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1 XATE PALM ( CHAMAEDOREA SP. ) ENRICHMENT IN WESTERN BELIZE: THE ECOLOGICAL EFFECTS OF MANAGEMENT IN RELATION TO UNDERSTORY PLANT SPECIES RICHNESS, DIVERSITY, AND COMPOSITION By CARLY N. VOIGHT A THESIS PRESENTED TO THE GRAD UATE SCHOOL OF THE UNIVERSITY OF FLORIDA IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DE GREE OF MASTER OF ARTS UNIVERSITY OF FLORIDA 2011
2 2011 Carly N. Voight
3 To the people, animals, plants, and ecosystems of the Mayan Forest
4 ACK NOWLEDGMENTS This research would not have been possible without the support of a multitude of professors, colleague s and friends. I would like to thank my advisor, Dr. Nigel Smith, and my committee members, Dr. Walter Judd and Dr. Emilio Bruna for their a ssistance in plant identification statistical analyses, guidance, constructive criticism, and enthusiasm. I acknowledge the Center for Latin American Studies, the Tropical Conservation and Development Program, the Tinker Foundation Field Research Grant, a nd the Foreign Language Area Studies Award at the University of Florida for funding and guidance. Words cannot begin to express my gratitude to James Mesh and Antonio Mai for their tireless help in contacting xate farmers, field assistance, kindness, and s upport. I would also like to thank the Itzamna Society including Maria Garcia and the Belize Myra, Abby, and little Jimmy who welcomed me into their home. My thanks go to the xate farmers in San Antonio, El Progresso 7 Miles and Cristo Ray for being willing to discuss their xate management and allowing a crazy gringa to c ollect plants from their land. I also t hank Pio Saqui for his assistance with contacts in Belize. I would l ike to thank my former supervisor at The Nature Conservancy, Dr. J. Dan Husband, as well as Kurt Neubig and Mario Blaco for their taxonomic assistance I t hank specialists Richard Abbott ( Polygalaceae ) Thomas Daniel (Acanthaceae) and Erin Tripp (Acanthac eae) for further plant identifica tion support in their respective areas. I appreciate the assistance of Kent Perkins and Trudy Lindler at the University of Florida Herbarium (FLAS).
5 I would like to deeply thank my family and friends who have provided encou ragement throughout my degree. I thank my parents, Jack and M arty Voight, and my friends, Christopher Strickland, Fern Toro, Samantha Alvarez, Amanda Hymansmith, Andrea Chaloux and my cats Smokey and Anja for their emotional support, advice, encouragement, willingness to listen, and hugs. I would not be where I am today without your love and support. Jach yum bo'otik!
6 TABLE OF CONTENTS page ACKNOWLEDGMENTS ................................ ................................ ................................ .. 4 LIST OF TABLES ................................ ................................ ................................ ............ 8 LIST OF FIGURES ................................ ................................ ................................ .......... 9 ABSTRACT ................................ ................................ ................................ ................... 11 CHAPTER 1 BA CKGROUND ................................ ................................ ................................ ...... 13 Introduction ................................ ................................ ................................ ............. 13 Xate Ecology and Harvest ................................ ................................ ...................... 15 Ecology and Distribution of Xate ( C. ernesti agustii ) ................................ ........ 16 Uses of Xate and the Xate Industry ................................ ................................ .. 16 Wild xate harvest in Mexico and Guatemala ................................ .............. 17 Wild xate harvest in Belize ................................ ................................ ......... 19 Shade grown NTFP plantations ................................ ................................ 23 Shade grown xate plantations in Belize ................................ ..................... 25 Landscape and Cultural Setting ................................ ................................ .............. 27 2 RESEARCH HYPOTHESES AND METHODOLOGY ................................ ............. 40 Research Hypotheses ................................ ................................ ............................. 40 Research Methodology ................................ ................................ ........................... 41 Experimental Design ................................ ................................ ........................ 41 Description of specific site locations ................................ .......................... 42 Data collection and species identification ................................ .................. 42 Data Analysis ................................ ................................ ................................ ... 45 Species richness ................................ ................................ ........................ 45 Species diversity ................................ ................................ ........................ 46 Species composition ................................ ................................ .................. 46 Canopy Cover ................................ ................................ ............................ 47 3 RESEARCH RESULTS ................................ ................................ .......................... 49 Species Richness ................................ ................................ ................................ ... 49 Species Diversity ................................ ................................ ................................ .... 49 Species Composition ................................ ................................ .............................. 50 Plant Species Dominance ................................ ................................ ................ 50 Non Forest Species ................................ ................................ .......................... 50 Rare Species ................................ ................................ ................................ .... 52 Canopy Regeneration ................................ ................................ ...................... 53
7 Canopy Openness ................................ ................................ ................................ .. 54 4 DISCUSSION ................................ ................................ ................................ ......... 69 Conclusions ................................ ................................ ................................ ............ 69 Research Implications ................................ ................................ ............................. 71 Implications for Sustainable Forest Management ................................ ............. 71 Implications for Wildlife Conservation ................................ ............................... 79 Limitations ................................ ................................ ................................ ............... 81 Conclusion ................................ ................................ ................................ .............. 82 APPENDIX A LIST OF SPECIES ................................ ................................ ................................ .. 87 B ESTIMATED RICHNESS RESULTS ................................ ................................ ...... 96 C LIST OF INFORMANTS ................................ ................................ .......................... 97 LIST OF REFERENCES ................................ ................................ ............................... 98 BIOGRAPHICAL SKETCH ................................ ................................ .......................... 107
8 LIST OF TABLES Table page 3 1 Rare species found in research study and corresponding rarity status. ............. 63 3 2 Canopy regeneration in forest sites ................................ ................................ .... 65 3 3 Canopy regeneration in unweeded xate plantation sites ................................ .... 66 3 4 Canopy regeneration in weeded xate plantation sites ................................ ........ 67
9 LIST OF FIGURES Figure page 1 1 Chamaedorea ernesti agustii locally known as xate or fishtail in Belize ............ 30 1 2 Chamaedorea ernesti agustii locally known as xate or fishtail in Belize ............ 31 1 3 Escalating xatero activity in the Chiquibul Forest (1991 2004) ........................... 32 1 4 Leafless C. ernesti agustii due to illegal Guatemalan xate harvest .................... 33 1 5 Life less and brown C. ernesti agustii due to illegal Guatemalan xate harvest ... 34 1 6 with the xate ................................ ................................ ................................ ....... 35 1 7 Xate plantation located west of San An tonio, Belize ................................ ........... 36 1 8 Monoculture xate plantation. This plantation was not sampled in this study ....... 37 1 9 Map of study area in rela tion to the country of Belize. ................................ ........ 38 1 10 Map of study sites within the landscape. ................................ ............................ 39 2 1 Map of study site locations. ................................ ................................ ................ 48 3 1 Understory plant species richness in xate plantations and forest ....................... 55 3 2 Estimation of understory plant species richness in xate plana tations and forest ................................ ................................ ................................ ................. 56 3 3 Mean understory plant species richness ................................ ........................... 57 3 4 vated xate has been removed from the analysis ................................ ................................ ................ 58 3 5 Relative abundance of dominant understory plant species for unweeded xate plantations ................................ ................................ ................................ ......... 59 3 6 Relative abundance of dominant understory plant species for weeded xate plantations ................................ ................................ ................................ ......... 60 3 7 Relative abundance of dominant understory plant species for forest sites ........ 61 3 8 Relative abundance of invasive species, Oeceoclades maculata ...................... 62 3 9 Relative abundance of rare species ................................ ................................ .. 64
10 3 10 Mean percent canopy cover ................................ ................................ .............. 68 4 1 Weeded xate plantation, established west of San Antonio, Cayo, Belize ........... 84 4 2 Unweeded xate plantation, established west of San Antonio, Cayo, Belize ....... 85 4 3 Rustic xate ( Chamaedorea elegans ) plantation in Veracruz Mexico ................. 86
11 Abstract of Thesis Presented to the Graduate School of the University of Florida in Partial Fulfillment of the Requirements for the Degree of Master of Arts XATE PALM ( CHAMAEDOREA SP. ) ENRICHMENT IN WESTERN BELIZE: TH E ECOLOGICAL EFFECTS OF MANAGEMENT IN RELATION TO UNDERSTORY PLANT SPECIES RICHNESS, DIVERSITY, AND COMPOSITION By Carly N. Voight May 2011 Chair: Nigel J.H. Smith Major: Latin American Studies Xate ( Chamaedorea sp p .) a non timbe r forest product (NTF P) in the palm f amily, is declining at a serious rate in various areas of Guatemala due to increased harvest for the floral arrangement industry. Although xate grows in Belize, the country does not currently harvest it in the wild. Due to the decline in Gu atemala, harvesters have turned to the illegal harvesting of xate across the Belizean border. Belizeans claim that Guatemalan xate harvesters are damaging the xate population and the forest in Belize. There is a recent global trend towards establishing pla ntations of previously wild gathered NTFP species, such as aa and Brazil nut, in an attempt to alleviate population level pressure from wild harvest. Xate is one of these species that is currently in a state of transition from wild harvest to cultivation This study addresses the ecological impacts at the community level on NTFPs under cultivation, using xate palm ( Chamaedorea ernesti agustii ) in Belize as a case study. Although xate plantations have been implemented to reduce population level impacts in the wild, the community lev el impacts within these plantations are unknown. The xate plantations in Belize utilize management practices at a relative low intensity
12 level. L andowners retain the natural forest canopy and practice organic agricultural methods The xate plantations in this study actually resemble a secondary growth forest that has been enriched with xate. Currently, there are two different management practices in xate plantations in western Belize: unweeded and weeded. The weeded plantations are the same as the unweeded plantations except that the understory is weeded approximately every six to eight months. I compared unweeded xate plantations, weeded xate plantations, and secondary growth forests in western Belize by assessing the difference in understory plant species richness, diversity, and composition. I found no significant difference in understory plant species richness and diversity between the two xate plantation management types and secondary growth forests. I found rare species in all management types and no s ignificant difference in relative abundance of an invasive species ( Oeceoclades maculata ) between management types. The domina nt species present in forest sites were also domina nt or present at weeded and unweeded xate enrichment plantings T his research p rovides evidence that the management utilized at these xate plantations has minimal negative effects at the ecological understory community level. This suggests that the intensity of management practices employed by the Belizean xate plantation owners in t his study was not considerable enough to lead to significant differences in the understory species community. I n essence this study found that xate enrichment plantings cultivated under a secondary growth canopy in western Belize are sustainable at the un derstory plant community level. Additional research on plantations with greater management intensities should be conducted to establish realistic management level thresholds.
13 CHAPTER 1 BACKGROUND Introduction Tropical forests encompass some of the highes t biodiversity levels and species level endemism i n the world but are currently facing alarming rates of deforestation and degradation in many areas This loss of species has unknown long term effects on ecosystem processes and functions, climate patterns, and the existence of important resources such as medicines and crop relatives ( Anderson et al. 2002 ; Wilson 2010 ). A range of strategies to protect these ecosystem services h as been promoted including the establishment of protected areas and the integrat ion of land protection with economic development, each with their own positive and negative aspects. While protected areas are often touted as being the most effective strategy for conserving biodiversity ( Locke and Dearden 2005 ; Terborgh 1999 ) regulation is often difficult to enforce in tropical countries and critics maintain that reserves will only be sustainable if local communities participate in decision making ( Wells et al. 1992; Wilhusen et al. 2002). Some studies support the idea that sustainable management practices protect ecosystems while supporting human communities socially and economically ( Anderson 1990; Gomez Pompa and Kaus 1990; Peters et al. 1989; Shackleton 2001). However critics maintain that in reality many programs are not actually ec ologically, socially and economically sustainable (Arnold and Perez 2001; Belcher et al. 2005; Kusters et al. 2006). In practice, this should not be a black and white issue since a multitude of solutions are required to combat deforestation and degradation Managed and unmanaged forests should be viewed as complementary ecosystems in a broad conservation scheme (Faria et al 2007 ; Schroth et al. 2004 ).
