normally the dry season. The irrigated

plots received natural rainfall plus 10 liters of water every 15 days (=20 mm/mo) starting on 21

September 2005 and finishing in at the end of November when rainfall were more frequent. Soil

water tensions at 0-10 cm depth in one droughted and one irrigated plot in each of four blocks

were monitored hourly with Watermark@ soil moisture sensors attached to data loggers; rainfall

inputs were monitored adj acent to the four irrigated plots (Figure 2-1). Droughted plots

simulated years with prolonged dry seasons whereas the irrigated plots simulated wet years with

short dry seasons.

To exclude seed and seedling predators, half of the plots were surrounded by 60 cm high,

1-mm mesh wire netting fastened to the ground. The exclosures were erected before the seeds

were sown and were maintained for the duration (15 mo) of the experiment. In each treatment

plot, I randomly sowed seeds of all tree species described above (Table 2-1). Each seedling

establishing from a planted seed was marked, mapped, and its height measured five times

between September 2005 and December 2006.

Treatment effects on seedling abundance were tested using repeated measures ANOVA for

a split plot design (Scheiner and Gurevitch 2001) with bromeliads (present or absent) as the main

factor, and irrigation, predator exclosure, and time as subplot factors. Seedling heights (RGR) for

each time period were compared between treatments for each factor using t-test with sequential

Bonferroni corrections (Sankoh et al. 1997). Due to seedling mortality over the study, the sample

sizes varied so I could not run repeated measures ANOVA on the RGR data. I ran the analysis

for all 6 species combined and for each species separately.









Results

Response of Regeneration to Management Intensities

Three years after logging and silvicultural treatment, seedling densities varied with

management intensities and among microsites. Although 22 species are included in Table 2-1,

only data from the most abundant 11 species are presented. Densities were higher in the control

than in the harvested areas, but did not differ between the normal harvesting and the intensive

management plots (Figure 2-4). However, seedling densities without Acosmium, was higher in

normal harvesting and lower in the control plot (Figure 2-4). This pattern was maintained five

years after logging for all species combined, but for nine of the 11 species evaluated, there were

treatment differences and temporal trends (Table 2-2). For example, Copaifera seedling density

increased with increasing management intensity. In contrast, Machaerium acutifolium Vogel

seedling densities did not differ between the two logged plots but were significantly higher in the

normal logging plot than in the control plot. In contrast, seedling densities were lower in the

intensive management plot than in the normal logging or control plot for four Acosmium,

Phyllostylon rhamnnoides (J. Poiss.) Taub., and Machaerium scleroxylon Tul.

Seedling densities varied markedly among microsites. Overall, undisturbed sites had the

highest seedling density (Table 2-3). This pattern was observed at the species level for

Acosmium, Caesalpinia pluviosa,, M scleroxylon and Phyllostylon. In contrast, Centrolobium

microchaete seedling densities were higher on logging roads than in other microsites. The other

nine species had statistically similar seedling densities among microsites (Table 2-3).

Overall, seedling mortality rates were higher in undisturbed areas than in harvested areas

and the mortality rate was higher in 2004 than in 2005 (Figure 2-5). In 2004, 47% and 66% of

the seedlings died in the normal and intensive management plot, respectively. In 2005, mortality

was still higher in undisturbed control plots, but the ratio was reduced (Table 2-4). At species









level, mortality rates were generally higher in undisturbed control plots than in either of the

logged plots. On average, the highest (22-26%/y) mortality rates were for Sweetia f~uticosa

Spreng., Caesalpinia, and M~ scleroxylon. At the other end of the spectrum, Gallesia integrifolia

(Spreng.) Harms and Piptadenia~~ttt~~~~ttt~~~ viridifolia (Kunth) Benth, suffered no mortality (Table 2-4).

Over all species, the recruitment of new seedlings was 0.8 seedlings /m2 /y. Recruitment

was three times higher in undisturbed areas than harvested areas (Figure 2-5) and marginally

higher (0.91 seedlings /m2 /y) in 2004 than in 2005 (0.63 seedlings /m2 /y). Acosnzium had the

highest recruitment rate (0.26 seedlings/m2/y), followed by Phyllostylon (0.13 seedlings/m2/y

and M~ acutifolium (0.08 seedlings/m2/y). Acosnzium and M~ acutifolium recruited mostly in

undisturbed forest, Phyllostylon recruits were common in the normally logged plot, and

Pterogyne and Aspidospernza recruited most new seedlings in the intensive management plot

(Table 2-4).

Factors Affecting Seedling Growth and Establishment

Seedling growth

Relative growth rates of seedlings differed among the three logging microsites either in

2004 (F=4.19, P=0.01) or in 2005 (F=10.76, P=<0.0001). In both years, RGRs were higher on

loging roads and logging gaps than in undisturbed microsites. RGRs were higher in 2004 than in

2005 (F=5.64, P=0.018) and were higher for long-lived pioneer species RGRs than partial shade

tolerant and shade tolerant species (F=15.63, P=<0.0001).

RGRs differed among years, species, and microsites following patterns that were not

consistent. For example, the RGRs of Acosnzium, Aspidospernza, Copaifera, and Sweetia were

higher in 2004 than in 2005 (F=7.74, P=0.005). For Aspidospernza and Caesalpinia, RGRs did

not differ among undisturbed sites, logging roads, and logging gaps. Among microsites, RGRs

for M acutifolium were higher in logging roads and logging gaps than in undisturbed microsites










in 2004 but not in 2005. RGRs of Acosnzium and Sweetia were similar among microsites in 2004

whereas in 2005, seedlings grew faster in logging roads and logging gaps than in undisturbed

microsites.

Monthly RGRs of seedlings in the experimental plots were similar in bromeliad-free and

bromeliad-infested plots at different times (tl=0.14, P1=0.89; t2=2.01, P2=0.05; t3=0.48, P3=0.63;

t4=0.04, P4=0.39). Similarly, RGRs were did not differ in the droughted and irrigated plots

(tl=0.89, P1=0.37; t2=0.87, P2=0.39; t3=0.93, P3=0.35; t4=0.88, P4=0.38) or between the exclosure

and non-exclosure plots (tl=0.01, P1=0.99; t2=0.12, P2=0.90; t3=0.40, P3=0.69; t4=0.36, P4=0.72;

Figure 2-10). At the species level, RGRs were also similar in bromeliad-free and bromeliad-

infested plots, droughted and irrigated plots, and exclosure and non-exclosure plots (Table 2-6,

2-7, 2-8).

Seedling establishment

In general, tree seedling establishment was suppressed by bromeliads, but responses to

bromeliad cover varied among species (Figure 2-6). Based on the six species for which there are

sufficient data, four had similar numbers of seedlings in bromeliad patches and bromeliad-free

areas. For the other two species (Hynzenaea and Pterogyne), seedling abundances were lower in

bromeliad covered plots (Table 2-5, Figure 2-7).

The effects of the drought and irrigation treatments varied over time (Figure 2-6).

Seedlings were initially about 150% more abundant in the irrigated than in the droughted plots

(Figure 2-8), but this difference diminished over time. This pattern was consistent among species

except for Aspidospernza, which had more seedlings in the droughted than in the irrigated plots

(Table 2-5).

Excluding seed and seedling predators generally resulted in higher seedling densities

(Figure 2-6), with some variation among species (Figure 2-9). Anaburana, Copaifera, and










Hymenaea seedlings were more common within the excloures than in the open-access plots

(Table 2-5). In contrast, the exclosure treatment had no effect on seedling densities of

Aspidosperma, Ceiba, and Pterogyne.

There were some significant interactions among factors both overall and at the species

level. In particular, irrigated plots with bromeliads from which predators were excluded had

higher seedling densities than plots that were irrigated and open to seed and seedling predators

(Table 2-5).

At the species level there were significant interactions among treatments only for Ceiba

and Copaifera. For Ceiba in the bromeliad covered plots, irrigation and predator exclosure

increased seedling densities relative to the droughted and open-access plots. For Copaifera in the

bromeliad-infested plots, irrigation and predator exclosure resulted in higher seedling densities.

In the bromeliad-free plots, drought coupled with predator exclosure resulted in higher seedling

densities than drought alone.

Discussion

Logging Effects on Seedlings Dynamics

Logging usually affects seedling densities due to direct effects and creation of microsites

that differ in environmental conditions, especially light intensities (Beaudet and Messier 2002,

van Rheenen et al. 2004). In most tropical forests, seedling densities of light-demanding species

increase on open microsites. In this study, however, overall seedling densities were higher in

undisturbed forest than in selectively logged areas (Figure 2-2). Similarly, total seedling densities

were higher on undisturbed microsites compared to microsites created by logging. In contrast, at

a site only 40 km from INPA, with a similar forest type and rainfall regime Fredericksen and

Mostacedo (2000) reported that seedlings of commercial timber species increased in abundance

on log landings, roads, and logging gaps.









At the species level, the maj ority of species (64%) had similar seedling densities among

microsites; only three shade-tolerant species had higher seedling densities in undisturbed

microsites. One light-demanding species, Centrolobium, had higher seedling densities on skid

trails and log landings compared to other microsites.

There are several potential explanations for observed differences between the results of this

study and those of Fredericksen and Mostacedo (2000). The forest canopy of Las Trancas where

the latter conducted their work is dominated by Anadenanthera, a light-demanding species,

whereas INPA Parket is dominated by Acosnzium, which is more shade tolerant. Results without

Acosnzium show that harvested plots have higher seedling densities than control plot (Figure 2-

4). The simple difference in dominants might account for some of the overall differences

observed between the sites. Another difference between the two studies is that Anadenanthera

seedlings emerging from seeds dispersed immediately before logging were extremely abundant

at Las Trancas (100-200 individuals/m2; perSonal observation). In contrast, seedlings of

Acosnzium were never as abundant at INPA and those that were present were apparently the

result of several years of recruitment.

Open microsites created by logging may be disadvantageous for seedling recruitment in

INPA Parket because such sites are extremely hot and dry, especially during the dry season. On

the other hand, it is possible that even though there were disturbances associated with fallen

trees, roads, and skid trails, the canopy could still be too closed at INPA to promote abundant

seedling recruitment (Jackson et al. 2002). In addition, the densities of understory plants and

other competitors might remain high in gaps, thus suppressing seedling recruitment in disturbed

areas, especially of shade-intolerant species (Beckage et al. 2000, Schnitzer et al. 2000). On the

log extraction paths, soil compaction might negatively affect seedling establishment and growth.









Finally, several species in this study are shade-tolerant, including the dominant (Acosmium), and

do not require disturbed areas for regeneration.