14 Non timber forest products (NTFPs), such as nuts, fruits, resins and foliage, have been promoted as one com ponent of sustainable tropical forest management that can be an effective means to rainforest conservation and poverty alleviation (Anderson 1990; Plotkin and Famolare 1992; Shackleton 2001). However, as with all sustainable management practices, NTFP mana gement is not without ecological and economic challenges. Even though it is frequently assumed that NTFP extraction does not alter or only slightly alters ecological processes, harvest may produce an impact at various ecological levels. NTFP extraction in the wild can affect the population of the managed species including changes in species growth, survival, reproduction, or regeneration (Hall and Bawa 1993; Salafsky et al. 1993 ; Ticktin 2004 ). The harvest of NTFPs can also have a negative influence at the ecological community level, such as changes in biodiversity or species abundance. Theoretically, NTFP harvest in forests may have negative population and community level effects as compared to forests that are not harvested (Ticktin 2004). Even if NTFPs ar e gathered in an ecologically sustainable manner, local communities may not gain enough income from the resource or consumers may not generate significant demand for a product (Arnold and Perez 2001; Belcher et al. 2005; Kusters et al. 2006). The role of N TFPs is further complicated by the level of management intensity. Plantations of previously wild harvested NTFPs are being implemented in an attempt to alleviate population level pressure from wild harvest (Anderson 1990 ; BBG 2005 ; Carpentier et al. 2000; Endress et al. 2004 ) Plantations may have lesser population level effects but greater negative community level effects as compared to wild harvest (Anderson and Putz 2002; Endress et al 2004) Understanding the impacts of forest
15 management throughout the landscape is critical. Therefore, biodiversity studies should also be conducted in areas such as managed forests and plantations to determine the effectiveness of biodiversity conservation at a landscape scale (Lindenmayer and Franklin 2002; Tarrega et al. 2006) The level of change within the community is most likely directly related to the level of management intensity. The understanding of management intensity levels and their affects on ecological communities is crucial in biodiversity conservation and developing management techniques that are truly sustainable. Xate palm ( Chamaedorea sp p ), which is in a state of transition from wild harvest to cultivation, is a model for understanding these management intensity levels. This study addresses the ecological impacts at the community level on NTFPs under cultivation, using xate palm in Belize as a case study. Although xate plantations have been implemented to reduce population level impacts in the wild, the community level im pacts within these plantations are unknown. The community level effects of various plantation management intensities are also not known. Therefore, I assessed the difference in understory plant species richness, diversity, and composition by comparing unwe eded xate plantations, weeded xate plantations, and secondary growth forests in western Belize. Xate Ecology and Harvest Xate (sha tay) is the common name in Mayan used for several different palm species in the Chamaedorea genus. This study focuses on C ham aedorea ernesti agustii (Figure s 1 1 and 1 2 ) as it is the main species planted in xate plantations in western Belize.
16 Ecology and Distribution of Xate ( C. ernesti agustii ) Chamaedorea is the largest Neotropical palm genus, encompassing between 77 to 100 species, depending on the recognition of horticultural hybrids (Henderson et al. 1995). Chamaedorea species occur in rain forest and cloud forest ecosystems from Central Mexico to Bolivia (Hodel 1992). C. ernesti agustii ranges from the Mexican Atlantic sl ope to Honduras (Henderson et al. 1995). All Chamaedorea species are understory palms and require shade. They prefer well drained calcareous sandy loam soils, abundant organic matter, and limestone bedrock (CEC 2002; BBG 2005). Some xate species, including C. ernesti agustii Chamaedorea species are dioecious in that individuals produce female or male flowers but not both (Henderson et al. 1995). Scienti sts have traditionally considered most palms t o be wind pollinated. W hile some Ar ec aceae genera may be, xate palm species are most likely not since the understory of the rainforest is floristically and structurally not well adapted to wind pollination (Hod el 1992). Most Chamaedorea species are undoubtedly insect pollinated because their flowers are brightly colored, aromatic, and contain sticky pollen (Fisher and Moore 1977). Chamaedorea ernesti agustii palms are pollinated by thrips, tiny insects in the or der Thysanoptera (Porter Morgan 2007). Uses of Xate and the Xate Industry American, Canadian, and European florists have imported xate from Mexico and Guatemala since the 1950s for Palm Sunday church services and foliage in floral arrangements ( CEC 2002 ; H odel 1992 ) because of their attractiveness and ability to remain green and vibrant for up to 45 days after cutting (BBG 2005) Fourteen percent
17 of the U.S. cut greens industry market is comprised of Chamaedorea leaves (Wilsey and Endress 2007). Entire seed lings are also sold for landscaping and as houseplants (BBG 2005) Chamaedorea palms were not traditionally used by the ancient Maya in Belize, Guatemala, or Mexico (Schlesinger 2001). Considering that most harvested Chamaedorea species are insect pollinat range requires hand pollination or the purchase of naturally raised seeds and therefore is generally not considered economically viable. Furthermore, Chamaedorea products have a low production cost and a low m arket price. Therefore, the majority of xate utilized outside of Latin America has been imported instead of grown domestically (CEC 2002). Wild x ate harvest in Mexico and Guatemala Consequently, xate harvest has contributed to the income of those living n ear or in tropical forests in Mexico and Guatemala. The total xate harvest process, including collectors and sorters, provides over 10,000 full time and seasonal jobs in Mexico (Vovides and Garcia Bielma 1994) and about 6,000 to 10,000 jobs in Guatemala (R ainforest Alliance 2004). Chamaedorea leaves are the second most commercially Reserve, xate harvest is the main source of income to the surrounding local communities (Radacho wsky and Ramos 2004). I n general the palms have been intensively harvested in these natural forests without regulation (CEC 2002). People who harvest xate from wild populations, or xateros as they are locally called, are paid by the number of leaves colle cted as opposed to the quality of the leaves. Sixty to seventy six percent of the leaves harvested are discarded due to the lack of industry quality standards ( Radachowsky
18 and Ramos 2004). Xate palms are needlessly damaged since leaves that are unusable by the industry but physiologically important to the plant are collected. Chamaedorea species are threatened due to this overharvest and the destruction and degradation of their rainforest habitat (Johnson 1997; Vovides and Garcia Bielma 1994). Of the forty seven species of Chamaedorea found in Mexico, twenty one are commercially important (CEC 2002). Of these twenty one, three are endangered and thirteen are considered threatened in Mexico, including C. ernesti agustii (Secretar a de Medio Ambiente y Recurs os Naturales 2001). In Uaxactun, Guatemala the density of C. oblongata palms have decreased by over two percent for adults and over thirteen percent for juveniles in only one year (Radachowsky and Ramos 2004). Xate harvesting in locations that are considered to be sustainably managed (Bridgewater et al. 2006). In several areas in Guatemala, xate palm individuals have ceased flowering or fruiting, possibly due to expending en ergy on leaf production (Radachowsky and Ramos 2004). This decline in xate palm populations in Guatemala and Mexico has led to four events: ( 1 ) harvest regulation attempts; ( 2 ) the illegal harvest of xate by Guatemalan xateros in Belize; ( 3 ) the promotion of enrichment planting in wild populations; and ( 4 ) the promotion of xate plantations. The government agency responsible for xate harvest regulation, Consejo Nacional de Areas Protegidas (CONAP), has attempted to improve collection methods. Xateros must on ly collect high quality palm fronds and only two leaves should be cut per plant per year (Radachowsky and Ramos 2004). However the enforcement of this regulation is almost impossible since CONAP only employs one NTFP technician (Bridgewater et al. 2006).
19 W ild x ate harvest in Belize Twelve Chamaedorea species are found in Belize, of which four are considered C. elegans C. ernesii augustii C. oblongata and C. tepejilote Belize has not traditionally utilized xate as a NTFP until recently. D ue to the decrease in the population of Chamaedorea species in Guatemala and the high abundance of C. ernesti agustii in Belize individual Guatemalans began crossing the border into Belize to illegally harvest xate (Bridgewater et al. 2006). Guatemalan xa tero activities were first noticed in 1991, about 5 kilometers from the border (Figure 1 3 ; Bol 2007 pers comm. ). By 2000, the staff at the Las Cuevas Research Station noticed Guatemalan xateros activity near the research station, which is about 20 kilom eters form the border (Bridgewater et al. 2006). Not only is xate extraction illegal since the palm products grown in Belize are smuggled over the border and sold in Guatemala, but they are also generally harvested in protected areas. The majority of the illegal xate activity occurs in the Chiquibul National Park and Chiquibul Forest Reserve. Chiquibul National Park is strictly managed for conservation and no economic activities are allowed. NTFP extraction, as well as timber harvest, is permitted in Chiqu ibul Forest Reserve. However no xate harvesting licenses that were issued have been activated. A ll of the xate harvested within this area therefore, has been illegal (Bridgewater et al. 2006). Currently, the problem has become more severe and pervasive As of 2005, an estimated 600 700 Guatemalans were illegally harvesting xate in Belize (BBG 2005). Guatemalan xatero activity had been noticed in the Manatee Forest Reserve as of 2008, 60 kilometers from the Guatemalan border (Bol 2007 pers comm. ). Furth ermore,
20 the illegal xateros are constantly observed in Chiquibul and their footpaths and horse tracks have become permanent trails (Bridgewater et al. 2006). Chamaedorea ernesii augustii is currently the only xate species illegally harvested in Belize, mo species (Bridgewater et al. 2006). Although many protected areas in Belize, such as Elijo Panti National Park and Cockscomb Basin Wildlife Sanctuary, appear to contain healthy C. er nesii augustii plants, many individuals of this species in Chiquibul are leafless (Figure 1 4 ) or even brown and near death as of 2008 (Figure 1 5 ; personal observation). Research estimates that 37.8 million leaves of C. ernesii augustii have been harveste d from the Chiquibul Forest Reserve from 1999 to 2004, worth U.S. $0.3 million to the xateros. As of 2004, about 75 percent of C. ernesii augustii palms in Chiquib ul Forest Reserve had been cut (Bridgewater et al. 2006) Repeated harvesting can result in a decrease in size of recently produced leaves, a decrease in the health and size of flowers and fruits, and a rise in mortality (Porter Morgan 2004). Harvested C. ernesii augustii palm individuals averaged about 2.5 leaves cut per plant (Bridgewater et al. 2006). Wicks (2005) found that C. ernesii augustii leaf production rates are about 1.56 new leaves annually per plant in Chiquibul. As a result, the illegal In addit ion to damaging the Chamaedorea population s xateros harvest other wild plants from forests and protected areas for food and shelter. They hunt a variety of animals for food such as peccaries, deer, kinkajous, tapirs, guans and tinamous (BBG 2005) The illegal xate harvest exemplifies how NTFP extraction can lead to a degradation of the resource. This extreme example most likely occurred because the
21 xateros do not live in communities near the xate they are harvesting. They feel no sense of ownershi p, responsibility, or stewardship to the land and they have no incentive to protect the resources within it. In fact, some Guatemalans may even feel they have a right to the xate resource since they refuse to acknowledge the Belizean border and claim a por The Belize Guatemala border dispute is a historical disagreement that is still heatedly debated today. The border between Belize (then British Honduras) and Guatemala has been in question since the colonial era. Great Britain, which at the time governed Belize, and Guatemala signed a treaty in 1859 establishing the current boundaries between the two countries. The British were required to construct a cart road between Guatemala and the Caribbean Sea in order for the treaty to be valid ( Clegern 1958) Guatemala declared that the treaty was broken in 1940 since Great Britain never built the road. Guatemala even claimed that Belize was a part of the Guat emala threatened to invade British Honduras on numerous occasions but British military presence has prevented the invasions ( Perez et al. 2009). Although Guatemala autonomy fr om Great Britain, the Guatemalan government still claims over half of Present day conflicts in addition to illegal xate harvest include illegal settlements, illegal farming, ambush attacks, and murder. The remoteness of t he border at Chiquibul Forest Reserve, lack of personnel, lack of finances, and high danger of armed Guatemalans are barriers to enforcement by the Belize Defense Force ( Perez et al.
22 2009). Considering the population of Guatemala is over three times that o f Belize and their large, well (Berman 2007), Belize police are hesitant to control illegal activity, including illegal xate harvest because of the potential threat of war ( Perez et al. 2009) have gained interest in the possibility of xate harvest as an NTFP in Belize. The Government of Belize created a Xate Technical Committee consisting of the Ministries of Na tural Resources, Agriculture, and Foreign Affairs to determine the feasibility of a sustainable xate harvest industry in Belize. Separately, the Darwin Initiative (a United Kingdom based agency) funded a four year grant to the Natural History Museum, Londo n to research xate in Belize (BBG 2005). Research on the long term effects of xatero activities and the abundance and health of wild Chamaedorea species in Belize (e.g. Bridgewater et al. 2006; Pickles 2004; Porter Morgan 2005; Wicks 2005) result ed from t his grant (Bridgewater et al. 2007). The Belize Forest Department has established xate harvest regulations, based upon the Darwin Initiative research results. These regulations include: (1 ) only one leaf per individual palm per year may be harvested; ( 2 ) o nly adult palms may be harvested; ( 3 ) the manager of the area must create and implement a management plan that includes annual permissible cuts based on inventory data; ( 4 ) the harvesting area should not be entered more than once per year to reduce disturb ance; ( 5 ) the xateros must be paid based on the quality of cut leaves, not quantity; and ( 6 ) only market quality leaves may be harvested from plants. However these regulations cannot be
23 implemented in the majority of natural forest areas in Belize due to i llegal harvesting (Bridgewater et al. 2007). In areas that xate is being illegally harvested, it is impossible for the Belize Forest Department to regulate the sustainability of the harvest as well as for the manager to follow a sustainable management plan The manager will have no incentive to abide by a sustainable harvesting regime unless personal benefits exist for said manager (Bridgewater et al. 2007). Considering the current and potential future pervasiveness of illegal xate harvest, xate plantations are being promoted as a way to conserve wild xate populations and effectively monitor sustainable xate harvest. Shade grown NTFP plantations Xate is one of many previously wild gathered NTFPs such as aa and Brazil nut, which are in a state of transition towards c ultivation ( Bridgewater et al 2006 ; Smith et al 1992; Ticktin 2004 ). These plantations are being promoted as alternatives to deforestation because they alleviate population level harvest pressure (Anderson 1990; BBG 2005 ; Carpentier et al. 2000; Endress et al. 2004 ) The sustainable wild harvest of xate in Belize is most likely not possible. Even if the illegal harvest is controlled in the future, it most likely has caused significant long term damage to the Chamaedorea population (Bridgewater et al. 2007). However, the ecological impacts of NTFP cultivation are relatively unknown despite interest in NTFPs for sustainable development and concerns about over over three quarters of the seventy papers surveyed focused on the population level. Only twelve percent of the papers focused on the community level. Of these investigations, the majority focused on the ecological consequences in respect to
24 mammalian, avi an and entomological diversity and population alterations. Only about six percent of the research Ticktin surveyed focused on the impact of NTFP gathering upon plant diversity and abundance. Very few studies have been conducted on NTFPs that are currently harvested in plantations at the community level (Trauernicht and Ticktin 2005), the majority being cacao plantations (Rice and Greenberg 2000). As a NTFP becomes commercialized, the system in which it grows usually becomes increasingly altered in order to enhance the production of the NTFP. Differences in community condition between managed and unmanaged forests are most likely directly related to the level of anthropogenic intensity. The majority of biodiversity in the tropics exists in areas managed and i nhabited by humans (Alcorn 1995) and is essential for human and economic well being. Therefore the conservation of biodiversity should not only be implemented in protected areas but also throughout the landscape (Lindenmayer and Franklin 2002; Tarrega et a l. 2006). Consequently, biodiversity studies should also be conducted in areas such as plantations to determine the effectiveness of biodiversity conservation at a landscape scale (Faria et al 2007). In order to uphold NTFPs potential for biodiversity cons ervation and to develop sustainable management techniques, an understanding of management intensity and their affects on ecological communities is crucial. Plantations may have lesser population level effects, such as greater growth and survival rates of t he cultivated NTFP, as compared to wild harvested forests. However plantations may contain harsher negative community level effects, such as less biodiversity and fewer species as well as an altered composition of species, as compared to natural forests or wild harvested forests (Anderson and Putz 2002;
25 Endress et al 2004) Some plantations contain fewer forest specialists and more early successional species than unmanaged forests ( Alves 1990 ; Leston 1970; Room 1975 ). Epiphytes and hemiparasites are removed in some plantations, significantly affecting important microhabitats of ants and other invertebrates (Room 1971; Young 2007) as well as food for birds and butterflies ( Greenberg et al. 1997). Canopy regeneration is an important factor in the plant communi ty. In some NTFP plantations, the biotic and abiotic conditions for regeneration are not met and it is effectively eliminated. In these situations, trees that naturally occur in the forest overstory are gradually replaced by other species in the plantation s (Rice and Greenberg 2000). The type of trees that replace natural forest species usually relates to management practices implemented by plantation owners. Plantation owners may plant specific trees or selectively weed all understory plants except for spe cific tree seedlings usually trees that provide optimal shade, fix nitrogen, or are economically useful which may eventually replace the natural forest canopy. Shade grown xate plantations in Belize In contrast to other NTFPs under cultivation, the ma nagement intensity of xate plantations in western Belize is relatively low. During the establishment of xate plantations, landowners retain the natural forest canopy for this shade dependent species (Bridgewater et al 2006) and practice organic agricultura l methods (BBG 2005). While the natural overstory of most coffee and cacao plantations is routinely thinned and pruned, the canopies of xate plantations in this study are not. Epiphytes and hemiparasites are not removed. Currently, two different management practices exist in xa te plantations in western Belize : unweeded and weeded. The unweeded and weeded plantations are essentially
26 secondary growth forest (20 30 years old) in which the farmer clears the forest u nderstory when the plantation i s established a nd planted xate in t he understory. The understory i s allowed to regenerate naturally. The weeded plantations are the same as the unweeded plantations except that the understory is weeded approximately every six with a machete in which the plant is cut at the forest floor instead of removing or rooting out the plant. Additionally, not all of the plants are cut in a plantation. Plantation owners stated that they only cut the plants that with the xate (Figure 1 6 ) Therefore, while the weeded plantations have less understory plant life as compared to the unweeded plantations, sampled in this study actually resemble enrichment plant ings (Figure 1 7) instead of monoculture xate plantations (Figure 1 8) Considering these plantings are called xate plantations in Belize but are Belize has not utilized xate as an NTFP until recently. Although Guatemalans have been able to garner full or part time employment from xate harvest production, Belizeans have never regarded it as a feasible income source beca average standard of living is considered to be much lower than a typical Belizeans (Bol 2007, pers. comm.; Mesh 2008, pers. comm. ). According to a World Bank (2010) study, the per capita income in 2009 of Guatemala is about U.S. $4,570 whil e Belizeans receive about U.S. $5,990. Xateros earn about U.S. $5.15 per day in Guatemala ( Radachowsky and Ramos 2004 ). Considering the large amount of time wild xate harvest requires, Belizeans have never esteemed it to be an economically viable
27 activity. However, xate plantation management requires much less time commitment and therefore small landowners are currently considering it as a viable investment for supplemental income ( Bol 2007, pers. comm.; Mesh 2008, pers. comm. ). The Darwin Initiative xate g rant also funded research on Chamaedorea species germination and plantation management techniques. They awarded money to the Belize Botanic Gardens to give xate seedlings and xate plantation management trainings to local Belizeans (BBG 2005). The Belize Bo tanic Gardens has collaborated with several small community based organizations, such as the Itzamna Society, to assist in working with local farmers (Mesh 2008 pers comm. ). The Itzamna Society has helped over twenty farmers in San Antonio, El Progresso 7 Miles, and Cristo Rey establish plantations of 40,000 seedlings total (Bridgewater et al. 2007). All of the data for this research w ere collected within the xate enrichment plantings that resulted from the assistance of the Darwin Initiative, Itzamna Soc iety, and Belize Botanic Gardens. Landscape and Cultural Setting This study was conducted within the Cayo District of Western Belize (Fig ure 1 9 ). The xate plantations and adjacent forest sites in this study are located near the villages of San Antonio and El Progresso 7 Miles, which lie south of the town of San Ignacio (Figure 1 10 ) Presently, this landscape consists of secondary growth forests (10 75 years old), cleared agricultural areas, and small towns (Garcia 2002). The annual rainfall in t he area t otals about 1 524 mm with a wet season from June to October and a dry season from November to May (Tour 2006). The predominate soil parent material is limestone, which forms rolling hills in this region. The soil is relatively fertile from the accumulatio n of organic matter in limestone cracks and the weathering of mineral soil (Meerman and Sabido 2001).
28 The forest surrounding San Antonio is considered subtropical moist forest, as classified by the Holdridge (1967) life zone system. Meerman and Sabido ( 2001) further classified Belize into 85 ecosystem types using Landsat imagery and field surveys. The The l shifting, low intensity agriculture in Belize around 3000 2000 BC. From about 500 BC to 900 AD, the Maya transitioned to intensive agriculture with short fallow periods in order to sustain their elaborate civilization (Tour 2006). Pacbitun, an ancient Mayan site located about three kilometers east of San Antonio, was occupied by the Maya from 90 0 BC to 900 AD (Healy 1990). S everal ancient Mayan sites Mayan Ranchos, Cristo Rey Tipu Negroman, and Macaw Bank lie within a nine k ilo m eter radius of Pacbitun (Graham et al. 1985). The area surrounding the Mayan sites was farmed through slash and burn ag riculture and hillslope terracing (Healy 1990). The majority of the lands used by the ancient Maya are the same lands preferred by farmers today (Tour 2006). The area is prone to natural disturbances such as hurricanes and fires. Furthermore, the forests within this study site have been logged for mahogany and cedar sporadically for the last 300 years, as is the case with forests throughout Belize (Primack et al. 1998). Several protected areas have been established near San Antonio and El Progresso 7 Miles Noj Kaax Meen Elijio Panti National Park, a reserve of 5,263 hectares contains hardwoods of about 75 years old (Garcia 2002). The name noj kaax or large forest in Yucatec Maya, prefaces the name of the park to stress the sacred
29 forest (Manzano and Rosa do 2007). The park is named after Don Elijio Panti, a world renowned Mayan healer, or meen in Yucatec Maya. Meen Elijio Panti was from San Antonio and passed away in 1996 at the age of 103 (Garcia 2002). While small in terms of size, Elijio Panti National Park is a part of a large patchwork of connected protected areas colloquially known as the Mayan Mountains. The Mountain Pine Ridge Forest Reserve ( 51,324 hectares ) and Chiquibul National Park and Forest Reserve ( 184,924 hectares ) are located south of Eli jio Panti National Park and are the three closest protected areas to San Antonio and El Progresso 7 Miles ( Berman 2007; Bridgewater et al. 2006). The people who live within this landscape are primarily Yucatec Mayan and mestizo The majority of the 2,350 people who live in San Antonio are Mayan (Berman 2007). Pacbitun was abandoned around 900 AD (Healy 1990). T he Maya who currently live in San Antonio are not descendents of the Maya who previously lived in the region but are actually descended from the Ma ya of Quintana Roo, Mexico. They are the ancestors of immigrants who fled from the Yucatan Penensula of Mexico in 1842 (Garcia 2002). The majority of the people who live in El Progresso 7 Miles are mestizo immigrants from Guatemala (Mesh 2008, pers. comm. ) Ninety percent of the people who live in the region surrounding San Antonio and El Progresso 7 Miles are farmers (Garcia 2002). The landscape is dotted by milpas of corn, beans, tomatoes, and peanuts; productive forests managed for xate, calhone palm ( At talea cohune ) and sabal palm ( Sabal mauritiiformis ) ; and patches of protected forests managed for tourism and conservation (personal observation).
30 Figure 1 1. Chamaedorea ernesti agustii locally known as xate or fishtail in Belize. Photo is taken at Co c k scomb Basin National Park Carly Voight
31 F igure 1 2 Chamaedorea ernesti agustii locally known as xate or fishtail in Belize. Photo is taken at Caracol Archeological Park Carly Voight
32 Figure 1 3 Escalating xatero ac tivity in the Chiquibul Forest Belize (1991 2004) 2004 Guatemala Belize
33 Figure 1 4 Leafless Chamaedorea ernesti agustii in Chiquibul Forest Reserve, Belize due to illegal Guatemalan xate harvest Carly Voight
34 Figure 1 5 Life less and brown Chamaedorea ernesti agustii in Chiquibul Forest Reserve, Belize as a result of illegal Guatemalan xate harvest Carly Voight
35 Figure 1 6 with the xate. After investigation, t he farmer weeds these sel ect plants by cutting them at the forest floor with a machete. This plantation is located west of San Antonio Belize (Location A, Figure 2 1). Carly Voight
36 Figure 1 7 Xate plantation located west of San Antonio Belize (Lo cation B Figure 2 1). Carly Voight
37 Figure 1 8 Monoculture x ate plantation This plantation was not sampled in this study. Natural History Museum, London
38 Figure 1 9 Map of study area in relation to the country of Beli ze Source of aerial photograph : Color satellite image mosaic for Belize, prepared by the Division of Applied Research & Development of the Water Center for the Humid Tropics of Latin America and the Caribbean (CATHALAC) f over Belize on March 28, 2 000 (Path 19, Rows 47 49). This dataset is in 16).