Seedling recruitment rates were 70% lower in the logged plots than in the unlogged control

plot and recruitment decreased with management intensity. Because of increases in light

availability associated with logging, I expected more recruitment in logged plots than in

unlogged plots. One explanation for this result could be that logging decreased seed-tree

densities, which limited seed inputs (Forget et al. 2001, Makana and Thomas 2004, Grogan and

Galvao 2006). Similarly, in French Guiana, Forget et al. (2001) reported that for all but one

species, recruitment was not favored by logging. The importance of retention of seed trees was

shown by Grogan and Galvao (2006), who reported that mahogany (Swietenia macrophylla) seed

production near gaps was important for recruitment. In addition, even if seed production is not

limiting, logging may create microsites that are not appropriate for seed germination and

seedling growth due to soil compaction (Pinard et al. 1996) or other factors. Site preparation

(e.g., soil scarification) in logging gaps and skid trails can enhance regeneration, as shown by

Pinard et al. (1996) in a dipterocarp forest in Malaysia, but such treatments do not always have

beneficial effects (Heuberger et al. 2002).

Seedling mortality rates were nearly 50% lower in the logged plots than in the unlogged

plot, which was the opposite of what I expected. Several studies report that seedling mortality

usually increases as harvesting intensity increases (Chapman and Chapman 1997, Saenz and

Guariguata 2001), but the logging intensities in my plots were all low. Nevertheless, mortality of

some plant species decreases as light intensity increases. For example, in a tropical rain forest in

Costa Rica, Kobe (1999) found that mortality rates of three of four canopy species monitored

decreased as light levels increased in the understory.









There was considerable interspecific variation in both recruitment and mortality rates. For

example, for three tree species (Acosnzium, Aspidospernza, and Copaifera) that are in the same

shade tolerant guild the percentage of recruitment ranged from 7.5-15.5 %/y. The mortality rates

for the same species ranged from 0.019-0.25 5 individuals/m2/y. Acosnzium, because of its

dominance, drives the overall patterns of recruitment and mortality.

Apparently, while recruitment rates are reduced by disturbance, the seedlings that do

establish grow sufficiently during the rainy season to survive the subsequent dry season. In

contrast, in the relatively closed conditions of the control plot, few of the abundant new recruits

survive the drought stress of their first dry season. Similarly, seedlings of a light-demanding

species (Anadenanthera) established abundantly, but died rapidly in the control plot.

Temporal changes in recruitment and mortality are related to variation in seed production

and rainfall regime (Kitajima and Fenner 2000, Khurana and Singh 2001). In this study, seed

production, seedling recruitment, and seedling mortality rates were all higher during 2003-2004,

a dry year, than during 2004-2005, a relatively wet year with a short dry season.

Within the logged plots, seedling RGRs were typically higher in disturbed microsites, but

there was some variation among species. Over the 2 y monitoring period, seedlings grew faster

on log extraction paths than undisturbed microsites. In a similar study in the same area but using

large transects, seedlings in skid trails and logging gaps grew faster than seedlings in undisturbed

forest (van Andel 2005). Contrary to my expectations, Acosnzium, a shade-tolerant species, had

the highest RGR on log extraction paths while RGRs of Anadenanthera, a light-demanding

species, not vary among microsites. Overall it appears that disturbed microsites are favorable for

seedlings, but this pattern seems to be driven by Acosnzium, the most abundant species. For most

of the species it seems that, in contrast to wetter tropical forests, RGRs of tropical dry forest









seedlings are not governed primarily by light availability and have a different relationship to

disturbance than wet and moist forest tree species (Khurana and Singh 2001).

Factors Affecting Seedling Establishment and Growth

Soil moisture availability is a key factor influencing seedling growth in dry forests

(Khurana and Singh 2001). In my study, seedlings that lost their leaves during the dry season

(e.g., Amburana and Aspidosperma) grew slowly, while the evergreen species (e.g., Copaifera

and Pterogyne) grew faster. I also found that bromeliad cover significantly reduced seedling

growth rates.

Ground bromeliads interfered with the establishment and growth of tree seedlings in this

study, as has been described for another forest in Bolivia (Fredericksen et al. 1999), as well as on

Barro Colorado Island in Panama (Brokaw 1983). As in those other two studies, I also observed

substantial interspecific variation in seedling responses to bromeliads. The observation that the

irrigation treatment eliminated the generally negative effect of bromeliads on seedling survival

and growth suggests that the rain-trapping effect of bromeliad rosettes, coupled with any water

they draw from the soil, is the cause of their deleterious effect. Perhaps the species that were not

sensitive to the presence of bromeliads (Amburana, Ceiba, Copaifera, and Pterogyne) are more

drought-tolerant than the other species.

I expected that experimentally shortening the dry season by irrigation would increase the

establishment and growth of tree seedlings, while exacerbating dry season drought would have

the opposite response. Instead, what I found that the principal impact of the drought treatments

was the timing of seed germination, but not all species were affected. Seeds in the droughted

plots did not germinate until after I removed the roofs and allowed rain to fall on the soil;

seedlings then reached the same densities as in the irrigated plots. A similar study in mahogany









in a dry forest in Guanacaste, Costa Rica reported the same effect of water availability on

seedling density (Gerhardt 1996b). Similarly, RGR were not affected by drought stress.

The tendency for species overall (except Aspidospernza) was higher densities in the

irrigated than in the droughted plots at the time of germination. This tendency suggests that

longer rainy seasons, such as those occurring during El Nifio years, can have dramatic effects on

seedling establishment (Gilbert et al. 2001, Engelbrecht and Kursar 2003). The same pattern of

increased mortality during dry years was revealed by the mortality of naturally established

seedlings during this study (Figure 2-5).

Due to a variety of morphological, physiological, and phenological traits, tree species

differ in the susceptibility of their seedlings to drought stress (Table 2-4; (Reader et al. 1993,

Khurana and Singh 2001, McLaren and McDonald 2003). For example, Anaburana seedlings

reduce their moisture requirements by being dry season deciduous, leaf areas ofAspidospernza

seedlings decline substantially during the dry season, and Acosnzium is noteworthy for its deep

root system (Wright 1991, Engelbrecht and Kursar 2003).

Seed or seedling predation seems to be a very important factor for seedling establishment

in tropical dry forests (Janzen 1971, Hulme 1996). I found that plots with exclosures had higher

seedlings densities than open plots. Exclosures prevented seedling herbivory, but not all types of

seed predation because insects and other small seed predators were not excluded (Grol 2005). In

tropical dry forests, herbivory has the most impact on tree regeneration when seedlings are

emerging, not after roots and stems are lignified (Kitajima and Augspurger 1989, Lucas et al.

2000). Once browsed, established seedlings usually resprout, but only lignified seedlings

survive. In some cases, herbivory damage could be superficial and not dramatically impact the

growth and survival of seedlings (personal observation).









There were some species in which exclosure treatments had an effect on established

seedlings, while for others there were similar. For example, Hymenaea, a species with a large

seed and seedlings larger than Ceiba, had a higher established seedling caused by the exclosure

plots. Several studies support the hypothesis that large seeds are usually most predated (Blate et

al. 1998, Dalling and Hubbell 2002), but see Moles et al. (2003). Likewise, Aspidosperma and

Pterogyne have seeds that probably were predated mostly by small insects (i.e., bruchid beetles)

which were not deterred by the exclosures.

Conclusions

Five years after selective logging and silvicultural treatments, overall tree seedling

densities were higher in the control plot than in the two logged plots. It is important to point out

that this result was heavily influenced by the reduced seedling densities in the logged plots of the

dominant species, the shade-tolerant Acosmium. Other species, which were less common

everywhere (e.g., Centrolobium, Copaifera, and Machaerium cf: acutifolium), actually increased

in response to forest management activities, and many other species were not affected. Within

the logged plots, undisturbed microsites had higher seedling densities except for Centrolobium, a

light-demanding species that sprouts readily from damaged lateral roots, which was more

abundant on log extraction paths.

Seedling recruitment rates were higher in the unlogged control plot and on undisturbed

microsites in the logged plots, but seedling in these areas also suffered higher mortality rates than

seedlings in disturbed areas. The higher turnover rates of seedlings under closed canopy

conditions coupled with their observed lower growth rates suggests that rapid growth under open

conditions during the rainy season is critical for seedling survival during the subsequent dry

season. This result is somewhat surprising given that, even undisturbed forest in my study area,

the canopy is 30-40% open after leaf fall.









Mortality rates of naturally established seedlings varied greatly among species. Seven of

22 species suffered no mortality during the 2-y monitoring period, whereas relatively high

mortality rates were observed for Caesalpinia (26%/y), Sweetia (25%/y), and Machaerium

scleroxylon (22%/y). Mortality rates of these species in the control and logged plots followed

similar patterns.

Bromeliad competition and seed/seedling predators greatly reduced seedling recruitment in

this tropical dry forest. Experimentally augmented soil moisture also increased seedling

establishment, but only in interaction with bromeliad removal or predator exclosure.

Experimental droughting delayed germination but did not influence seedling densities once the

treatment terminated and rainfall was allowed into the plots. Despite these general trends, species

varied in their sensitivities to bromeliads, drought stress, and predators.

Finally, seedling growth was not promoted by experimentally lengthening the rainy season.

Nevertheless, the larger seedlings that developed during the extended rainy season were more

likely to survive the subsequent dry season. Given that global change models consistently predict

that the study region will receive less rainfall and suffer longer dry seasons in the future (e.g.,

Mayle et al. 2007), regeneration failures are likely to become more common.











Table 2-1. Spatial distributions, crown position, ecological group, geographical range, tree densities and basal area of timber species
in a Bolivian tropical dry forest.


Stem Density
(#/ha >10 cm
DBH)
159.25
0.005
13.67
12.92
0.08
15.17
0.67
11.42
16.67
5.33
2.83
0.17
27.17
4.25
4.42
0.17
0.25
12.8
1.83
2.75
142.90
434.00


Spatial Crown
Distribution Position


Ecological
Group

TS
PS
L
PS
L
PS
L
L
L
TS
L
PS
LL
PS
TS
LL
LL
PS
LL
PS


Basal Area
(nr/ha)

6.89
0.006
2.00
0.40
0.07
1.18
0.07
0.55
0.82
0.51
0.05
0.06
0.34
0.68
0.63
0.01
0.144
0.22
0.27
0.04
4.67
19.60


Scientific Name

4cosiniuin cardenasii
dinburana cearensis*
4nadenanthera inacrocarpa
4spidosperina rigiduin.*
4stroniuin urundeuva
Caesalpinia pluviosa
Ceiba sainauina*
Centrolobiuin iicrochaete
Chorisia speciosa
Copaifera chodatiana*
Cordia alliodora
Hyinenaea courbaril*
Machaerium acutifolium
Machaerium scleroxvion
Phyllostvilon rhainnoides
Pterogvne nitens*
Schinopsis brasiliensis
Sweetia fruticosa
Tabebuia .,~, ruagest..w
Zeyheria tuberculosa
Others
Total
Crown position: (EM)
Ecological group: (L) =


Family

Fabaceae
Fabaceae
Mimosaceae
Apocynaceae
Anacardiaceae
Caesalpinaceae
Bombacaeae
Fabaceae
Bombacaceae
Caesalpinaceae
Boraginaceae
Caesalpinaceae
Fabaceae
Fabaceae
Rhamnaceae
Caesalpinaceae
Anacardiaceae
Fabaceae
Bignoniaceae
Bignoniaceae


Deciduous Range


Homogeneous
Random
Homogeneous
Clumped
Clumped
Homogeneous
Random
Clumped
Random
Random
Random
Random
Homogeneous
Random
Clumpled
Homogeneous
Random
Random
Homogeneous
Random


None
May-Aug
Jun-Oct
Jul-Sep
Jul-Sep
Aug-Sep
Jul-Aug
Jun-Oct
Jul-Aug
None
Jun-Aug
None
None
Jul-Sep
None
None
Aug-Sep
Jul-Sep
Jul-Sep
Oct-Nov


emergent, (CA) = canopy, (SC)
light-demanding pioneer, (LL) =


= sub-canopy.
Long-lived pioneer, (PS)


partially shade tolerant, (TS) = shade tolerant.