39 Figure 1 10 Map of study sites within the landscape Source of aerial photograph : Color balanced L satellite image mosaic for Belize, prepared by the Division of Applied Research & Development of the Water Center for the Humid Tropics of Latin over Belize on Ma rch 28, 2000 (Path 19, Rows 47 49). This dataset is in San Ignacio San Antonio El Progresso 7 Miles
40 CHAPTER 2 RESEARCH HYPOTHESES AND METHODOLOGY The goal of my research is to better understand the ecological effects of xate plantation management in Bel ize. This chapter presents my research hypotheses and methodologies that were used to determine any differences in plant species richness, diversity, or composition among the study sites Research Hypotheses My study assesses the difference in plant specie s richness, diversity, and composition by comparing different xate enrichment management schemes and secondary forests containing natural xate populations (as a control). Considering that environmental factors (topography, slope, soil, etc) and previous la nd use are similar among the three areas: H1. Considering the level of understory clearance and compaction, plant richness should decrease in the following order: unmanaged secondary growth forest sites, unweeded xate plantations, and weeded xate plantatio ns. Since these xate plantations resemble enrichment plantings, the severity of the decrease is unknown. H2. growth forest sites, unweeded xate plantations, and weeded xate plant ations. H3. There should be a change in composition. For example, early successional species and exotic species should increase in the following order: unmanaged secondary growth forest sites, unweeded xate plantations, and weeded xate plantations. F orest dependent species and rare species should decrease in the above order.
41 Research Methodology This study compares unmanaged secondary growth forests with xate enrichment plantings of two different management types: weeded and unweeded. The unweeded and weede d understory plantations both have a natural secondary growth forest cover. The weeded plantations are similar to the unweeded plantations except that the understory is weeded approximately every six to eight months. All of the weeded plantations in this s tudy were weeded approximately five months prior to data collection. While the weeded xate enrichment plantings have less understory plant life as compared to the unweeded xate enrichment plantings they are not denuded. R efer to Chapter 1 ( p. 26) for more info rmation on xate plantation management practices. Experimental Design Considering that the plantations are very small ranging from 0.1 to 2 hectares long transects were not possible. I established a n area of 30 m x 30 m in each xate enrichment area and unmanaged forest area to account for this difference among the plantations All plots were placed within th ese 30 m x 30 m areas I established fifteen sites in total five sites were placed in the adjacent natural forest that surrounds the xate plantati ons, five sites were established in weeded plantations, and five sites were placed in unweeded plantations. The sites are similar in terms of elevation, slope, aspect, age, rainfall, soils, previous land use and surrounding land use All fifteen sites ar e located in western Belize, near the villages of San Antonio and El Progresso 7 Miles, which lie south of the town of San Ignacio ( Figure 2 1). All of the xate enrichment plantings utilized in this study were established with the assistance of the Belize Botanic Gardens and the Itzamna Society and funded by the Darwin Initiative
42 Grant. Therefore, all of the plantations were established during the same time period, with genetically similar xate seeds, and under similar management guidelines (except for the use of weeding). Consequently, the xate in all of the enrichment plantings in this study are about the same age and height. Each location is described below. Since natural forest sites were established adjacent to the xate plantation each location contains more than one site. Description of specific site l ocations Location A includes five sites northwest of San Antonio. Site W1, W2, and W3 are weeded xate plantations separated from each other by strips of secondary growth forest. Site U4 is an unweeded xa te plantation. Site F1 is an unmanaged secondary growth forest control area. Location B includes two sites northeast of San Antonio. Site U2 is an unweeded xate plantation and Site F2 is an unmanaged secondary growth forest control area. Location C inc ludes three sites south of El Progressive 7 Miles. Site U3 and U5 are unweeded xate plantations owned by two different farmers and Site F3 is an unmanaged secondary growth forest control area. Location D includes two sites northwest of San Antonio. Site W4 is a weeded xate plantation and Site F4 is an unmanaged secondary growth forest control area. Location E includes three sites northwest of San Antonio, established at the Belize Botanical Gardens. Site U1 is an unweeded plantation, W5 is a weeded plant ation and Site F5 is an unmanaged secondary growth forest area. Data collection and species identification Ten 1x1 m plots were situated randomly within each 30 m x 30 m site. To avoid confounding edge and tree fall gap effects, no plot was placed within 5 m of the
43 plantation edge and within 10 m of a tree fall gap edge All vascular plants less than 1.30 m tall within sampling plots were identified to species level, following Beattie and Oliver ( 1994 ) Scandents were recorded as long as the understory clim bers, vines, or lianas were similar to understory herbs, shrubs or tree seedlings in that they were rooted in the soil and were less than 1.30 meters. For clonal species such as some grasses, each clump was recorded as one individual. According to various studies, this methodology provides meaningful understory plant species richness data ( Beck et al. 2002; Schulze et al 2004) In order to estimate percent canopy openness, I recorded the percent canopy cover in the center of every plot in all management types with a densiometer. Densiometers have been found to be the best tool for accurate percent canopy cover measurements that are also time efficient ( Korhonen et al. 2006) I identified as many species in the field as possible. Belizean field botanists who have been trained by New York Botanical Garden assisted with the field identification. I collected, dried, and pressed voucher specimens of every species recorded, except for Convention on International Trade in Endangered Species ( CITES ) listed and pr otected species. No CITES species were collected nor imported into the United States. The collection was transported to the University of Florida. I spent a considerable amount of time identifying as many specimens to the species level as possible. To iden tify specimens, I used a variety of taxonomic keys and floras ( Balick et al 2000 ; Hamilton 1989; Parker 2008; Standley and Steyermark 1958; Swallen 1955), the University of Florida herbarium, and a number of online herbariums that specialize in Neotropica l specimens (New York Botanical Garden, Missouri Botanical Garden, The Field Museum of Chicago Instituto Nacional de Biodiversidad, and University of Michigan ). T he
44 majority of the collected specimens were sterile and many were seedlings and young sapling s. Also some of the seedlings were morphologically different than adults because they were immature. Therefore some specimens could only be identified to genus or plant family. In these cases, the specimen was grouped into morphospecies within the genus or family (i.e. Mikania sp. #1 or Rubiaceae #2). Specimens that were unable to be identified to family were sorted to morphospecies (i e indet #1). This methodology has been useful for sterile specimens in various other plant species diversity studies ( Gree nberg et al 1997 ; Kessler et al 2005; Schulze et al 2004 ) Dr. Walter Judd, a plant systematist and professor, confirmed all of my identifications and assisted in the identification of specimens I was unable to identify. Several specialists assisted in ide ntifying or identification confirmation, as listed in the acknowledgements section. All species encountered in this study are presented in Appendix A. For several pairs of morpho species, sterile individuals could not be distinguished and therefore they wer e treated as one species in the analyses. For example, the vine Paullinia pinnata has five leaflets. Individuals that were definitely identified as Paullinia pinnata were recorded as Paullinia pinnata However other individuals that closely resembled Paull inia pinnata and did not closely resemble any other vine species of Sapindaceae found in Belize but had fewer than five leaflets (most likely because they were immature ) were recorded as Paullinia cf. pinnata Considering the individuals were most likely t he same species, they were treated as the same for the species richness, diversity and composition analyses.
45 Data Analysis Species richness H1. By what degree does species richness decrease within the xate plantation sites as compared to unmanaged forest sites? The number of species was recorded within all 1x1 m plots established. The number of species observed strongly relates to the number of collected individual plants, especially within tropical communities (Magurran 2004). Care must be taken in compa ring richness samples to avoid pitfalls such as sample size, sampling effort, and size of surveyed plots (Gotelli and Colwell 2001). Most richness data sets are typically not inclusive in tropical forest field surveys. Consequently, an attempt at evaluati ng diversity between different locations encounters the predicament that an unknown number of species were not collected in the sample. A number of species richness estimators have been designed to compensate for this limitation, all with their own set of drawbacks and assumptions (Colwell and Coddington 1994). I estimated the true species richness using a computer program called EstimateS, which has been effective in estimating species richness in over 1,800 studies (Colwell 2009). This program computes se veral different species richness estimators including Chao 1, Chao 2, first order jackknife, second order jackknife, bootstrap, and Michaelis Menten. Chao 2 and second order jackknife have shown to be the least biased species richness estimates for studies with small sample sizes (Colwell and Coddington 1994). However, EstimateS does not calculate the standard deviation for seco nd order jackknife (Colwell 2009 ). In correlating the results from the species richness estimators Chao 2 and second order jackkni fe, I found the results to have a 0.86 correlation (p < 0.00001). As the
46 results between Chao 2 and second order jackknife appear to have a strong correlation, I decided to only present the results from Chao 2 in the main text and figures All species rich ness estimator results are presented in Appendix B Chao 2 richness estimation data met the ANOVA assumptions of homogeneity of variances and normality. I used SPSS to perform a one way ANOVA with species richness as the dependent variable following Schulz e et al (2004). Species diversity H2. By what degree does species diversity decrease within the xate plantation sites and as compared to unmanaged forest sites? The number of species and number of individuals within each species were recorded within all 1x 1 m plots established. The two most common diversity indices are Shannon and Simpson (Hill et al 2005). Although the Shannon index is the most commonly used index, it has many disadvantages. It is based on the assumption that all species are represented in the sample plots and all individuals are randomly sampled from an infinitely large community. Conversely, the Simpson index is insensitive to sample size. Therefore, I used the Simpson index to calculate diversity due to its ability to determine the varia nce of species abundance distribution (Magurran 2004). Simpson diversity index data met the ANOVA assumptions of homogeneity of variances and normality. I compared the values of the Simpson diversity index between the three management types using a one wa y ANOVA Species composition H3. What is the change in plant species composition (i e exotics, rare species, dominate species) between plantations sites and unmanaged forest sites?
47 I calculated the relative abundance of each species (the proportion of th e number of individuals within each species to total individuals; see Krebs 1999 ) within each management type. The composition of several categories such as dominant species and rare species was compared graphically. The relative abundance of exotic specie s met the ANOVA assumptions of homogeneity of variances and normality. I compared the relative abundance values of nonnative species between the three management types using a one way ANOVA. An in depth canopy regeneration study is beyond the scope of thi s study. Although I was not able to gather detailed data on all of the canopy species present, I did record the common overstory species that I was able to identify in the field. The relative frequencies of the understory species recorded were numerically compared with the presence of common overstory species. Canopy Cover The recorded percent canopy cover data met the ANOVA assumptions of equality of variances and normality. I compared the percent canopy cover between forest sites, unweeded xate plantation s, and weeded xate plantations using a one way ANOVA.
48 Figure 2 1. Map of study site locations (see p. 40 41 for location descriptions) Source of aerial photograph : Color satellite image mosaic for Belize, prepared b y the Division of Applied Research & Development of the Water Center for the Humid Tropics of Latin over Belize on March 28, 2000 (Path 19, Rows 47 49). This dataset is in rd projection (NAD27, Zone 16).
49 CHAPTER 3 RESEARCH RESULTS Species Richness I found 379 species, representing 73 families in my 15 study sites Fabaceae was the most diverse family, followed by Rubiaceae and Sapindaceae ( Appendix A). Individuals that cou ld not be identified to any taxonomic family were sorted into twenty seven morphospecies based on stem, leaf, and root traits Results from the species richness estimator Chao 2 showed that understory species richness increased in the following order from lowest to highest: weeded xate enrichment plantings unweeded xate enrichment plantings and secondary growth forest sites ( Figures 3 1 3 2 and 3 3 ). However this difference w as not significant (p = 0.474). Species Diversity function of richness and evenness, was highest in the forest sites, followed by unweeded xate enrichment plantings and weeded xate enrichment plantings A one way ANOVA found the difference to be significant (p = 0.047). Since the richness was not signific ant between the sites, the sites varied due to a difference in evenness. Once planted Chameadorea ernesti agustii was removed from 0.333 ; Figure 3 4 ). These findings i mply that management techniques such as weeding or clearing the understory prior to planting xate did not appear to result in a significant difference in diversity Instead, the significant difference in diversity is most likely related to the mere presenc e of planted C. ernesti agustii.
50 Species Composition Plant Species Dominance Graphs representing the dominant understory plant species for all areas are presented in Figures 3 5, 3 6, and 3 7. All of the dominant understory plant species found in the secon dary growth forest sites were either dominant in both xate plantation management types (such as Rourea glabra Cupania belizensis and Paullinia cf. costaricensis or Psychotria sp. #2) or present (such as Psychotria tenuifolia ). Conversely, all of the domi nant understory plant species found in the unweeded xate plantations were also dominant or present in the for est areas, such as Chamaedorea ernesti augustii Chrysophyllum mexicanum and Paullinia pinnata The same is true for the dominant understory plant species found in weeded xate plantations, such as Chamaedorea ernesti augustii Piper jacquemontianum and Acacia sp. #1, which were also dom inant or present in the forest areas. Non Forest Species The majority of the species observed in this study for w hich information existed on their distribution are recorded as species characteristically encountered in lowland tropical forests of Belize and the Peten, Guatemala. I found species in both the forest sites and xate enrichment plantings that are: ( 1 ) commo nly found in both forest and open areas but are more typical of disturbed areas such as Oplismenus hirtellus (Swallen 1955) and Bidens squarrosa ( Meerman and Sabido 2001); ( 2 ) generally restricted to wet and moist forests such as Odontonema tubaeforme (Dan iel 2010), Gaussia maya Astrocaryum mexicanum and Protium copal (Parker 2008); or ( 3 ) typically encountered in secondary growth forest such as Bursera simaruba Piper jacquemontianum and Metopium brownei (Parker 2008). However, the majority of
51 species f ound in this study primarily occur in lowland rainforest both secondary growth forests and forests (those that have not been clea red in approximately 75 years) This is not surprising considering that xate plantations were carved out of secondar y growth forests and are located within a landscape of unmanaged forests, managed forests and agricultural areas. Meerman and Sabido (2001) classified Belize into 85 ecosystem types using Landsat imagery and field surveys. The natural ecosystem type that was identified for common woody plants of the vegetation classification for the are a, such as Attalea cohune Cupania belizensis Cordia sp p ., Pimenta dioica, Pouteria sp p ., Sabal mauritiiformis Protium copal, Spondias mombin, and Trophis racemosa This vegetation type lists palms, lianas, and shrubs in the Rubiaceae famil y as common, c onsistent with my findings in both forest and xate plantation sites. A comparison of forest specialist species verses early successional species between sites cannot statistically be addressed in this study. Not all of the species in this study could be i dentified to the species level. As a result, they could not be classified into categories such as forest specialist, forest generalist or disturbance generalist. The only non forest species encountered in this study that was identified to the species lev el is Oeceoclades maculata an aggressive nonnative terrestrial orchid. This species, which is native to Africa (Stern 1988), was found in both xate plantation and forested sites. Oeceoclades maculata was most abundant in unweeded plantations,
52 followed by forested sites, and w eeded plantations (Figure 3 8 ). However there was no significant difference in percent relative abundance (p = 0.301). Rare Species A comparison of rare species between sites cannot statistically be addressed in this study. All of the species richness estimators in EstimateS computed drastically higher species richness estimations than my observed richness and therefore I most likely did not record all of the species that are present in a site. Rare species are typically sensitive to s ampling effort. As a result, it can be assumed that if a rare species was present at a site, it may not have been recorded. The rarity status for recorded rare species is presented in Table 3 1 and the relative abundances observed for certain rare species are presented for discussion in Figure 3 9 Rare species were found in weeded xate plantations, unweeded xate plantations, and forest sites. Gaussia maya was observed three times more frequently in forested areas. I could not test for a possible significa nt difference in relative abundance of Gaussia maya between management types due to the reasons mentioned above Red cedar ( Cedrela odorata ), which is categorized as vulnerable due to timber harvest exploitation, was only recorded in forested areas. Howeve r since it was observed extremely infrequently, it might also be pr esent but not recorded in xate enrichment sites Calophyllum brasiliense, a species that is threatened in Guatemala, was only found in xate plantations. As is the case with red cedar, since C. brasiliense was found very infrequently in the xate enrichment plantings it might also be present in the forest but not recorded. Zamia cf. polymorpha could not be confirmed to the species level because all individuals found in the field were sterile and I could not collect it due to its highly
53 endangered status. There are three species of Zamia in Belize, all of which are rare. The IUCN Red List considers the status of Zamia polymorpha Zamia prasina Zamia variegata Considering that Zamia prasina is endemic to southern Belize and the only area Zamia variegata is found in Belize is along the coast (IUCN 2010), the species that I encountered was most likely Zamia polymorpha. Z amia cf. polymorpha was the most commonly observed rare species in all sites. It was most frequent in forest sites. The individuals of Louteridium chartaceum found in this study are a new population of this species. This rare species, endemic to Belize, h as previously only been found on limestone hills in the southern part of the Belize District (Daniel 1997). This population is the first found in western Belize (Cayo District), located about 70 km from any known population. It was only found at one xate p lantation and one unmanaged forest site. Thomas Daniel, an expert on Acanthanceae in Belize, confirmed the identification of the specimen. Louteridium chartaceum was observed more frequently in the one unweeded plantation than the adjacent forest site. Ca nopy Regeneration An in depth canopy regeneration study is beyond the scope of this research. While I was not able to collect detailed data on all of the canopy species present, I did record certain frequent overstory species. The majority of these species were also found in the understory as seedlings and consequently it appears that the common canopy species are regenerating in the xate enrichment sites ( Tables 3 2, 3 3, and 3 4). The only concerning factor would be that Cedrela odorata was found in the o verstory of unweeded plantations but not in the understory. However it was found very infrequently
54 in the overstory and, as mentioned earlier, it might also be present in the understory but not recorded in xate plantations due to its rarity status. Canopy O penness The canopy became increasingly open with management intensity weeded xate plantations were the most open, followed by unweeded xate planatations and forest sites. However, this difference was not significant between management types (p = 0.127 ). The percent canopy cover recorded in each management type is presented in Figure 3 10
55 Figure 3 1. Understory plant species richness in xate planta tions and forest. Observed richness is the total number of species recorded in the field for each site. Estimated richness is the number of species calculated with the species richness estimator Chao 2. W = weeded xate plantations, U = unw eeded xate plantations, F = secondary growth forest sites
56 Figure 3 2. Estimation of the understory plant species ri chness in xate planatations and forest. Estimated richness is the number of species calculated with the species richness estimator Chao 2, with standard deviation W = weeded xate plantations, U = unweeded xate plantations, F = secondary growth forest site s
57 Figure 3 3. Mean understory plant species richness (with standard error) in forest sites, unweeded plantations, and weeded plantations. Observed richness is the total number of species recorded in the field for each site. Estimated richness is the number of species calculated with the species richness estimator Chao 2. The standard error does not account for the individual variances of the Chao 2 estimator presented in Figure 3 2. Forest = secondary growth forest sites U nweeded = unweeded xate pla ntations, Weeded = weeded xate plantations.
58 Figure 3 (with standard error) in forest sites, unweeded xate plantations and weeded xate plantations; before and after cultivated xate has been removed from the analysis. Fo rest = secondary growth forest sites U nweeded = unweeded xate plantations, Weeded = weeded xate plantations.
59 Figure 3 5. Relative abundance of dominant understory plant species for unweeded xate plantations and the corresponding relative abundance fo r the same species recorded at weeded xate plantations and forest sites. Forest = secondary growth forest sites U nweeded = unweeded xate plantations, Weeded = weeded xate plantations.
60 Figure 3 6. Relative abundance of dominant understory plant species for weeded xate plantations and the corresponding relative abundance for the same species recorded at unweeded xate plantations and forest sites. Forest = secondary growth forest sites U nweeded = unweeded xate plantations, Weeded = weeded xate plantations
61 Figure 3 7. Relative abundance of dominant understory plant species for forest sites and the corresponding relative abundance for the same species recorded at unweeded xate plantations and weeded xate plantations. Forest = secondary growth fores t sites U nweeded = unweeded xate plantations, Weeded = weeded xate plantations.
62 Figure 3 8. Relative abundance of invasive species, Oeceoclades maculata for secondary growth forest sites, unweeded xate plantations, and weeded xate plantations.
63 Tab le 3 1. Rare species found in research study and corresponding rarity status. Rare Species Rarity Status Source Calophyllum brasiliense Threatened in Guatemala* CONAP Cedrela odorata Vulnerable** IUCN Red List Gaussia maya Vulnerable** IUCN Red List Lo uteridium chartaceum Endemic Belize Checklist Zamia cf. polymorpha Near Threatened*** IUCN Red List *CONAP (2001) places this tree in category 3 on the list of plants threatened with extinction, a species that is threatened and possibly endangered with extinction in the near future. **IUCN (2004) defines Vulnerable as a species facing a high risk of extinction in the wild in the medium term future ***IUCN (2004) defines Near threatened as a species that is close to qualifying for a threatened category in the near future
64 Figure 3 9. Relative abundance of rare species for secondary growth forest sites, unweeded xate plantations, and weeded xate plantations.