Geographical range: (R) = restricted, (I) = intermediate, (W) = widespread. Restricted species are those that are found only in one type
of forest and are limited to the Chiquitano dry forest in Bolivia. Intermediate species are those that are found in two or three
forest types but are either restricted to Bolivia or are only rarely found in neighboring countries. Widespread species are found in
several forest types and in other countries.
Species marked with asterisks were used in the seedling survival experiment.











Table 2-2. Means of seedling densities in an unharvested control plot, a plot subjected to normal timber harvesting, and a plot
subj ected to more intensive harvesting followed by silvicultural treatments. Subplots within the main treatment plots are
treated as replicates. Different letters indicate statistical differences (P<0.05) among harvesting treatments as indicated by
repeated measures ANOVAs and LSD post hoc tests.


Intensive
Management


Mean
Square
64.89
0.67
1.11


Scientific Name


Unharvested


Normal Logging


(0.09)b
(0.01)b
(0.02)a


Acosnzium cardenasii
Anadenanthera nzacrocalpa
Aspidospernza rigidunt
Caesapinia pheviosa
Centrolobium naicrochaete


1.42
0.09
0.01

0.12


(0.09)c
(0.01)b
(0.02)b


1.03
0.06
0.06


0.63
0.01
0.11


(0.09)a
(0.01)a
(0.02)a


18.90
8.15
5.33

11.50
1.89

2.31

12.33
22.17
48.46
64.74


0.0001
0.0001
0.005

0.001
0.170

0.100

0.0001
0.0001
0.0001
0.0001


0.09 (0.02)b
0.02 (0.01)a

0.01 (0.003)a


0.02 (0.02)a
0.01 (0.01)a

0.002 (0.003)a


1.17
0.03

0.01


Chorisia speciosa


0.005 (0.003)a


(0.04)b
(0.13)c
(0.01)b
(0.02)b


(0.04)b
(0.14)b
(0.01)a
(0.02)a


(0.04)a
(0.13)a
(0.01)a
(0.02)a


Copaifera chodatiana
Machaeriunt acutifolium
Machaerium scleroxylon
Phyllostylon rha~nnoides


0.10
0.04
0.12
0.31


0.05
1.31
0.004
0.00


0.32
0.72
0.001
0.01


8.40
166.75
1.85
12.68


(0.01)a 0.02 (0.01)b 0.05 (0.01)a


Sweetia fiuticosa 0.03


0.07 2.53 0.080










Table 2-3. Mean seedling densities (#/m2) for 3 years (2003-2005) of commercial tree species in microsites created by logging in a
Chiquitano dry forest. Sample size for each microsite type is noted within brackets; standard errors of the means are noted
in parenthesis. Seedling densities in the different microsites were compared with repeated measures ANOVA with
microsites as the between-subject factor and time as the within-subj ect factor. Different letters indicate different
significance between treatments using pairwise Least Significant Difference (LSD) tests.
Scientific Name Logging Gap [100] Log Extraction Path [54] Undisturbed [337] Mean Square F P

4cosiniuin cardenasii 0.701 (0.103)b 0.389 (0.14(,) 0.945 (0.056)a 24.92 7.88 <0.0001

4nadenanthera inacrocarpa 0.018 (0.015) 0.049 (0.021) 0.050 (0.008) 0.119 1.71 0.18

4spidosperina rigiduin 0.044 (0.024) 0.026 (0.033) 0.056 (0.013) 0.07 0.39 0.68

Caesalpinia pluviosa 0.024 (0.015)b 0.037 (0.020)b 0.073 (0.008)a 0.32 4.71 0.009

Ceiba sainauina 0.000 (0.001) 0.000 (0.001) 0.001 (0.000) 0.0001 0.68 0.50

Centrolobiwn inicrochaete 0.008 (0.005)b 0.022 (0.007)a 0.005 (0.003)b 0.02 2.86 0.05

Chorisia speciosa 0.004 (0.003) 0.002 (0.004) 0.004 (0.002) 0.001 0.23 0.79

Copaifera chodatiana 0.092 (0.045) 0.086 (0.061) 0.146 (0.024) 0.51 0.85 0.43

Gallesia integrifolia 0.000 (0.003)b 0.000 (0.004)b 0.007 (0.002)a 0.009 3.69 0.03

Machaerium acutifolium 0.609 (0.133) 0.668 (0.181) 0.519 (0.072) 2.13 0.40 0.67

Machaerium scleroxvion 0.008 (0.011)b 0.000 (0.015)b 0.048 (0.006)a 0.29 7.89 <0.0001

Phyllostylon rhainnoides 0.007 (0.025) 0.000 (0.035) 0.122 (0.014) 2.203 11.34 <0.0001

Pterogvne nitens 0.000 (0.004) 0.006 (0.005) 0.005 (0.002) 0.003 0.91 0.41

Sweetia fruticosa 0.015 (0.009) 0.011 (0.012) 0.031 (0.005) 0.05 2.24 0.11

Overall Species 1.543 (0.180)b 1.302 (0.245)b 2.019 (0.098)a 52.87 5.44 0.005











Table 2-4. Establishment and mortality rates of seedlings of commercial tree species monitored over a 3 y period in a control plot and
plots subj ected to two harvesting intensities (N=144 subplots/treatment plot).


Scientific Name Mortality Rate (% of Seedlings / y) Number of New Seedlings (#/m2 /y
Normal Intensive Normal Intensive
Undisturbed Logging Management Overall Undisturbed Logging Management Overall

Acosmium cardenasii 17.0 11.7 9.8 12.8 0.489 0.176 0.101 0.255
Anadenanthera macrocarpa 21.6 5.4 18.8 15.2 0.106 0.079 0.003 0.062
Aspidosperma rigidum 14.5 2.5 5.5 7.5 0.007 0.013 0.037 0.019
Caesalpinia pluviosa 35.8 23.2 18.8 25.9 0.117 0.064 0.011 0.064
Ceiba samauma 0.0 0.0 0.0 0.0 0.001 0.001 0.001 0.001
Centrolobium microchaete 0.0 10.7 43.8 18.2 0.000 0.006 0.000 0.002
Chorisia speciosa 0.0 0.0 0.0 0.0 0.004 0.004 0.000 0.003
Copaifera chodatiana 24.1 8.2 14.3 15.5 0.125 0.015 0.050 0.063
Gallesia integrifolia 0.0 0.0 0.0 0.0 0.034 0.000 0.000 0.011
Machaerium c~f acutifolium 16.1 12.4 12.6 13.7 0.003 0.138 0.106 0.082
Machaerium scleroxylon 34.7 30.8 0.0 21.8 0.157 0.012 0.001 0.056
Phyllostylon rhamnoides 19.5 10.0 0.0 9.8 0.381 0.004 0.011 0.132
Pterogyne nitens 43.8 0.0 0.0 14.6 0.000 0.001 0.022 0.008
Sweetia fr~uticosa 41.4 24.8 8.3 24.8 0.020 0.010 0.004 0.011


Overall


22.0 12.4 11.2 15.2 1.446


0.523 0.354 0.775












Table 2-5. Results of repeated measure analysis of variance for a split plot design run for
seedling density for all species combined or six timber tree species analyzed
separately. Double asterisks indicate between subj ect factors while the unmarked
variables are within subj ect factors.
Type III
Sum of Degrees of Mean
Source Squares Freedom Square F P-value

All species
Time (Ti) 44.01 4 11.00 25.88 <0.0001
Bromeliad (Br)(**) 163.20 1 163.20 9.99 0.005
Time*Bromeliad (Br) 4.95 4 1.24 2.90 0.03
Irrigation (Ir) 27.83 1 27.83 2.74 0.11
Irrigation*Bromeliad 7.16 1 7.16 0.70 0.41
Exclosure (Ex) 79.66 1 79.66 13.01 0.002
Ex*Br 18.71 1 18.71 3.06 0.09
Ti*Ir 13.99 4 3.49 10.81 <0.0001
Ti*Ir*Br 1.92 4 0.48 1.48 0.22
Ti*Ex 7.75 4 1.94 5.76 <0.0001
Ti*Ex*Br 1.43 4 0.36 1.06 0.38
Ir*Ex 11.73 1 11.73 1.31 0.27
Ir*Ex*Br 16.60 1 16.60 1.86 0.19
Ti*Ir*Ex 0.58 4 0.14 0.39 0.81
Ti*Ir*Ex*Br 0.35 4 0.09 0.24 0.92

4mburana cearensis
Time (Ti) 1.74 4 0.44 7.72 <0.0001
Bromeliad (Br)(**) 0.83 1 0.83 0.43 0.53
Time*Bromeliad (Br) 0.13 4 0.03 0.58 0.68
Irrigation (Ir) 3.01 1 3.01 1.87 0.20
Irrigation*Bromeliad 1.09 1 1.09 0.68 0.43
Exclosure (Ex) 4.50 1 4.50 2.59 0.14
Ex*Br 2.52 1 2.52 1.45 0.26
Ti*Ir 1.35 4 0.34 6.03 0.001
Ti*Ir*Br 0.08 4 0.02 0.36 0.84
Ti*Ex 0.55 4 0.14 2.39 0.07
Ti*Ex*Br 0.22 4 0.05 0.95 0.44
Ir*Ex 8.87 1 8.87 3.10 0.10
Ir*Ex*Br 0.09 1 0.09 0.03 0.86
Ti*Ir*Ex 1.03 4 0.26 3.65 0.01
Ti*Ir*Ex*Br 0.08 4 0.02 0.29 0.88

4spidosperma riaidwn
Time (Ti) 1.16 4 0.29 8.59 <0.0001
Bromeliad (Br)(**) 0.59 1 0.59 0.25 0.63
Time*Bromeliad (Br) 0.04 4 0.009 0.28 0.89
Irrigation (Ir) 5.37 1 5.37 1.78 0.23
Irrigation*Bromeliad 1.31 1 1.31 0.43 0.53
Exclosure (Ex) 0.19 1 0.19 0.10 0.76
Ex*Br 2.6 1 2.6 1.38 0.28










Table 2-5. Continued.
Type III
Sum of Degrees of Mean
Source Squares Freedom Square F P-value
Ti*Ir 0.32 4 0.08 1.95 0.13
Ti*Ir*Br 0.08 4 0.02 0.47 0.76
Ti*Ex 0.12 4 0.03 1.18 0.34
Ti*Ex*Br 0.13 4 0.03 1.24 0.32
Ir*Ex 1.50 1 1.50 0.87 0.39
Ir*Ex*Br 1.15 1 1.15 0.67 0.44
Ti*Ir*Ex 0.08 4 0.02 0.76 0.56
Ti*Ir*Ex*Br 0.08 4 0.02 0.76 0.56

Ceiba samauma
Time (Ti) 0.53 4 0.13 2.02 0.12
Bromeliad(Br)(**) 7.75 1 7.75 4.65 0.07
Time*Bromeliad (Br) 0.05 4 0.01 0.21 0.93
Irrigation (Ir) 1.00 1 1.00 0.72 0.43
Irrigation*Bromeliad 0.53 1 0.53 0.38 0.56
Exclosure (Ex) 0.001 1 0.001 0.004 0.95
Ex*Br 0.06 1 0.06 0.54 0.49
Ti*Ir 0.30 4 0.08 1.06 0.39
Ti*Ir*Br 0.03 4 0.007 0.10 0.98
Ti*Ex 0.12 4 0.03 0.91 0.47
Ti*Ex*Br 0.05 4 0.01 0.42 0.79
Ir*Ex 1.69 1 1.69 9.28 0.023
Ir*Ex*Br 0.79 1 0.79 4.35 0.082
Ti*Ir*Ex 0.13 4 0.03 0.94 0.46
Ti*Ir*Ex*Br 0.09 4 0.02 0.67 0.62

Copaifera chodatiana
Time (Ti) 4.1 4 1.03 15.09 <0.0001
Bromeliad (Br)(**) 2.96 1 2.96 1.49 0.24
Time*Bromeliad (Br) 0.16 4 0.04 0.60 0.66
Irrigation (Ir) 1.88 1 1.88 3.64 0.07
Irrigation*Bromeliad 0.06 1 0.06 0.12 0.73
Exclosure (Ex) 3.53 1 3.53 3.18 0.09
Ex*Br 0.03 1 0.03 0.03 0.87
Ti*Ir 0.73 4 0.18 3.59 0.01
Ti*Ir*Br 0.41 4 0.10 2.03 0.10
Ti*Ex 0.53 4 0.13 2.25 0.07
Ti*Ex*Br 0.13 4 0.03 0.56 0.69
Ir*Ex 0.56 1 0.56 0.76 0.39
Ir*Ex*Br 5.13 1 5.13 6.96 0.02
Ti*Ir*Ex 0.12 4 0.03 0.59 0.67
Ti*Ir*Ex*Br 0.31 4 0.08 1.46 0.22

Hymenaea courbaril
Time (Ti) 1.85 4 0.46 16.09 <0.0001
Bromeliad (Br)(**) 1.63 1 1.63 3.16 0.09
Time*Bromeliad (Br) 0.09 4 0.02 0.83 0.51










Table 2-5. Continued.
Type III
Sum of Degrees of Mean
Source Squares Freedom Square F P-value
Irrigation (Ir) 0.37 1 0.37 1.33 0.27
Irrigation*Bromeliad 1.13 1 1.13 0.47 0.51
Exclosure (Ex) 5.28 1 5.28 13.23 0.003
Ex*Br 0.26 1 0.26 0.65 0.44
Ti*Ir 0.30 4 0.07 1.96 0.11
Ti*Ir*Br 0.14 4 0.03 0.90 0.47
Ti*Ex 1.4 4 0.35 8.53 <0.0001
Ti*Ex*Br 0.08 4 0.02 0.49 0.74
Ir*Ex 0.08 1 0.08 0.45 0.52
Ir*Ex*Br 0.28 1 0.28 1.58 0.23
Ti*Ir*Ex 0.42 4 0.10 2.95 0.03
Ti*Ir*Ex*Br 0.18 4 0.05 1.29 0.28

Pteroevne nitens
Time (Ti) 1.09 4 0.27 6.68 <0.0001
Bromeliad (Br)(**) 2.12 1 2.12 7.74 0.02
Time*Bromeliad (Br) 0.39 4 0.09 2.36 0.07
Irrigation (Ir) 0.26 1 0.26 1.40 0.25
Irrigation*Bromeliad 0.71 1 0.71 4.03 0.07
Exclosure (Ex) 0.05 1 0.05 0.21 0.65
Ex*Br 0.18 1 0.18 0.75 0.41
Ti*Ir 0.62 4 0.16 4.65 0.003
Ti*Ir*Br 0.08 4 0.02 0.63 0.64
Ti*Ex 0.09 4 0.02 0.62 0.65
Ti*Ex*Br 0.14 4 0.03 0.98 0.43
Ir*Ex 0.13 1 0.13 0.71 0.42
Ir*Ex*Br 0.01 1 0.01 0.06 0.81
Ti*Ir*Ex 0.11 4 0.02 0.45 0.77
Ti*Ir*Ex*Br 0.06 4 0.02 0.45 0.51











Table 2-6. Mean relative height growth rates (+ 1SE) of seedlings of commercial tree species in
response to bromeliad cover removal evaluated in 4 times. Significant differences
between treatments were evaluated using t-tests at 95% of confidence level with
sequential Bonferroni corrections (PB = 0.0125).


Species

Amburana cearensis





Aspidosperma rigidum




Ceiba samauma





Copalifera chodatiana


Date


Bromeliad
Mean (SE)
0.032 (0.019)
0.046 (0.029)
-0.022 (0.012)
0.026 (0.009)

-0.009 (0.021)
-0.0004 (0.018)
0.0145 (0.02)
0.027 (0.006)

0.170 (0.026)
0.090 (0.046)
-0.038 (0.015)
0.022 (0.008)

0.038 (0.015)
0.033 (0.010)
-0.014 (0.009)
0.009 (0.007)

0.346574
0.026 (0.046)
0.009 (0.016)


0.084 (0.007)
0.230 (0.078)
0.024 (0.024)
-0.009 (0.013)


No Bromeliad
Mean (SE)
-0.025 (0.026)
0.063 (0.021)
-0.015 (0.010)
0.024 (0.008)

0.020 (0.019)
0.096 (0.027)
-0.003 (0.004)
0.014 (0.003)

0.080 (0.035)
0.103 (0.028)
-0.022 (0.008)
-0.010 (0.011)

0.029 (0.015)
0.056 (0.017)
0.005 (0.007)
0.018 (0.006)

0.141 (0.101)
0.028 (0.023)
-0.012 (0.019)
0.016 (0.010)

0.109 (0.058)
0.107 (0.027)
0.032 (0.014)
-0.036 (0.014)


t-test
1.57
0.43
0.37
0.14

0.89
2.22
1.57
1.84

1.02
0.20
0.93
1.65

0.36
0.99
1.44
0.76


P
0.13
0.67
0.72
0.89

0.38
0.03
0.13
0.08

0.33
0.84
0.37
0.14

0.72
0.33
0.15
0.45


JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06


Hymenaea courbaril


0.68 0.52
0.05 0.96
0.62 0.54


Pterogyne nitens


0.20
1.49
0.53
0.53


0.85
0.15
0.81
0.25










Table 2-7. Mean relative height growth rates (+ 1SE) of seedlings of commercial tree species in
irrigated and droughted plots evaluated 4 times. Significant differences between
treatments were determined using t-test at 95% of confidence level with sequential
Bonferroni corrections (PB = 0.0125).


Species


Irrigated
N Mean (SE)
25 -0.006 (0.021)
21 0.044 (0.018)
19 -0.024 (0.012)
15 0.025 (0.008)

7 -0.006 (0.066)
7 0.152 (0.072)
7 0.004 (0.017)
6 0.024 (0.008)

13 0.074 (0.026)
11 0.086 (0.033)
14 -0.029 (0.009)
6 -0.013 (0.009)

35 0.039 (0.011)
37 0.040 (0.013)
31 0.001 (0.008)
18 0.008 (0.006)

9 0.164 (0.092)
15 0.020 (0.023)
12 -0.001 (0.011)
1 0.026

10 0.107 (0.052)
10 0.085 (0.036)
8 0.025 (0.021)
3 -0.036 (0.029)


Droughted
N Mean (SE)
4 0.015 (0.035)
11 0.084 (0.035)
10 -0.002 (0.005)
8 0.024 (0.013)


Date


P
0.69
0.28
0.19
0.92

0.51
0.02
0.85
0.36

0.01
0.46
0.46
0.09

0.47
0.51
0.76
0.09


0.71
0.74


0.86
0.15
0.70
0.61


t-test
0.40
1.11
1.34
0.10

0.66
2.39
0.19
0.93

2.82
0.76
0.75
1.89

0.72
0.66
0.30
1.76


0.38
0.11


0.18
1.53
0.40
0.53


Amburana cearensis





Aspidosperma rigidum




Ceiba samauma





Copalifera chodatiana





Hymenaea courbaril




Pterogyne nitens


JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06


0.017 (0.081)
0.041 (0.014)
0.002 (0.004)
0.016 (0.003)

0.346
0.125 (0.037)
-0.017 (0.011)
0.019 (0.015)

0.022 (0.024)
0.056 (0.021)
-0.003 (0.009)
0.024 (0.007)


0.037 (0.042)
-0.012 (0.029)
0.005

0.077
0.165 (0.037)
0.035 (0.014)
-0.023 (0.009)










Table 2-8. Mean relative height growth rates (+ 1SE) of seedlings of commercial tree species in
response to mammal exclosure evaluated 4 times during 2006. Significant differences
between treatments were evaluated using t-test at 95% of confidence level with


sequential Bonferroni corrections (PB


0.0125).