65 Table 3 2. Cano py regeneration in forest sites Canopy species Frequency observed in overstory Relative abundance in understory Acacia sp. #1 I 1.795 Allophylus cominia I 0.066 Attalea cohune C 1.729 Bursera simaruba C 1.363 Byrsonima sp. #1 I -Cecropia peltata I -Chrysophyllum mexicanum I 1.164 Coccoloba cf. tuerckheimii I 0.499 Cord ia alliodora I 0.133 Cryosophila stauracantha C 0.698 Cupania belizensis C 5.452 Eugenia sp. #2 C 2.626 Salicaceae #3 I 0.033 Gaussia maya I 0.399 Hampea trilobata I 0.133 Lauraceae #8 I 0.033 Pouteria sp. #1 I 0.166 Protium copal I 0.499 Sabal m auritiiformis C 0.233 Spondias mombin C 0.598 Trophis mexicana I 0.199 Trophis racemosa I 0.964 C = Common, I = Infrequent
66 Table 3 3 Canopy regeneration in unweeded xate plantation sites Canopy species Frequency observed in overstory Relative abun dance in understory Acrocomia aculeata I 0.031 Allophylus cominia I 0.061 Allophylus sp. #1 I 0.031 Astronium graveolens I 0.214 Attalea cohune C 0.398 Bursera simaruba C 0.980 Caesalpiniaceae #1 I 0.031 Cecropia peltata I -Cedrela odorata I -Chrysophyllum mexicanum C 5.668 Coccoloba cf. tuerckheimii C 0.643 Cordia alliodora I 0.153 Cryosophila stauracantha C 1.164 Cupania belizensis C 2.819 Eugenia sp. #2 C 1.746 Eugenia sp. #3 I 0.092 Hampea trilobata I 0.092 Lauraceae #8 I 0.031 Metopium brownei I 0.858 Pimenta dioica I 0.276 Protium copal I 0.306 Sabal mauritiiformis C 0.490 Simarouba glauca I -Spondias mombin C 0.460 C = Common, I = Infrequent
67 Table 3 4 Canopy regeneration in weeded xate plantation sites Canopy speci es Frequency observed in overstory Relative abundance in understory Acacia sp. #1 I 3.016 Annona reticulata I 0.195 Attalea cohune C 0.531 Bursera simaruba C 1.145 Cecropia peltata I -Chrysophyllum mexicanum C 2.206 Coccoloba cf. tuerckheimii C 0.195 Cojoba graciliflora I 0.028 Cordia alliodora I 0.223 Cryosophila stauracantha C 0.475 Cupania belizensis C 3.742 Eugenia sp. #2 C 0.475 Gaussia maya I 0.056 Metopium brownei C 4.328 Nectandra sp. #1 I 0.363 Sabal mauritiiformis C 0.363 Spo ndias mombin C 0.531 Trophis mexicana I 0.028 Trophis racemosa I 0.782 C = Common, I = Infrequent
68 Figure 3 10. Mean percent canopy cover for secondary growth forest sites, unweeded xate plantations, and xate weeded plantations, with standard error
69 CHAPTER 4 DISCUSSION Conclusions The objective of this study is to develop a broader understanding of the ecological impacts of xate palm enrichment plantings on the understory plant species of secondary growth forests in Belize. While it is beyond the scope of this study to assess the long term ecological effects of Chamaedorea cultivation in the forest understory, the results indicate that there is no significant difference between xate enrichment sites and adjacent secondary growth forest sites in we stern Belize in terms of understory plant species richness, diversity, and composition. Oeceoclades maculata was encountered at about the same relative abundance in weeded xate plantations, unweeded xate plantations, and forest sites. The exotic orchid ha s been found to be most abundant in areas with moderate disturbance levels but also grows in areas that have not been clear cut or managed for agriculture (Cohen and Ackerman 2009). This conclusion leads one to predict that while it would be expected at un managed forest sites, it may be significantly more abundant in areas with higher management intensity. However, I found no significant difference in the relative abundance of Oeceoclades maculata in the forest sites as compared to the xate enrichment sites A direct relationship between canopy openness and understory composition is beyond the scope of the study. However, since I did not find a significant difference in canopy cover between weeded xate plantations unweeded xate plantations, and secondary g rowth forest sites, it is possible that the t hree management types do not significantly differ in terms of abiotic factors such as light availability, temperature, and
70 soil moisture. These factors can influence plant growth and survival ( Augspurger 1984; K obe 1999) and affect forest regeneration and community composition (Clark et al. 1996; Montgo mery and Chazdon 2002) I did not find a significant difference in understory plant richness or diversity between forest sites and xate plantations, which indicat es that the capability for recovery of this community was high. This is especially true for sites located within weeded plantations as the understory layer is cut about every six months. This may be because this area of Belize is adapted to natural disturb ance of frequent fires and hurricanes (Primack et al. 1997) Based upon the results presented in the previous chapter, this research provides evidence that the management utilized at these xate plantations has minimal negative effects at the ecological und erstory community level This suggests that the intensity of management practices employed by the Belizean xate plantation owners in this study was not significant enough to lead to significant differences in the understory species community. Further studi es should be conducted to substantiate these findings; however this study supports the conclusion that xate enrichment plantings that retain a natural forest canopy in western Belize are sustainable at the understory plant community level. The process of cutting the vegetation under the forest canopy prior to establishment and periodic weeding with a machete could be predicted to result in very visible differences in the understory community. In reality, the border between the surrounding secondary growth forest and the xate plantation is actually difficult to distinguish ( Figures 4 1 and 4 2). The xate plantations in western Belize as a result,
71 resemble enrichment plantings. They are very different from typical monoculture plantations (Figure 1 8). In int erviewing plantation owners, I found that some farmers needed an economic use for the secondary growth forest on their land. If the option had not been available to the farmers to convert the land into xate plantations, they stated that they might have con verted the secondary growth forest into a milpa, the local name for small farm. In this way, the xate plantation concept may deter deforestation among small land owners in Belize. Research Implications Implications for Sustainable Forest M anagement The tran sition from harvesting NTFPs in the wild to growing NTFPs as crops in plantations is a growing trend ( Bridgewater et al 2006 ; Smith et al 1992; Ticktin 2004 ). For xate in Belize, this conversion became a reality due to the pressure put on wild populations from illegal harvesting. A sustainable harvest of xate in the wild is not possible as long as illegal harvest continues (Bridgewater et al. 2007) While plantations generally have lesser population level effects as compared to wild harvest, care must be ta ken to minimize negative community level effects ( Ticktin 2004) Considering that biodiversity is essential for not only ecosystem health but also for human health and economic well being, the conservation of biodiversity should not only be implemented in protected areas. Instead, forest biodiversity should be maintained in areas throughout the forested landscape (Lindenmayer and Franklin 2002; Tarrega et al. 2006). In general, anthropogenic changes, such as certain management practices, can be so great th at they may result in the disappearance of a large number of establishe d species and their replacement by immigrant species. Some species may tolerate the
72 disturbance while others may vanish (Sheil 1999; Sagar et al. 2003). The species richness present at an area is therefore clearly related to the intensity and type of the anthropogenic change. In sustainable management, the understanding of these patterns of change along the gradient of management intensity is crucial in determining the management intensi ty threshold level and developing strategies to conserve biodiversity while providing goods to humans. The levels of management intensity within different management schemes must be understood and analyzed in order to inform management decisions. The under standing of management intensity levels and their affects on ecological developing management techniques that are truly sustainable. I will examine a few of these manage ment practices implemented in NTFP plantations and relate them to my research. The ideas presented in this discussion are intended to be interpreted with caution as only a few studies exist on Chamaedorea plantations, understory plant species community com position in NTFP plantations, or the influence of specific management techniques on ecological communities. These findings do point to possible reasons for differences in the ecological community found by various studies. A multitude of studies have found that plantations with a shade canopy contain a significantly higher diversity for numerous taxa including understory plants than Estrada et al. 1994; Greenberg et al. 1997 ; Lest on 1970; Room 1971 ). Many other studies suggest plantations, maintain significantly higher biodiversity and richness than plantations with
73 a shade canopy of just on Greenberg et al. 1997 ; Mas and Dietsch 2 004 ). These results lead one to conclude that a significant factor in diversity richness an d composition of plantations is the amount and type of overstory shade trees. Yet, while some studies indicate that forests contain a significantly higher diversity and richness than rustic plantations ( Alves 1990; Epps 2009 ; Rolim and Chiarello 200 4; Trau ernicht and Ticktin 2005 ), others indicate that there is no significant difference between the two types (Mas and Dietsch 2004 ; Perfecto et al 1996 ). The discrepancy between these conclusions may relate to a combination of factors but they most likely re late to variations in management practices, including but not limited to the number and type of shade trees. This issue has not been properly addressed in scientifically published articles even though it is a key factor that relates to the inconsistencies in the study of NTFP cultivation (Greenberg et al. 1997). While some rustic plantations are very conspicuous with patchy canopy cover and extremely open understories ( Figure 4 3), others are difficult to distinguish f rom the adjacent forest (Figures 4 1 an d 4 2 ). All of the rustic xate plantations sampled in this study have a canopy cover and understory similar to secondary growth forests and are difficult to distinguish from the adjacent vegetation. The variation of conditions present in plantations is dir ectly related to the range of plantation management techniques that the plantation owner considers both in the beginning phases of plantation establishment and afterwards during regular maintenance. These decision making differences depend on the ecologica l knowledge, environmental ethic, and forethought of the plantation owner.
74 One factor that must be taken into consideration is the level of shade the cultivated plant requires. Chamaedorea ernesti augustii is one of the most shade tolerant species under c ultivation. Some species that can be grown in rustic conditions such as coffee and cacao do not require shade and can be grown in sun plantations ( Perfecto et al 1996 ; Rice and Greenberg 2000 ). Other species such as Chamaedorea hooperiana do require shade (Trauernicht and Ticktin 2005) but need more sun than C. ernesti augustii (Mesh 2007 pers. comm. ). Many of the coffee, cacao, and Chamaedorea plantation studies I have encountered in scientific articles that discuss plantation management state that the o verstory, regardless of type of shade tree, is routinel y pruned (Greenberg et al. 1997; Mas and Dietsch 2004). Trimming is p erformed to increase the amount of light and therefore increase the production of the cultivated plant. In C. hooperiana plantations large leaves are pruned that may cover and kill seedlings when shed (Trauernicht and Ticktin 2006). This seemingly slight difference in periodic pruning may explain differences in species diversity, richness, and composition between studies conducted at different plantations Studies on rustic cacao plantations in Brazil with pruned overstories found significantly higher plant species richness and diversity in unmanaged forests as compared to the rustic plantations. The plant species composition was sign ificantly altered, with pioneer and early secondary species becoming more dominate in the rustic plantations ( Epps 2009 ; Rolim and Chiarello 2004 ). study on rustic Chamaedorea hooperiana plantations with routinely pruned ov erstories in southern Mexico found that the stem density, species richness, and species diversity of understory plants was significantly lower in plantation sites as compared to the
75 adjacent forest. The canopy in C. hooperiana plantations was significantly more open as compared to the adjacent forest, most likely a result of the periodic pruning. The greater openness of the canopy may have led to higher light levels, lower soil moisture, and higher temperatures in the understory as well as different levels of soil nutrients, all which may have influenced the change in the community. I found through interviews that xate plantation owners in western Belize do not prune the overstory. No significant difference in canopy cover was recorded between plantations an d adjacent forest sites. I did not find significant differences in species richness or diversity between forest sites and plantations in this study, which may be related to a maintenance in percent canopy cover in xate plantations. Greenberg et al. (1997) noted that bird species richness seemed to decrease in the following order: forests, rustic coffee plantations that had not been trimmed, and rustic coffee plantations that had been heavily trimmed. Mas and Dietsch (2004) found no significant diversity an d richness levels of birds and butterflies in forested areas as compared to rustic coffee plantations. The rustic coffee plantation canopy cover in this study is not pruned and it contains a significantly similar percent canopy cover as the forest. Therefo re, this study provides evidence that a lack of pruning may result in a similar level of bird and butterfly diversity and richness between forest sites and rustic coffee plantations. Significant pruning of overstory trees may affect the degree of a reliabl e source of food for frugivorous birds and butterflies. Pruning of limbs also affects epiphyte and hemiparasite abundance, which certain species such as euphonias depend on for fruit (Greenberg et al. 1997). Additional studies should be conducted that inve stigate possible community changes of multiple taxa in relation to pruning.
76 Trauernicht and Ticktin (2005) also found that one of the forest dominant understory palms, Astrocaryum mexicanum was significantly less abundant in rustic plantations as compared to forest sites. Astrocaryum mexicanum was systematically removed by plantation owners due to its trunk with a multitude of spines and large fronds that can kill planted seedlings. In interviewing xate plantation owners in western Belize, I found that the y do not select any one particular species to remove over any others. This includes the give and take palm ( Cryosophila stauracantha ), which has a trunk with long sharp spines and large leaves. The relative abundance of C. stauracantha was comparatively si milar in xate plantations as compared to forested sites in this study ( Tables 3 2, 3 3, and 3 4) Another factor plantation owners consider is the location of the plantation. Some studies that found significant differences between rustic plantations and fo rests in terms of diversity were conducted in rustic plantations that were established under an old growth canopy (Trauernicht and Ticktin 2005). Other studies, such as this one, that found no significant differences between rustic plantations and forests in terms of diversity were conducted in rustic plantations that were under secondary growth canopy ( Rice and Greenberg 2000). Plantations established under a secondary growth forest canopy theoretically will most likely contain a diversity, richness, and c omposition more similar to secondary growth forests than plantations established under old growth forests when compared to old growth forests themselves. Secondary growth forests are more acclimated to disturbance than primary forests and therefore the spe cies that inhabit secondary growth forests are more likely to be able to adapt to management techniques such as periodic weeding. To my knowledge, not a single study has been
77 conducted comparing rustic plantations that have been established under an old gr owth canopy with rustic plantations that have been established under a secondary growth canopy. Future research is needed to address this question. The richness, diversity, and composition of an ecological community are also significantly related to the su rrounding landscape. Research suggests that rustic agroforestry systems located adjacent to natural forest areas tend to have high richness and diversity of forest understory plants, mammals, birds, and invertebrates as compared to those rustic agroforestr y plots isolated from forest patches ( Alves 1990; Estrad a et al. 1994, 1997; Faria et al 2007; Pardini 2004 ; Young 2007 ). This research supports this idea since the xate enrichment sites sampled in this study were located within a forested landscape. Rusti c plantations may provide corridors for animals foraging and migrating between forest tracts (Rice and Greenberg 2000). Xate plantations with plant species richness and diversity levels similar to forest sites and also located adjacent to forest patches ma y also support the same level of diversity in terms of birds, mammals, insects, amphibians and reptiles as secondary growth forest areas. This study, supported by data from other studies, provides evidence that xate plantation owners who apply the followi ng practices will most likely not significantly affect the understory plant community in western Belize: (1 ) establish plantations near natural forests; ( 2 ) retain the natural forest canopy to provide shade for the xate, maintain the regeneration of plant species, and maintain similar forest abiotic factors (i.e. soil moisture, light availability, and nutrients from leaf litter) that may affect understory plant species diversity and richness; ( 3 ) establish plantations in the
78 understory of secondary growth f orests as opposed to old growth forests; ( 4 ) very infrequently weed or do not weed the understory in order to not only lessen the cutting of individuals but also to minimize compaction; ( 5 ) weed only the plants that are located near the Chamaedorea which m ight be competing with the Chamaedorea ; ( 6 ) weed only using a machete to cut the plant at the forest floor, taking care to not remove or root out the plant; ( 7 ) do not selectively remove specific species when weeding; ( 8 ) do not prune the overstory; and ( 9 ) utilize organic methods such as natural fertilizers and pesticides. employed in xate plantations in order to conserve biodiversity based on the data in this study and a review of the literature. This discussion is limited for a number of reasons. Only one study on plant community level effects in addition to this study has been conducted within Chamaedorea plantations Few studies have been conducted on understory plant species communities in NTFP plantations or NTFP enrichment sites The majority of the research on this topic is conducted over the short term and therefore strong predictions on future community composition patterns are not possible. Most management practices oth er than the composition of the tree overstory are not mentioned in the majority of the research on NTFP cultivation. Due to the wide range of management practices implemented by plantation owners, researchers should interview farmers and report these pract ices in scientific articles. I emphasize that further studies should be conducted in xate plantations with different management schemes in different locations for a number of t axa in order to form conclusive
79 and possible certifi cation criteria. Data on other plantations with greater management intensities will need to be collected to establish realistic management level thresholds. The eco certification of xate has been the subject of numerous studies and reports (Bridgewater et al. 2007; Pickles 2004; Wilse y 2008; Wilsey and Endress 2007 ). All express a need for research on the ecological community level effects of xate wild harvest and cultivation in order to establish certification criteria. As it is highly improbable that wild due to illegal activity (Bridgewater et al. 2007), eco certification in Belize may need to focus on cultivated xate harvest. This study could aid in establishing criteria for certification in terms of xate cultivation. This research can only provide evidence towards the sustainability of xate plantations in western Belize that are grown under natural forest canopies and employ the above management practices. A number of xate plantations in Mexico, Guatemala, and Belize employ more intensive management practices such as pruning, intensive weeding, or shaded monoculture/polyculture overstories ( Bridgewater et al. 2007 ; Wilsey and Endress 2007) Further studies should be conducted to determine the ecological effects of xate cultivation that are under more intensive management level practices in order to establish certification criteria. Implications for Wildlife Conservation Many studies have shown that rustic plantations with high plant species richness and diversity also support a high diversity of other bio ta (Perfecto et al. 1996; Rice and Greenberg 2000; Carlo et al 2004). Other research suggests that rustic agroforestry systems located adjacent to natural forest areas tend to have high rich ness and diversity of forest mammals, birds, and invertebrates as compared to those agroforestry plots isolated from forest patches (Estrada et al. 1994, 1997; Faria et al 2007 ; Young
80 2007 ). Some threatened and endangered species endemic to the Atlantic fo rest of Brazil such as the golden headed lion tamarin ( Leontopithecus chrysomelis ) and the pink legged graveteiro ( Acrobatornis fonsecai ) actually prefer rustic cacao plantations to forested areas (Pacheco et al. 1996 ; Raboy et al 2004 ). Rustic plantation s may provide corridors for animals foraging and migrating between forest tracts (Rice and Greenberg 2000). It follows that xate plantations with plant species richness and diversity levels similar to forest sites and located adjacent to forest patches may also support the same level of diversity in terms of birds, mammals, insects, amphibians and reptiles as secondary growth forest areas. While conducting research in the Chamaedorea enrichment sites I observed many birds such as keel billed toucan ( Ramph astos sulfuratus ), pale billed woodpecker ( Campephilus guatemalensis ), black headed trogon ( Trogon melanocephalus ), collard trogon ( Trogon massena ), rufous tailed hummingbirds ( Amazilia tzacatl ), euphonias ( Euphonia spp.), flycatchers, and tanagers. A humm ingbird nest was found in a shrub located in the understory of one xate plantation site. Many birds depend on the fruits and flowers of understory plant species and these include species that I recorded at xate plantations. Ochre bellied flycatchers ( Mion ectes oleaginous ) eat the fruits of Psychotria sp p and vines in the family Dilleniaceae, rufous tailed hummingbirds feed on the nectar of Costus sp p and Passiflora sp p ., black headed trogons consume the fruits of vines in the family Sapindaceae, buff thr oated saltators ( Saltator maximus ) and yellow tailed orioles ( Icterus mesomelas ) eat the fruits of Inga sp p (Piaskowski et al. 2006), and a variety of hummingbirds feed on the nectar of Odontonema tubaeforme (Daniel 2010 pers comm. ). Numerous bird speci es also
81 depend on fruit trees, including species of trees that I recorded in the overstory and in the understory as seedlings at xate plantations. For example, many birds including keel billed toucans and crested guans ( Penelope purpurascens ) feed on the f ruits of Sabal mauritiformis olive throated parakeets ( Aratinga nana ) consume the seeds of Acacia sp p ., black faced grosbeaks ( Caryothraustes poliogaster ) and red lored parrots ( Amazona autumnalis ) feed on the fruits of Protium copal euphonias eat the fr uits of Cupania belizensis and many birds including the black headed trogon, bright rumped attila ( Attila spadiceus ), keel billed toucan, masked tityra ( Tityra semifasciata ), rose throated becard ( Pachyramphus aglaiae ), and slaty tailed trogon ( Trogon mas sena ) depend on Bursera simaruba (Piaskowski et al. 2006). I also observed many butterflies in Chamaedorea plantations such as the zebra longwing ( Heliconius charitonia ), postman ( Heliconius erato ), five spotted longwing ( Heliconius hecalesia ), malachite ( Siproeta stelenes ), cook ( Anartia fatima ), and owl ( Caligo eurilochus sulanus ). I found host plants necessary for these butterflies in the xate enrichment plantings including Justica breviflora (host plant for Anartia fatima and Siproeta stelenes ), Passifl ora biflora (host plant for Heliconius hecalesia and Heliconius erato ) and Heliconia sp p (host plant for Caligo eurilochus sulanus ). While the majority of these species are quite common in rainforest, forest edges, and plantations, Heliconius hecalesia an d Caligo eurilochus sulanus typically prefer unmanaged rainforest to disturbed areas (DeVries 1987). Limitations It is not feasible from this study to determine other possible community level effects such as changes in forest structure. It was beyond the s cope of this study to measure the height and diameter at breast height (dbh) of every individual in the shrub
82 and understory layer. Nor was it possible to record species within the mid story (i.e. greater than 1.30 m ) due to time and funding constraints. However I observed that xate plantations might lack medium sized woody plants as compared to forest sites, especially between forest sites and weeded xate plantations. Since the weeded plantations are only cut with a machete every six months and the recove ry of the community is high, the herbaceous and woody plants are able to regrow in the understory, not affecting richness and diversity. I did observe several herbaceous plants in flower and fruit in both plantation types; however this was rarer for woody species. As a result, the weeding may prevent plants from reaching the maturity needed for the production of reproductive structures and consequently the ability for some woody species in the shrub layer to reproduce and provide nectar or fruits to animals Although this study indicates that the common canopy species in xate plantations are most likely regenerating as understory tree seedlings, they may be cut before growing into canopy sized trees. Additional research is needed to understand possible diffe rences in structure between the xate plantations and forest sites. Conclusion Xate plantations, as well as other NTFP plantations, cannot provide the same functions, processes, community composition, and diversity for all taxa as old growth forests. As a r esult, old growth forests must be protected in reserves (Faria et al 2007). Conversely, forests throughout the world are being utilized by humans and a strategy to conserve biodiversity strictly through ecological protected areas will most likely fail ( And erson 1990 ; Lindenmayer and Franklin 2002 ). Secondary growth forests and agroforests are important dispersal and gene flow pathways and aid in landscape scale conservation ( Brown and Lugo 1990; Chazdon et al. 2009; Faria et al 2007; Ribeiro et