Species


Exclosure
Mean (SE)
-0.011 (0.023)
0.059 (0.018)
-0.013 (0.007)
0.018 (0.007)

0.015 (0.018)
0.075 (0.034)
-0.002 (0.006)
0.020 (0.004)

0.094 (0.039)
0.141 (0.032)
-0.018 (0.007)
0.002 (0.013)

0.022 (0.017)
0.058 (0.012)
0.002 (0.007)
0.014 (0.006)

0.178 (0.103)
0.028 (0.023)
-0.005 (0.017)
0.005

0.142 (0.060)
0.082 (0.032)
0.028 (0.017)
-0.025 (0.017)


No Exclosure
Mean (SE)
0.021 (0.019)
0.052 (0.041)
-0.026 (0.022)
0.047 (0.012)

0.004 (0.027)
0.056 (0.021)
0.008 (0.008)
0.014 (0.006)

0.092 (0.051)
0.069 (0.034)
-0.031 (0.011)
-0.001 (0.013)

0.047 (0.011)
0.029 (0.022)
-0.006 (0.011)
0.018 (0.007)

0.053
0.026 (0.026)
-0.013 (0.025)
0.026

0.039 (0.073)
0.159 (0.041)
0.032 (0.018)
-0.029 (0.015)


Date


P
0.45
0.84
0.48
0.09

0.73
0.67
0.30
0.33

0.97
0.15
0.39
0.87

0.26
0.21
0.49
0.70

0.70
0.98
0.88


0.32
0.15
0.89
0.89


t-test
0.76
0.20
0.72
1.79

0.35
0.43
1.05
0.99

0.04
1.52
0.88
0.17

1.13
1.28
0.69
0.39

0.40
0.02
0.16


1.05
1.51
0.14
0.15


Amburana cearensis





Aspidosperma rigidum




Ceiba samauma





Copalifera chodatiana





Hymenaea courbaril




Pterogyne nitens


JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06

JanO6
March 06
May 06
Dec 06




















200 -\









O100 -E
EO

-20 -,Y C


-00


Moth

Fiue21 otl aifl A n olmosuetninmaurdb aemr@si
sesr (B.As hw i stewte de ote riae ol lt.Vria
lie niat 1sadad( =)











100


80 -I I ... .~. .. ... .. ... ...







S40-



20-




0-10 10-20 20-30 30-40 40-50 50-60 > 60

Canopy Openness (%)

Figure 2-2. Canopy openness in an unlogged control plot, an area subjected to normal timber
harvesting (4.7 m3/ha harvested), and an area subj ected to intensive harvesting (8.2
m3/ha) 8 months (left-hand bar) and 42 months (right hand bars) after logging.
Canopy openness measures were made with a spherical densitometer at the end of the
rainy season at 1 m above ground at 144 equally spaced points in each 10 ha plots.












Bromeliads No Bromeliads


Block 1 R R2 R R2W-XNW-NE


No BromeliadsBr


Block 2 R1 R2 R1 Ii R2
NW-EX WA-N

Bromeliads No Bo

Block 3 R1 R2 R1 R2 WA = WATER
NW = NO WATER
EX =EXCLOSURE
NE = NO EXCLOSURE

IR =Replications
SP1 SP2 SP3 SP4 SP5 SP6
.. SP =Species
No Bromeliads Bromeliads

Blc 0 R1 R2 R1 i R2







Figure 2-3. Design of the experiment on the effects of ground bromeliads, irrigation, extended
drought, and seed and seedling predator exclosures on seedling establishment.





1.0




0.5-




0.0
2003 2004 2005
Time (Years)


(A) AII Species _


_ Undisturbed
0- Normal Harvesting
7- Intensive Management




I ~a


4 -

4.0 -



2 -

3.0 -



0 -
2.0 -



1.0 -


(B) Without Acosmium


Figure 2-4. Seedling densities of 1 1 timber species (A) and 10 species without Acosmium
cardena~sii (B), the most dominant species, in a control plot, a plot subj ected to
normal timber harvesting, and a plot subj ected to intensive timber stand management.
The forestry treatments were carried out in 2001, 19 mo prior to the first census. Data
are from 4 m2 Subplots distributed regularly in a grid with 25 m spacing through each
10 ha treatment plot. Vertical lines indicate standard errors. Different letters indicate
different significance between treatments using pairwise LSD tests at 95%
confi dence.














M U0iltlUI
r 30 Normal Harvesting
V) E Intensive Harvesting

25-


20 -0


15-


10 -0


0 5-

0

2.0

S1.8-

c 1.6-



S1.2-

1.0-

S0.8-

S0.6-

0.4-

0.2-

0.0
2004 2005

Time (years)
Figure 2-5. Temporal changes in seedling recruitment (#/m2/y) and mortality for commercial
tree species in a Chiquitano dry forest in Bolivia.













-*- Bromeliad
3 -1 -0- No Bromeliad













1-


0 --r




4 C ` Exlsr









(C) -0- Exclosure













Sep5 Nov5 Jan6 Mar6 May6 Dec6

Time

Figure 2-6. Seedling density over time in response to (A) bromeliad cover, (B) irrigation or
drought, and (C) mammalian seed predators. Arrow indicates the time when I stopped
the irrigation and shielding plants from rainfall. Vertical lines show standard errors of
the means (N=40).











































































Figure 2-7. Mean densities (+ 1SE) of seedlings of commercial timber tree species in 2 m2 plOts
(N=40) with bromeliads (filled dots) and without bromeliads (open dots). The
number of seeds sown in each 1 m2 plOt is indicated after each species name.


Amburana cearensis (7)


Aspidosperma rigid )


0 -

0.6 -


0.4 -


0.2 -


0.0 -


0.6-


0.4-


0.2-


0.0-





0.8Copaifera choda


0.6-


0.4-


0.2-


0.0-





-0.8Ptrgnni


0.6-


0.4-


0.2-


0.0-



Dec6 Sep5 Nov5Jan6 Mar6 May6


-9 Bromeliad
-0 No Bromeliad


I I I I I


Ceiba samauma (8)


0.8


0.6


0.4


0.2


0.0







.-

0.6


0.4


0.2


0.0 -


I I I I I


~ns (8)


Hymenaea courbaril (3)


Dec6


Sep5 Nov5 Jan6Mar6 May6

















































































Figure 2-8. Mean of seedling densities (+ 1SE) in irrigated and droughted experimental plots.
Note differences in y-axis scales. Arrows indicate the time when I stopped the

irrigation and shielding plants from rainfall.


Amburana cearensis


Ceiba samauma





















Hymenaea courbaril









-j?--


Pterogyne nitens


-


-


-


-


-


1.0


0.8


0.6


0.4


0.2


0.0




















0.8





0.4


S0.2







0.0


-* Irrigation
-0 No irrigation


Copaifera chodatiana


Dec6 Sep5 Nov5Jan6 Mar6 May6


I I I I I


Dec6


Sep5 Nov5Jan6 Mar6 May6












Amburana cearensis


Aspidosperma rigidum


1.0 .


0.8 -1 T0.8-


0.6 -1 O 0.6-


0.4 -1 / 0.4-


0.2 -1 ~0.2-

-0 Exclosure
.0.0 -.0 e No exclosure





Ceiba samauma Copaifera chodatiana
1.0 1.0


0.8 -1 0.8-


S0.61 -0.6-


0.4 -0.4-


S0.2 -0.2-


0.0 0.0 -





Hymenaea courbaril Pterogyne nitens
1.0 1.0


0.8 -1 0.8-


0.6 -1 10.6-


0.4 -0.4-


0.2 -( T 0.2 i -- -


0.0 -0.0 -



Sep5 Nov5 Jan6Mar6 May6 Dec6 Sep5 Nov5 Jan6 Mar6 May6 Dec6

Figure 2-9. Mean seedling densities (+1 SE) in control plots (closed dots) and in plots from
which mammals were excluded (open dots).




























































Figure 2-10. Mean relative growth height growth rates (+ 1SE) of seedlings of commercial tree
species in response to three experimental treatments: (A) irrigation or drought; (B)
mammalian seed predator exclusion; and, (C) bromeliad cover. Numbers indicate the
number of seedlings used for each evaluation; numbers in brackets are the seedlings
in irrigated, exclosure, or bromeliad-cover plots while numbers in parenthesis are for
droughted, non-exclosure, and no-bromeliad plots. There were no significant
differences between treatments for each time calculated with t-test at the 95%
confidence level with sequential Bonferroni corrections (repeated-measures ANOVA
was not suitable because sample sizes declined over the study period due to seedling
mortality).


[99] [101] [91] [49]
-(45) (87) (84) (54)



-* Droughted
-0- Irrigated




(A)

[92] [1 20] [111] [64]
-(52) (68) (64) (41)


-*- No Exclosure
-0- Exclosure





(B)

[44] [58] [48] [28]
-(100) (130) (127) (77)


-*- No Bromeliad
-0- Bromeliad





(C)


0.14
0.12
0.10
0.08
0.06
0.04
0.02
0.00


0.14
0.12
0.10
0.08
0.06
0.04
0.02
0.00


0.14
0.12
0.10
0.08
0.06
0.04
0.02
0.00


I I


Jan6 Mar6


May6
Time


Dec6











CHAPTER 3
CONTRIBUTION OF ROOT AND STUMP SPROUTS TO NATURAL REGENERATION IN
A LOGGED TROPICAL DRY FOREST IN BOLIVIA

Introduction

Securing sufficient natural regeneration of commercial tree species after logging is critical

for sustainable forest management. Most studies of tropical forest regeneration focus on tree

recruitment from seeds and, consequently, regeneration is often viewed as depending on seed

production, seed dispersal, seed viability, and the environmental requirements for seed

germination and seedling establishment (Holl 1999, Dalling and Hubbell 2002, De Steven and

Wright 2002). In tropical dry forests, many tree species produce abundant and well-dispersed

seeds with high viability, but due to seed predation (Janzen 1971), water stress (Gerhardt 1994),

and a multitude of other factors, successful recruitment from seed is rare for many species in

these forests (Mostacedo and Fredericksen 1999). Furthermore, in forests in general and in dry

forests in particular, tree seedlings that do become established often grow more slowly than

sprouts (Miller and Kauffman 1998, Khurana and Singh 2001).

Given the limitations on seed dispersal and germination, as well as on seedling

establishment and survival, successful dry forest regeneration after logging, severe windstorms,

or fires may depend greatly on contributions from stump and root sprouts (Kruger et al. 1997,

Kammesheidt 1998, Miller and Kauffman 1998, Gould et al. 2002, Homma et al. 2003). The

general capacity of dry forest tree species to sprout may be an adaptive response to a history of

fire (e.g., Bond and Midgley 1995) or other disturbances. Whatever the ultimate cause, in a

variety of seasonal tropical forests, logging reportedly stimulates abundant stump sprouting of

felled and broken trees, and root sprouting from superficial roots damaged by heavy equipment

(Kauffman 1991, Kammesheidt 1998, Miller and Kauffman 1998, Kammesheidt 1999, Bell










2001). In the Chiquitano dry forest in Bolivia, although sprouting has been reported few times

following logging (Fredericksen et al. 2000) and fires (Gould et al. 2002, Kennard and Putz

2005), little is known about the overall contributions of sprout-origin plants to forest recovery.

Whereas interspecific comparisons of sprouting ability are numerous for Mediterranean

ecosystems (Bellingham 2000, Pausas 2001), how this ability varies with light requirements and

other ecological attributes is less clear for tropical dry forest species (but see Paciorek et al. 2000

for resprouting across the spectrum of shade tolerance of trees on Barro Colorado Island).

Sprouting is of interest to forest managers and ecologists because sprouts often have more rapid

grow rates than true seedlings (Daniel et al. 1979, Clark and Hallgren 2003).

The purpose of this study was to examine the contribution of sprouts to the natural

regeneration of a tropical dry forest following logging. More specifically, I (1) characterized

stump and root sprouting features of the commercial canopy tree species. I (2) quantified the

effect of logging on relative abundances and growth rates of stump sprouts, root sprouts, and true

seedlings. I (3) related the species-specific probabilities of stump sprouting as a function of

stump diameter and stump height; and I (4) explored how sprouting varies with the ecological

requirements of canopy tree species.

Methods

Study Area

This study was conducted on the property of INPA Parket (hereafter INPA), a 30,000-ha

tract of privately-owned seasonally dry tropical forest 30 km NE of the town of Concepci6n (160

6' 45"S, 610 42' 47"), 250 km northeast of the city of Santa Cruz de la Sierra, Bolivia (Figure 1-

1). The study area is flat to gently sloping, with an altitude of approximately 380 m, mean annual

temperature of 24.3 OC, and mean annual precipitation of 1 100 mm. During the 5-mo dry season

(May-October), most trees are deciduous and many tree species flower and fruit following rain









events in the mid- to late-dry season. The forest canopy is 20-25 m tall with common species

including Acosmium cardena~sii, Tabebuia impetiginosa, Anadenanthera macrocarpa,

Astronium urundeuva, and Centrolobium microchaete (Pinard et al. 1999); after first mention,

species will be referred to by their generic names (for full names see Table 3-1). Currently, 21

tree species, including those mentioned above, are harvested for timber processed mostly into

parquet flooring.

During the rainy season, canopy openness, as measured 1 m above the ground with a

spherical densiometer, was 8% and 14% in control and logged areas, respectively. During the dry

season, canopy openness triples because many tree species are deciduous. The understory is

dense, partially due to the abundance of lianas, and typically 30-40% of the ground is covered by

the bromeliad, Pseudananas~ddd~~~ddd~~~dd sagenarius.

Experimental Design and Data Collection

Stump sprouts

I sampled sprouts from the stumps of harvested trees in three areas that varied in time since

logging. The first area (50 ha) was selectively logged by INPA Parket (- 4 trees/ha and 4 m3/ha,

and 10-12 species harvested) 1 y before I began my study. Here I mapped, marked, and

measured the diameters and heights of the stumps and all stump sprouts of the five most

commonly harvested tress (Anadenanthera, Centrolobium, Copaifera chodatiana, Tabebuia, and

Zeyheria tuberculosa; tree densities reported in Table 2-1) and monitored sprout survival and

height growth for one year. The second study area (40 ha) is located in the 20-ha permanent plots

maintained by the Instituto Boliviano de Investigaci6n Forestal (IBIF) for monitoring forest

dynamics after logging. Two years prior to my study, 4-8 trees/ha (5.3-6.4 m3/ha, 14 species)

were logged from these plots. I monitored sprouting as described above. The third area (30 ha)

was logged (2-3 trees/ha, -3 m3/ha, and 5-7 species harvested) 5 y before my study. I measured









the frequency and height of stump sprouts of six tree species (Anadenanthera, Caesalpinia,

Centrolobium, Copaifera, M~ scleroxylon, and Tabebuia) in this area, using a 7 m telescoping

measuring rod.

In 2003 I checked for sprouts on the stumps of trees harvested in 1998 (6 species),

2001(10 species), and 2002 (6 species). The 498 stumps evaluated in the three areas were from

trees > 40 cm DBH that were felled with chainsaws 10-90 cm height above ground. Each

species was placed in one of the following four ecological guilds based on field observations and

the literature (Whitmore 1998, Mostacedo and Fredericksen 1999, Poorter et al. 2006): light-

demanding pioneers have light requirements and are short-lived; long-lived pioneers also are

light-demanding but are longer lived; somewhat shade tolerant species establish in the shade but

only mature under moderate to high light intensities; and, shade-tolerant species can establish

and survive in the shade. I counted all sprouts and measured the heights of the two tallest on each

stump (from the point of origin) as well as the height and diameter of each stump dating from the

2001 and 2002 harvests.

Comparison of different juvenile types in relation to microsites created by logging

In two of IBIF's 20-ha experiment plots, I compared the densities and sizes of seedlings

and sprouts < 2 m tall in the following microsites created by an episode of selective logging that

occurred 1.5 years prior to my study: logging gaps (280-330 m2, N = 16); logging roads (N =

16); log landings (N = 8); primary skid trails (N = 16); and, secondary skid trails (N = 16).

Secondary skid trails were those used to extract a single log, while primary skid trails were those

where skidder had extracted > 2 logs. In each microsite, all plants < 2 m tall of 16 canopy tree

species in 10 x 4 m plots were classified as having developed directly from a germinated seed or

sprouted from a root or stem; determination of plant origin often involved excavation, but was

generally unambiguous.









Data Analysis

I used logistic regression to determine the probability of a stump sprouting in relation to its

diameter and height for each the Hyve species that sprouted frequently (Anadenanthera,

Centrolobium, Copaifera, Tabebuia, and Zeyheria). Nagelkerke R-square values were used to

determine the percentage of variance explained by each regression and a Hosmer and Lemeshow

X2 gOodness-of-fit tests was used to determine the significance of each relationship (Field 2000).

To determine whether there are relationships between stump diameter and height

(independent variables) with the number and maximum heights of sprouts (response variables), I

used linear regressions or nonlinear regression analyses based on linear, quadratic, cubic, and

inverse models. For each species, the simplest (i.e., fewest parameters) model with a high R2

value was selected in which each parameter had a reasonable biological explanation.

Analyses of variance (ANOVAs) followed by Tukey's post-hoc comparisons were used to

compare densities of seedlings and root or stem sprouts among logged microsites. Absolute

relative annual height growth rates of stump sprouts were calculated for 10 species based on their

height 2 y after logging, and heights after 1, 2, and 5 y after logging for 6 species. Stump sprout

heights were compared among ecological guilds using ANOVAs and Tukey's post-hoc tests. For

each species and ecological guild, an ANOVA and then Tukey's post-hoc comparison were used

to compare mean growth among origin types (i.e., stump sprout, root sprout, or true seedling).

All analyses were carried out with SPSS 12.0 for Windows.

Results

Sprout Characterization.

Stump sprouting was common after logging in the dry forest studied; 27 of the 3 1 species

monitored at least occasionally reprouted from stumps; 62% did so frequently (Table 3-1).

Centrolobium, Zeyheria, and Tabebuia were the most frequent stump sprouters (Figure 3-1).










Among the six commercial tree species monitored, the proportion of stumps with living sprouts

decreased with time since logging (Figure 3-2). Overall, for stumps censused 1, 2, and 5 years

after logging the proportion of stumps with live sprouts was 55%, 43%, and 38%, respectively,

but the rate of stump sprout mortality varied by species. In particular, 80% of the Caesalpinia

stumps and 73% of the Centrolobium stumps had live sprouts 5 y after the trees were felled.

Whereas high proportions of Anadenanthera and Copaifera stumps initially sprouted (23 and

13%, respectively), neither species had living stump sprouts in the plot logged 5 y prior to my

study .

Root sprouting was also common after logging in the tropical dry forest of INPA. Of the

31 tree species monitored (Table 3-1), 16 sprouted from lateral roots, 7 species at high

frequencies. Acosmium cardena~sii, Centrolobium, and Casearia gossypiosperma were the most

frequent root sprouters.

Most of the 27 species that frequently sprouted from roots or stumps were shade tolerant

(9) or at least partially shade-tolerant (8). Light-demanding pioneer species sprouted

infrequently, if at all (e.g., Astronium, Piptadenia~~ttt~~~~ttt~~~ viridifolia, Acacia bonariensis, and Schinopsis

brasiliensiss. The most frequent sprouters were the long-lived pioneer species, Centrolobium and

Tabebuia, and the partial shade-tolerant species, Zeyheria (Table 3-1).

Caesalpinia pluviosa had the higher (20.014.4) number of sprouts per stump, followed by

Centrolobium (15.411.47) and Zeyheria (14.211.5). The other four of the seven species

monitored had < 5 sprouts/stump (SE=0.5), with the absolute lowest numbers observed in

Copaifera and Anadenanthera (Figure 3-3)

Considering all sprouted stumps, there was a significant negative linear relationship

between the number of sprouts per stump and stump diameter (Table 3-2). In contrast, when










species were considered separately, the only species that showed a significant (quadratic)

relationship between number of sprouts and stump diameter was Copaifera, and that relationship

was positive (i.e., opposite from the overall trend). For all species considered together, there was

a negative linear relationship between the number of sprouts and stump height, but the

relationship varied among species. The number of Caesalpinia sprouts increased with stump

height whereas in Centrolobium and Tabebuia, the relationship was negative (Table 3-2).

Sprout height growth, based on measures of the tallest stump sprouts 1-5 y after the trees

were cut, generally decreased with stump diameter, but species varied in this relationship. Only 3

of the 7 species for which I have sufficient data showed significant trends: Caesalpinia showed a

positive cubic relationship; in Machaerium the relationship was inverse positive; and, Copaifera

had a quadratic and positive relationship (Table 3-2). Sprout height growth rates decreased with

stump height when all of the species were considered together. In contrast, at the species level,

only Caesalpinia and Machaerium showed significant relationships between sprout growth rates

and stump height, but in the former the relationship was negative and quadratic and the latter,

positive and inverse (Table 3-2).

The probability of stump sprouting as related to stump diameter varied among species

(Figure 3-3). In Copaifera, Anadenanthera, Tabebuia, and Centrolobium, the proportions of

sprouted stumps were approximately 0.17, 0.24, 0.57, and 0.97, respectively, and did not vary

with stump diameter. In Zeyheria, sprouting reached 98% of the stumps 38-40 cm diameter but

decreased to only 40% among stumps 90 cm in diameter (Figure 3-4).

The probability of sprouting in relation to stump height also varied among species Figure

3-5). The proportions of sprouted stumps of Copaifera, Anandenanthera, and Centrolobium did

not vary with stump diameter. In contrast, a Tabebuia stump 10 cm tall was almost certain to










sprout (0.98) whereas this probability declined to 0.13 for a 72 cm tall stump. In Zeyheria, the

probability of sprouting was high (0.91) and did not vary with stump height (Figure 3-5).