83 al. 2010; Ri ce and Greenberg 2000). The paradigm between protecting forests in reserves and protecting forests through management should not be proposed as a black or white scenario. Instead, these rustic xate plantations and other agroforests, ought to be regarded as complementary ecosystems in wider conservation scheme in which the foundation is the native forest (Schroth et al. 2004). The conservation of biodiversity ultimately depends on the conservation of a multitude of habitats within the landscape.
84 Figure 4 1. Weeded xate p lantation at Location A (Figure 2 1), established west of San Antonio, Cayo, Belize Carly Voight
85 Figure 4 2. Unweeded xate plantation at Location A (Figure 2 1), established west of San Antonio, Cayo, Belize Carly Voight
86 Figure 4 3. Rustic xate ( Chamaedorea elegans ) plantation in Catemaco, Veracruz (Mexico). This plantation was not sampled in this study. David Wilsey
87 APPENDIX A LIST OF SPECIES Family Species Common name Status Acanthaceae Acanthaceae #1 Acanthaceae #2 Acanthaceae #3 Acanthaceae #4 Acanthaceae #5 Acanthaceae #6 Acanthaceae #7 Aphelandra deppeana Justica breviflora au'kop zorillo macho Louteridium chartaceum Endemic Odontonema tubaeforme Adiantaceae Adiantium sp. #1 Adiantum tenerum maiden hair fern Amaranthaceae Amaranthaceae #1 Anacardiaceae Astronium graveolens jobillo Metopium brownei black poisonwood Spondias mombin hog plum Annonaceae Annona reticulata wild custard apple Annonaceae #1 Annon aceae #2 Apocynaceae Apocynaceae #1 Apocynaceae #2 Apocynaceae #3 Apocynaceae #4 Apocynaceae #5 Apocynaceae #6 Apocynaceae #7 Mandevilla sp. #1 Tabernaemontana alba grandpa's balls Tabernaemontana arborea horse's balls Tabernaemontana sp. #1 Thevetia ahouai dog's balls Araceae Anthurium schlechtendalii phe a sant's tail Araceae #1 Araceae #2 Monstera sp. #1 Syngonium cf. podophyllum Syngonium podophyllum Syngonium sp. #1 Syngonium sp. #2 Syngonium sp. #3 Arecaceae Acrocomia aculeata grugru palm Arecaceae #1 Attalea cohune cohune palm Chamaedorea ernesti augustii xate fishtail Chamaedorea oblongata xate jade
88 APPENDIX A, cont. Family Species Common name St atus Arecaceae Chamaedorea tepejilote tepe jilote, pacaya Cryosophila stauracantha give and take Desmoncus orthacanthos basket tie tie Gaussia maya palmasito Rare Sabal mauritiiformis sabal palm Aristolochiaceae Aristolochia maxima guaco As pleniaceae Asplenium sp. #1 Asteraceae Asteraceae #1 Asteraceae #2 Asteraceae #3 Asteraceae #4 Asteraceae #5 Bidens squarrosa Chromolaena odorata Critonia morifolia palo verde Lepidaploa sp. #1 Mikania sp. #1 Mikania sp. #2 Mikania sp. #3 Neurolaena lobata jackass bitters Bignoniaceae Bignoniaceae #1 Bignoniaceae #2 Bignoniaceae #3 Bignoniaceae #4 Bignoniaceae #5 Bignoniaceae #6 Bignoniaceae #7 Bignoniaceae #8 B ignoniaceae #9 Bignoniaceae #10 Bignoniaceae #11 Bignoniaceae #12 Stizophyllum riparium so bach cf. Mansoa sp. #1 cf. Mansoa sp. #2 Boraginaceae Boraginaceae #1 Boraginaceae #2 Boraginaceae #3 Boraginaceae #4 Cordi a alliodora salmwood Cordia sp. #1 Cordia sp. #2 Burseraceae Bursera simaruba red gumbolimbo Protium copal copal Protium glabrum Burseraceae #1
89 APPENDIX A, cont. Family Species Common name Status Calophyllaceae Calophyll um brasiliense Rare Cannabaceae Celtis iguanaea Celastraceae Crossopetalum gaumeri Chrysobalanaceae Chrysobalanaceae #1 Hirtella sp. #1 Combretaceae Combretaceae #1 Combretum sp. #1 monkey brush Terminalia amazonia nargosta Commelin aceae Tripogandra grandiflora Connaraceae Rourea glabra Convolvulaceae Ipomoea heterodoxa Ipomoea sp. #1 Ipomoea sp. #2 Ipomoea trifida Costaceae Costus sp. #1 Cucurbitaceae Cayaponia racemosa Cucurbita cf. lundelliana Rytid ostylis gracilis Cyperaceae Scleria sp. #1 Dilleniaceae Dilleniaceae #1 Dilleniaceae #2 Tetracera sp. #1 Dioscoreaceae Dioscorea sp. #1 Dioscorea sp. #2 Dioscorea sp. #3 Dioscorea sp. #4 Erythroxylaceae Erythroxylum bequaert ii Euphorbiaceae Acalypha sp. #1 Alchornea latifolia Cnidoscolus sp. #1 Croton sp. #1 Dalechampia heteromorpha Dalechampia sp. #1 Dalechampia sp. #2 Euphorbiaceae #1 Euphorbiaceae #2 Euphorbiaceae #3 Euphorbiaceae #4 Euphorbiaceae #5 Euphorbiaceae #6 Euphorbiaceae #7 Fabaceae Fabaceae #1 Fabaceae #2 Fabaceae #3 Fabaceae #4 Fabaceae #5
90 APPENDIX A, cont. Family Species Common name Status Fabaceae Fabaceae #6 Fabaceae #7 Fabaceae #8 Fabaceae Caesalpinioideae Bauhinia ungulata cow foot tree Caesalpiniaceae #1 Caesalpiniaceae #2 Fabaceae Mimosoideae Acacia sp #2 white cockspur Acacia sp. #1 red cockspur Cojoba arborea barba jelote Cojoba gra ciliflora john cow bead Inga sp. #2 Inga pinetorum Mimosa sp. #1 Mimosa watsonii var. recordii haulback Mimosoidceae #1 Pithecellobium hymenaeifolium clavos k'iix Fabaceae Papilionoideae Calopogonium sp. #1 Calopogonium mucun oides Centrosema virginianum Clitoria cf. ternatea false man vine Desmodium cf. metallicum Desmodium incanum strong back Lonchocarpus minimiflorus white cabbage bark Piscidia piscipula jabin Rhynchosia sp. #1 Rhynchosia sp. #2 Swartzia myrifolia var. standleyi bastard rosewood Gesneriaceae Gesneriaceae #1 Heliconiaceae Heliconia sp. #1 Heliconia sp. #2 Heliconia sp. #3 Lamiaceae Ocimum campechianum wild basil Lauraceae Lauraceae #1 Lauraceae #2 Lau raceae #3 Lauraceae #4 has che Lauraceae #5 Lauraceae #6 Lauraceae #7 Lauraceae #8 black timbersweet Lauraceae #9 Lauraceae #10 Nectandra sp. #1 white timbersweet Loganiaceae Spigelia humboldtiana Strychnos panamensis Malpighiaceae Hiraea fagifolia
91 APPENDIX A, cont. Family Species Common name Status Malpighiaceae Malpighiaceae # 1 Malpighiaceae #2 Malpighiaceae #2 Malpighiaceae # 3 Malpighiaceae #4 Malpighiaceae #5 Malpighiaceae # 6 sk unk root Malpighiaceae #7 Malvaceae Hampea trilobata moho Malvaceae # 1 moho Malvaceae # 2 Malvaviscus arboreus white moho Corchorus siliquosus Luehea speciosa bay cedar Melastomataceae Clidemia octona Mouriri myrtilloides Mel iaceae Cedrela odorata red cedar Rare Trichilia sp. #2 Trichilia sp. #1 Menispermaceae Cissampelos tropaeolifolia Menispermaceae #1 Menispermaceae #2 Moraceae Brosimum terrabanum Moraceae #1 Trophis mexicana Trophis racemosa Myrtaceae Eugenia capuli chi lun che Eugenia sp. #1 pigeon berry Eugenia sp. #2 pigeon berry Eugenia sp. #3 pigeon berry Eugenia sp. #4 pigeon berry Eugenia sp. #5 pigeon berry Myrtaceae #1 Myrtaceae #2 Myrtaceae #3 Myrtac eae #4 Myrtaceae #5 Myrtaceae #6 Myrtaceae #7 Myrtaceae #8 Myrtaceae #9 Pimenta dioica allspice Nyctaginaceae Nyctaginaceae #1 Nyctaginaceae #2 Nyctaginaceae #3 Pisonia sp. #1
92 APPENDIX A, cont. Family S pecies Common name Status Ochnaceae Ouratea sp. #1 Orchidaceae Oeceoclades maculata Invasive Passifloraceae Passiflora biflora Passiflora coriacea Passiflora mayarum Passiflora rovirosae Passiflora serratifolia Passiflora xiikzodz ssp. xiikzodz batwing passionflower Phyllanthaceae Phyllanthus sp. #1 Picramniaceae Alvaradoa amorphoides Piperaceae Piper jacquemontianum Piper marginatum Piper sp. #1 Piper sp. #2 Poaceae Oplismenus hirtellus Panicum sp. #1 Poaceae #1 Poaceae #2 star grass Poaceae #3 Poaceae #4 Poaceae #5 Poaceae #6 Poaceae #7 Poaceae #8 Poaceae #9 Polygonaceae Coccoloba acapulcensis Coccoloba cf. tuerckheimii Coccoloba sp. #1 Polygalaceae Polyg ala hondurana Securidaca diversifolia man vine Polypodiaceae Pleopeltis sp. #1 Polypodiaceae #1 Primulaceae Ardisia sp. #1 Ardisia sp. #2 Ranunculaceae Clematis sp. #1 Rhamnaceae Gouania polygama Rubiaceae Chiococca alba black sku nk root Geophila repens Hamelia patens ixcanan polly redhead Hamelia sp. #1 Morinda sp. #2 Morinda sp. #1 Psychotria fruticetorum Psychotria sp. #1 red anal Psychotria sp. #2 green anal Psychotria sp. #3
93 APPENDIX A, cont. Family Species Common name Status Rubiaceae Psychotria tenuifolia dark green anal Rubiaceae #1 Rubiaceae #2 Rubiaceae #3 Rubiaceae #4 Rubiaceae #5 Rubiaceae #6 Rubiaceae #7 Rubiaceae #8 Rubiaceae #9 Rub iaceae #10 Rubiaceae #11 Rubiaceae #12 Rubiaceae #13 Rubiaceae #14 Rubiaceae #15 Rubiaceae #16 Rubiaceae #17 Rubiaceae #18 Rubiaceae #19 Rutaceae Zanthoxylum caribaeum Salicaceae Casaeria sp. #1 ixim che Casaeri a sp. #2 Prockia crucis Salicaceae #1 Salicaceae #2 Salicaceae #3 Xylosma characantha Sapindaceae Allophylus cominia Allophylus sp. #1 Cupania belizensis grande betty Paullinia cf. costaricensis Paullinia costata Paullinia cf. pinnata Paullinia pinnata Paullinia sp. #1 Paullinia sp. #1 Paullinia sp. #2 Paullinia sp. #3 Sapindaceae #1 Sapindaceae #2 Sapindaceae #3 Sapindaceae #4 Sapindaceae #5 Sapindaceae #6 Sapindace ae #7
94 APPENDIX A, cont. Family Species Common name Status Sapindaceae Sapindaceae #8 Sapindaceae #9 Sapindaceae #10 Sapindus saponaria soap seed tree Serjania macrocarpa Serjania sp. #1 Talisia oliviformis white kanep Sapotaceae Chrysophyllum mexicanum wild star apple Pouteria sp. #1 wild canastel Sapotaceae #1 Sideroxylon cf. obtusifolium Sideroxylon sp. #1 Schizaeaceae Lygodium venustum wire whiss, alambre Scrophulariaceae Russelia sarmentosa Sim aroubaceae Simarouba glauca negrito Smilacaceae Smilax cf. spinosa Smilax mollis Smilax spinosa Smilax aristolochiaefolia Solanaceae Solanum diphyllum veneno xiu Thelypteridaceae Thelypteris sp. #1 Tiliaceae Heliotropium sp. #1 Urt icaceae Pilea sp. #1 Pourouma bicolor Urticaceae #1 Verbenaceae Petrea volubilis panuelo de sarah Rehdera penninervia hinge hinge Stachytarpheta cayennensis vervain Violaceae Rinorea guatemalensis Vitaceae Cissus microcarpa Vitis tiliifolia Zamiaceae Zamia cf. polymorpha Rare unidentified species indet #1 indet #2 indet #3 indet #4 indet #5 indet #6 indet #7 indet #8 indet #9 indet #10 indet #11 indet #12 indet #13 indet #14 indet #15 indet #16
95 APPENDIX A, cont. Family Species Common name Status unidentified species indet #17 indet #18 indet #19 indet #20 indet #21 indet #22 indet #23 indet #24 indet #25
96 APPENDIX B ESTIMATED RICHNESS R ESULTS Site Name ACE Mean ICE Mean Chao 1 Mean Chao 1 SD Chao 2 Mean Chao 2 SD Jack 1 Mean Jack 1 SD Jack 2 Mean Bootstrap Mean MMRuns Mean F2 96.07 121.85 119.56 24.75 122.17 22.68 104.6 5.88 123.27 87.54 112.37 F4 126 141.28 152.25 36.59 156.05 35.32 115.1 5.1 139.19 94.95 104.96 F5 113.71 134.72 110.53 10.22 117.63 11.77 122.5 5.24 134.14 105.92 126.5 F1 137.35 152.36 142.29 20.91 142.69 19.51 132.9 5.1 152.9 112.5 129.91 F3 122.4 144.84 131.88 19.74 132.25 18.01 125.1 5.6 143.5 105.9 128.07 U2 96.24 113.65 91.62 10.41 107.32 16.45 100.9 2.83 115.74 85.51 98.53 U1 100.69 113.71 109.25 16.89 114.96 17.84 105.9 7.03 121.46 90.55 105.89 U5 131.68 145.26 142.12 21.72 135.0 3 16.99 130.1 7.4 147.79 111.08 137.47 U4 133.51 163.76 125.17 12.92 144.66 19.84 134.7 7.35 155.59 113.33 125.86 U3 135.46 163.94 123.63 11.77 139.02 16.93 135.7 7.11 154.46 114.7 144.45 W1 92.38 102.05 93.04 9.03 104 13.49 101.4 4.49 112.96 88.88 102. 58 W4 97.7 112.93 92.53 9.01 113.5 18.5 103 5.2 118.47 88.13 100.56 W2 124.1 151.04 133.38 21.6 135.06 19.97 124 6.43 143.91 104.03 116.6 W3 113.09 127.08 114.04 14.8 125.88 19.74 113.7 5.7 130.86 97.29 110.89 W5 127.87 152.45 126.89 18.17 132.44 18.75 124 6.15 143.2 104.11 117.2 F = Secondary growth f orest site, U = unweeded xate plantation, W = weeded xate plantation, SD = standard deviation
97 APPENDIX C LIST OF INFORMANTS Oral or written information is cited as a personal communication (pers. comm. ) in the text. Bol N 200 7. Operations Manager, Las Cuevas Research Station. Daniel T F 2010. Curator, California Academy of Sciences. Mesh, J. 2008. Technical Assistant, Itzamna Society.
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107 BIOGRAPHICAL SKETCH Carly Voight has had a lifelong interest in tropical rainforest ecology and m anagement. She holds a Bachelor of Science in international ecosystem management from the University of Wisco nsin Stevens Point and a Master of Arts in Latin American s tudies with a tropical conservation and developmen t concentration. Carly has been intrigued with rain forests since the age of six and first received the opportunity to travel to the tropic s at seventeen. Along with six other high school students, Carly was selected by The Nature Conservancy and the Wisconsin Energy Corporation to study tropical ecology with the Belize Bound Program at Rio Bravo Conservation and Management Area in northwest Belize. During this study abroad program, Carly met a timber forest products. While in undergraduate school, she s tudied abroad through various programs to Belize, Mexico, Guatemala, and Costa Rica. She has previously worked for The Nature Conservancy in Florida, St. Croix (U.S. Virgin Islands), upstate New York, Arizona, Colorado, and Michigan. She strongly believes in a community based collaborative approach to conservation planning and adaptive management. In her spare time, she indulges in her dream of becoming a nature photographer by running a small business, Rainforest Reflections ( http://www.rainforestreflections.com ).