Juvenile Types and the Effects of Logging

In the plots censused 1.5 years after logging, 45% of juveniles < 2 m tall of canopy tree

species were root and stem sprouts, not true seedlings (Figure 3-6). At the species level there

was great variation in the proportions of true seedlings (Figure 3-7). All 15 species evaluated

were represented by some sprouts and sprouted at least occasionally from roots whereas only 9

species were represented by stem sprouts. Three species sprouted predominantly from roots

whereas stem sprouting was the predominant mode of regeneration in only one species. Light-

demanding species tended to regenerate more from seeds and root sprouts than from stem sprouts

(F=12. 10, P=<0.0001), while partially shade tolerant and shade tolerant regenerated more from

seeds (F=4.46, P=0.01; F=8.01, P=0.0004; respectively).

Densities of plants < 2 m tall of canopy tree species did not vary among the logging

microsites (F = 1.37, P = 0.24), but microsites differed in the relative contributions of true

seedlings and sprouts (Table 3-3). True seedlings were twice as abundant as root and stem

sprouts combined in logging gaps (F = 9.91, P = 0.0001). In contrast, there was no difference in

plant density by origin in logging roads (F = 0.38, P = 0.68). Densities of plants from root

sprouts and true seedlings were similar in log landings (F = 1.95, P = 0.18). On primary skid

trails most plants <2 m tall were true seedlings, with fewer root sprouts, and almost no stem

sprouts (F = 13.57, P < 0.0001). On secondary skid trails, true seedlings were much more

common than plants of either sprout type (F = 4.6, P = 0.01).

Growth of Stump Sprouts

Based on measures of the tallest sprout per stump, the growth rates of stump sprouts varied

among species by more than an order of magnitude (Figure 3-8). Anadenanthera (197 cm/y),










Centrolobium (195 cm/y) and Zeyheria (185 cm/y) had the highest growth rates, while

Aspidosperma (3.5 cm/y) and Copaifera (25 cm/y) had the lowest. Stump sprouts of light-

demanding pioneer species grew faster than those of shade-tolerant species, but there was a great

deal of within species variation, especially in the growth rates of the latter (Table 3-4). Stump

sprouts of long-lived pioneer species grew at about the same rates as shade tolerant species.

Among individuals of canopy tree species < 2 m tall, root and stem sprouts both grew

faster than seedlings and in 5 of 12 species monitored, root sprouts grew faster than stem sprouts

(Table 3-5). Centrolobium and Chorisia speciosa had the highest root sprout growth rates. I

found no Anadenanthera, Aspidosperma, Casearia arborea, M. scleroxylon, Phyllostylon

rhamnoides, or Piptadenia~~ttt~~~~ttt~~~ stem sprouts (Table 3-5). Among the root sprouters, light-demanding

and long-lived pioneer species grew faster than partially shade tolerant and shade tolerant

species. Among the stem sprouters and true seedlings, shade-tolerant species grew slower than

belonging to other light-syndrome classes (Table 3-6).

Stump sprout heights varied over time and among the six species censused 1, 2, and 5 y

after logging (Figure 3-9). Apparent absolute growth rates (cm/y) increased through the second

year and then height increments stopped except in Centrolobium, which continued to grow at a

rapid rate through the fifth year. Anadenanthera sprouts were apparently growing rapidly

through the second year, but I could find no live stumps in the plot logged 5 y prior to my

census.

Discussion

Of the 3 1 canopy tree species studied in a dry tropical forest in Bolivia, 27 (87%) have

some capacity to sprout from either roots or stumps. Sprouting is apparently characteristic of

many tropical dry forest tree species and helps them persist in an environment where stress is

severe and disturbances are frequent (Bellingham 2000, Bond and Midgley 2001). As observed









in the USA (Jones and Raynal 1988) and Mexico (Dickinson 1998), root sprouting was promoted

by logging damage to roots in my study species. The high proportion of species that sprouted

from broken and cut stems in INPA may be related to the high frequency with which stems lose

their terminal buds due to herbivore browsing during the dry season when other browse is scarce

as well as to the direct effects of drought stress (Bossard and Rejmanek 1994, Del Tredici 2001,

Groll 2005).

Natural Regeneration and Shade Tolerance: True Seedlings vs. Sprouts

Scarcity of natural regeneration from seeds is common for most tree species in tropical dry

forest in Bolivia (Mostacedo and Fredericksen 1999). The main reasons for this scarcity appear

to be high seed predation, low seed viability, and high seedling mortality during the dry season.

Sprouting from broken and cut stems, along with root sprouting, appears to be a very important

regeneration mechanisms in tropical dry forests in Brazil (Castellani and Stubblebine 1993),

Jamaica (Bellingham et al. 1994), and Venezuela and Paraguay (Kammesheidt 1999), including

the forest I studied in Bolivia where 45% of the regeneration of canopy trees originated from root

or stem sprouts.

Sprouting is a common mode of tree regeneration in forests around the world. For example

in oak forests in the USA (Clark and Hallgren 2003, Nyland et al. 2006) and in boreal forest in

Russia (Homma et al. 2003), trees reportedly regenerate mainly from sprouts. Sprouting seems to

represent the predominant mode of regeneration in forests frequently subj ected to logging, wind

damage, and fire (Bond and Midgley 2001).

Natural regeneration by sprouting from lateral roots was common in some commercial

species in my study site. In particular, Centrolobium, Tabebuia, Aspidosperma, and M~

scleroxylon regenerated mostly from root sprouts. Centrolobium was previously reported as a

root sprouting species (Fredericksen et al. 2000), but the importance of this mode of regeneration









in the other species has apparently been overlooked. It remains to be seen whether root sprouts

mature into sound trees, and there are reasons to suspect that they will not. First of all, given that

most new stems emerge from damaged stems or roots, sprouts of all sorts seem particularly

prone to butt and root rots. Second, I observed that several Centrolobium root sprouts that were

5-8 cm DBH 5 y after sprouting still had not developed their own root systems. Those that I

excavated emerged from large diameter roots running about 5 cm below the soil surface but had

developed almost no roots of their own and were thus mechanically unstable when pushed

perpendicular to the orientation of the source root. Given the general importance of root

sprouting after fires, logging, and other severe disturbances, such as found in tropical dry forest

in Paraguay and moist semi-deciduous forest in Venezuela (Kammesheidt 1999), root sprout

longevity is an issue that deserves more attention from researchers.

Partially shade-tolerant and shade-tolerant species were more likely to sprout than light-

demanding species. Most of the partially shade-tolerant species in INPA sprouted from either

roots or stems; similar findings were reported for a moist but seasonal tropical forest in Panama

(Paciorek et al. 2000). In contrast, in a moist tropical forest but after slash-and-burn agriculture

of eastern Paraguay, light-demanding species contributed more sprouts than shade-tolerant

species (Kammesheidt 1998). Although some light-demanding species in INPA did not sprout

(13%), others stump sprouted frequently, such as species in the Bombacaceae and

Flacourtiaceae. Furthermore, the light-demanding pioneer species that did sprout grew faster

than sprouts from other ecological groups.

Allometric Relationships with Stump Sprouting

The weak and inconsistent trends in the relationship between either stump diameter or

stump height and the growth rates or number of sprouts per stump means that I have little basis

on which to make firm recommendations for sprout management. Consistent patterns in sprout









responses are also not apparent in the literature. Whereas several studies mention that in the first

years after tree cutting there is a positive relationship between stump diameter and sprout growth

rates (Jobidon 1997), other studies report the opposite (Trani et al. 2005), and generally the

relationship does not remain significant after a few years. These results suggest that factors other

than stump size controls sprouting (McConnaughay et al. 1996, Masri et al. 1998).

The probability of sprouting varied substantially among species but I observed no effect of

stump diameter on the probability of sprouting in four of the five species studied. For example,

Centrolobium had the highest probability of stump sprouting (97%), while Copaifera had the

lowest (17%). In contrast, Zeyheria showed a decreasing probability of stump sprouting with

increasing stump diameter, a pattern also observed in a wetter but still seasonal tropical lowland

forest in Panama (Putz and Brokaw 1989) and in an oak forest in southern Indiana (Weigel and

Peng 2002). The probability of stump sprouting did not vary either with stump height except in

Tabebuia, in which the probability decreased with stump height. These results suggest that

harvesting trees of any tree size will promote same probability of sprouting, except for Zeyheria

in which it is better to cut smaller trees and in Tabebuia in which low stumps are preferred if

sprouting is to be encouraged.

Growth of Stem and Root Sprouts Compared with True Seedlings

One advantage of natural regeneration via sprouting is that sprouts typically grow more

rapidly than true seedlings, at least initially (Gould et al. 2002, Kennard et al. 2002), which was

confirmed by this study. I also observed that small plants of sprout origin typically appeared less

affected by drought than true seedlings (Personal Observation). Nevertheless, in my study forest

as well as in Australia (Enright and Goldblum 1999) and South Africa (Kruger et al. 1997), true

seedlings of some species grew just as fast as sprouts. In several species, especially light-










demanding pioneers (e.g., Cordia, Casearia), height growth rates were similar between true

seedlings and root or stem sprouts.

Conclusions

Due to the high costs and frequent failures of seed and seedling planting, natural

regeneration is critical for the sustainable management of tropical dry forest tree species in

Bolivia. Given that so many tree species sprout prolifically from stumps of all sizes or from

lateral roots, especially after mechanical damage, sprouts need to be considered as a source of

regeneration. The abundance of sprouts and their typically rapid growth rates, when compared

with those of true seedlings, adds to the potential value of sprouts for forest management. In

forests not designated for timber stand management, sprouts deserve at least as much attention

from researchers as seeds and true seedlings. That said, future studies should consider the long-

term fates of sprouts. The observation in this study that the stump sprouts of most species

essentially stopped growing after 2 y needs to be verified and otherwise explored, as do the

factors that cause high rates of mortality of the sprouted stumps of some species. In the case of

root sprouts, which were also abundant in my study area, long-term monitoring is needed to

determine whether they ever grow up to be sound, canopy trees.













Table 3-1. Frequency of root and stem sprouting and shade tolerance of commercial and non-commercial canopy tree species in a
tropical dry forest in Bolivia. Shade tolerance is base on Pinard et al. (1999), and Mostacedo and Fredericksen (1999):
L=Light-demanding pioneer, LL=Long-lived pioneer, PS=Partially shade tolerant, ST=Shade tolerant.


Resprout Type/ Frequency
ROOT STEM


Shade Tolerance


Species


Abbreviation Family


Commercial Timber Sp~ecies
Amburana cearensis (Allemlio) A.C. Sm.
Anadenanthera macrocarpa (Benth.) Brenan
Aspidosperma rigidum Rusby
Astronium urundeuva (Allemcro) Engl.
Caesalpinia pluviosa DC.
Can'niana ianeirensis R. Knuth
Cedrela fissilis Vell.
Centrolobium microchaete (Mart. ex Benth.) Lima ex G. P. Lewis
Copaifera chodatiana Hassl.
Cordia alliodora (Ruiz & Pay.) Oken
Hymenaea courbaril L.
M2/achaerium scleroxylon Tul.
Phyllostylon rhamnoides (J. Poiss.) Taub.
SPlartymiscium ulei Harms
Schinopsis brasiliensis Engl.
Sweetia fruticosa Spreng.
Tabebuia impetiginosa (Mart. ex DC.) Standl.
Tabebuia \r~rarrit;,idi (Vahl) G. Nicholson
Zeyhena tuberculosa (Vell.) Bureau

Non-commercial Sp~ecies
Acacia bonariensis Gillies ex Hook. & Arn.
Acosmium cardenasii H.S. Irwin & Arroyo
Aspidosperma cylindrocarpon Milll. Arg.
Capparis pnsca J.F. Macbr.
Caseana gossypiosperma Briq.
Ceiba samauma (Mart.) K. Schum.
Chon'sia speciosa A. St.-Hil.
Enotheca roseorum (Cuatrec.) A. Robyns
Gallesia integrifolia (Spreng.) Harms
M2/achaerium atr urit;,idimu Vogel
Piptadenia 1 n ,, 1,r /.., (Kunth) Benth.
Spondias mombin L.


AMCE
ANMA
ASRI
ASUR
CAPL
CAIA
CEFI
CEMI
COCH
COAL
HYCO
MASC
PHRH
PLUL
SCBR
SWFR
TAIM
TASE
ZETU


ACBO
ACCA
ASCY
CAPR
CAGO
CESA
CHSP
ERRO
GAIN
MAAC
PIVI
SPMO


Caesalpinaceae
Mimosaceae
Apocynaceae
Anacardiaceae
Caesalpiniaceae
Lecythidaceae
Meliaceae
Fabaceae
Caesalpiniaceae
Boraginaceae
Caesalpiniaceae
Fabaceae
Rhamnaceae
Fabaceae
Anacardiaceae
Fabaceae
Bignoniaceae
Bignoniaceae
Bignoniaceae


Mimosaceae
Fabaceae
Apocynaceae
Capparaceae
Flacourtiaceae
Bombacaceae
Bombacaceae
Bombacaceae
Phytolacaceae
Fabaceae
Mimosaceae
Anacardiaceae


No
No
Yes/Low
No
Yes/Low
Yes/Low
Yes/Low
Yes/High
Yes/Low
Yes/Low
Yes/Low
Yes/Low
No
No
No
No
Yes/Low
Yes/High
No


No
Yes/High
Yes/High
No
Yes/High
No
Yes/High
No
No
Yes/High
No
No


Yes/Low
Yes/Low
Yes/High
No
Yes/High
Yes/High
Yes/Low
Yes/High
Yes/High
Yes/Low
Yes/Low
Yes/High
Yes/High
Yes/High
No
Yes/Low
Yes/High
Yes/Low
Yes/High


No
Yes/High
Yes/High
Yes/High
Yes/High
Yes/Low
Yes/High
Yes/Low
Yes/High
Yes/Low
No
Yes/Low












Table 3-2. Summary of the best models and their significance from the regression analyses between stump diameter or stump
volume (independent variables) and the heights of stump and numbers of sprouts per stump (dependent variables). Tests
were made for species that were sampled with > 8 individuals at alpha = 0.05. Models: L=Linear, Q=Quadratic, C=Cubic,
I=Inverse. Signs mean positive (+) or negative (-) relationship between two variables.
Scintfi Nme# of sprouts Height of sprouts
Model R2 DF F P Model R2 F P
Stump Diameter
4nadenanthera inacrocarpa L 0.015 36 0.56 0.461 L 0.034 1.27 0.2680
Caesalpinia pluviosa Q(+) 0.267 18 3.27 0.061 C(+) 0.406 6.16 0.0090
Centrolobiwn inicrochaete L 0.005 90 0.41 0.522 L 0.012 1.09 0.3000
Copaifera chodatiana Q(-) 0.390 21 6.71 0.006 Q(-) 0.536 12.11 <0.0001
Machaerium scleroxyvon Q 0.365 5 1.44 0.321 I(+) 0.630 10.24 0.0190
Tabebuia ,,i, rs,.so L 0.026 54 1.45 0.234 L 0.010 0.54 0.4660
Zevheria tuberculosa L 0.001 62 0.05 0.819 I 0.036 2.33 0.1320
All species L(-) 0.020 301 5.61 0.020 L(-) 0.040 3.95 0.0400

Stump Height
4nadenanthera inacrocarpa L 0.01 36 0.49 0.49 L 0.001 0.03 0.87
Caesalpinia pluviosa L(+) 0.4 19 12.90 0.002 Q(-) 0.32 4.24 0.03
Centrolobiwn inicrochaete I(-) 0.36 90 49.96 <0.0001 I 0.02 2.06 0.16
Copaifera chodatiana L 0.03 22 0.66 0.42 L 0.02 0.42 0.52
Machaerium scleroxyvon L 0.006 6 0.04 0.85 I(+) 0.62 10.03 0.02
Tabebuia ,,i, rs,.so I(-) 0.34 59 29.82 <0.0001 L 0.02 0.90 0.35
Zevheria tuberculosa I 0.03 63 2.06 0.16 L 0.008 0.53 0.47
All Species L(-) 0.03 311 8.91 0.003 I(-) 0.16 56.6 <0.0001











Table 3-3. Mean (+ 1SE) densities of true seedlings, stem sprouts, and root sprouts in 10 x 4 m plots in microsites created during
selective logging. Different letters indicate differences between microsites in the densities of plants of different origins
using Tukey post hoc comparisons with 95% of confidence.


Microsites
Logging Gaps
Logging Roads
Landings
Primary Skid Trail
Secondary Skid Trail


Seedling
78.75 (10.47)a
28.43 (11.45)a
34.58 (15.84)a
50.15 (9.47)a
59.84 (11.49)a


Stem Sprot
21.87 (5.18)b
8.33 (3.33)a
0.62 (0.62)a
4.21 (1.87)c
31.41 (10.62)b


Root Spot
23.44 (8.82)b
26.09 (6.80)a
24.64 (7.83)a
24.06 (4.89)b
19.53 (5.17)b


Mean of Square
11341
513
1411
8493
6866


F
9.91
0.38
1.95
13.57
4.64


P
0.0001
0.68
0.18
0.0001
0.01













Table 3-4. Mean (+ 1SE) of stump sprout height growth rates (cm/year) by ecological groups.
Different letters indicate significant differences between ecological groups using
Tukey post hoc comparisons with 95% confidence.


Ecological Group
Light-demanding Pioneer
Long-lived Pioneer
Partially Shade Tolerant
Shade Tolerant


N Mean
47 194.7 (17.4)a
6 84.8 (17.8)"
24 159.5 (12.3)b
4 14.5 (8.8)"


Mean Square
57005.7


0.001











Table 3-5. Means (+ 1SE) of stem heights by species that sprouted from stems or roots compared to the heights of seedlings. Different
letters indicate significant differences in plant origins within species using Tukey post hoc comparisons at the 95%
confidence level.


Species


Seedling


Stem Resprout


Root Sprout


Mean
Square
28623
11042
568
6770
300
1844
32850
352
961
3040
23291

0.8
54643
2552
10001


Acosnzium cardena~sii
Anadenanthera nzacrocalpa
Aspidospernza rigidunt
Caesalpinia pheviosa
Casearia arborea
Casearia gossypiosperza
Centrolobium naicrochaete
Chorisia speciosa
Copaifera chodatiana
Cordia alliodora
Machaeriunt acutifolium
Machaerium scleroxylon
Phyllostylon rha~nnoides
Piptadenia~~ttt~~~~ttt~~~ viridifolia
Sweetia f~uticosa
Tabebuia inspetiginosa


18.3(2.3)"
25.1(1.5)b
30.0
54.7(16.5)
125.0(52.0)
102.7(10.9)
67.8(26.6)b
141.2(15.7)
31.7(16.6)
135.0(39.1)
33.6(2.5)b

14.0(2.27)
69.0(6.2)b
58.0(17.2)
44.5(36.8)


42.4(4.0)a 46.2(3.6)a
-72.6(10.6)a
-55.0(5.68)
100.5(29.8) 112.5(29.8)
-105.0(90.1)
100.0(19.5) 78.8(13.8)
138.7(39.9)a 154.2(4.8)a
-125.0(22.2)
61.4(12.9) 33.0(28.8)
155.0(67.6) 55.0(67.6)
59.2(5.3)a 60.7(4.1)a
-70.0(5.0)
~15.0
-140.0(19.1)a
110.0 120.0
90.0(52.0) 116.9(11.4)


27.35
19.6
1.76
1.90
0.037
0.97
5.15
0.20
1.15
0.66
21.09

0.03
12.51
1.23
1.85


0.0001
0.0001
0.22
0.18
0.86
0.39
0.006
0.67
0.37
0.60
0.0001

0.86
0.001
0.36
0.17












Table 3-6. Mean (+ 1SE) heights of trees I 2 m tall that were root sprouts, stem sprouts, or seedlings grouped by ecological guild.
Different letters indicate significant differences between ecological groups using Tukey post hoc comparisons at the 95%
confidence level.


Origin types

Root sprout
Stem sprout
Seedling


Light-demanding
pioneer
149.1 (3.9)a
86.4 (13.0)ab
44.7 (2.0)a


Long-lived
pioneer
116.9 (10.7)b
90.0 (34.5)a
44.5 (20.1)a


Partially shade
tolerant
64.6 (8.2)c
64.0 (7.3)ab
35.9 (2.8)a


Shade tolerant


Mean
Square
323997
9593
30298


46.7 (7.3)c
43.7 (5.7)b
18.5 (2.9)b


66.8
4.1
18.6


< 0.0001
0.009
< 0.0001













100



80



60



40



20



0


CEMI CAPL MASC ZETU TAIM ANMA COCH


Species
Figure 3-1. Proportions of stumps of commercial timber species that resprouted (number of
stumps noted in parenthesis). CEMI = Centrolobium microchaete, CAPL =
Caesalpinia pluviosa, MASC = Machaerium scleroxylon, TAIM = Tabebuia
impetiginosa, ANMA = Anadenanthera macrocarpa, COCH = Copaifera chodatiana.











100


Centrolobium Caesalpinia M. scleroxylon Tabebuia Anadenanthera Copaifera


Species
Figure 3-2. The proportions of stumps with live sprouts over time since logging. For complete
species names see Table 3-1.





25-



E 20-






15


10




CAPL CEMI ZETU TAIM MASC ANMA COCH

Species
Figure 3-3. Mean (+ 1SE) numbers of sprouts/stump for the most frequent sprouting species.
CEMI = Centrolobium microchaete, CAPL = Caesalpinia pluviosa, MASC =
Machaerium scleroxylon, TAIM = Tabebuia impetiginosa, ANMA = Anadenanthera
macrocarpa, COCH = Copaifera chodatiana.