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Lacandon Maya forest management: Restoration of soil fertility using native tree species
Stewart A.W. Diemont a , Jay F. Martin a, , Samuel I. Levy-Tacher b , Ronald B. Nigh c , Pedro Ramirez Lopez d , J. Duncan Golicher b
Ecological Engineering Group, Department of Food Agricultural and Biological Engineering, The Ohio State University, 590 Woody Hayes Dr., Columbus, OH 43210-1057, USA b Division of Conservation and Biodiversity, Department of Ecology and Terrestrial Systems, El Colegio De La Frontera Sur, San Cristobal de Las Casas, Chiapas, Mexico c Centro de Investigaciones y Estudios Superiores en Antropologia Social del Sureste, San Cristobal de Las Casas, Chiapas, M exico d Department of Agroecology, El Colegio De La Frontera Sur, San Cristobal de Las, Casas, Chiapas, Mexico
a

a r t i c l e
Article history:

i n f o

a b s t r a c t
In southern Mexico, where rainforests are being degraded rapidly, the Lacandon Maya use an agroforestry system that both restores and conserves the rainforest. Their system cycles through eld and fallow stages that produce food, medicines, and raw materials, and regenerates tall secondary forest. This investigation identied plants managed by Lacandon to restore soil fertility during fallow. Through interviews, Lacandon identied 20 plants managed for forest restoration. Leaf litter measurements and soil samples were taken near two of these species, Ochroma pyramidale and Sapium lateriorum. Leaf litter increased quicker

Received 23 June 2005 Received in revised form 4 October 2005 Accepted 28 October 2005

Keywords: Rainforest restoration Indigenous knowledge Soil ecology Ochroma pyramidale Sapium lateriorum

beneath O. pyramidales compared to other tree species (R = 0.48, P = 0.004), and total nematode concentrations increased with distance from this tree (R = 0.71, P < 0.001). Together, these two ndings indicated an inhibition of degradation that permits accelerated soil organic matter accumulation. Available phosphorus (P) concentrations beneath S. lateriorum were 16% higher than outside the canopy (P = 0.03), and increased with age of the tree, indicating P recovery from subsoil. Our research shows that the Lacandon are cognizant of the natural abilities of certain species to fulll the restoration needs in their systems. It demonstrates that Maya agroforestry and local knowledge could contribute to efforts to conserve and restore rainforests, and reduce deforestation by accelerating fallow in tropical agriculture. 2005 Elsevier B.V. All rights reserved.

1.

Introduction

Land areas of southern Mexico are being deforested and losing productivity at alarming rates. In Chiapas, Mexico, deforestation is claiming 7% of the forest each year, and erosion has moderately degraded 10% to 25%, and severely degraded 5% of the arable soil (Howard and Homer-Dixon, 1996). These problems are endemic throughout the tropics,

as increasing population densities stress the environment through demands on agricultural land (Lal, 1995; Alvarez and Naughton-Treves, 2003). Because these areas are experiencing high population growth and movement (Ram, 1997), these problems will be magnied in future years. Displaced and migrant populations, in particular, have had a large effect on the ecological stability of this region (Nicholson et al., 1995; Atran, 1999; Mas and Puig, 2001). Land

Corresponding author. Tel.: +1 614 247 6133; fax: +1 614 292 9448. E-mail address: 1130@osu.edu (J.F. Martin). 0925-8574/$ see front matter 2005 Elsevier B.V. All rights reserved. doi:10.1016/j.ecoleng.2005.10.012

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management of these groups tends to take the form of cattle pasture (Mas and Puig, 2001; Durand and Lazos, 2004) or shortterm milpa (OBrien, 1998). Uncontrolled grazing leaves the land compacted and incapable of production, even for grazing, after a very short time (Garciaoliva et al., 1994; Durand and Lazos, 2004). Following crop and cattle production, these areas do not return to a mature, enriched forest, but to degraded grass and brush vegetation. These lands have very little productive use and relatively low biodiversity (Miller, 1999). Lands devastated by inappropriate use intensify the demand for new lands, leading to further deforestation and social conict (Nicholson et al., 1995; Howard and Homer-Dixon, 1996). Ecosystem management and restoration in this area of the world is a complex social, economic, and ecological problem (Nicholson et al., 1995). Tools that offer monetary incentives or sustenance may be important to any long-term solution to rainforest loss and restoration (Nicholson et al., 1995; Foroughbakhch et al., 2001; Li, 2004). Numerous researchers state the importance of recording indigenous knowledge to better understand sustainable land management (Fox et al., 2000; Long and Zhou, 2001; Hardwick et al., 2004). Because indigenous swidden agroforestry systems can be productive (Long and Nair, 1999) while maintaining their ecological integrity (Wang and Young, 2003), swidden practices could contribute to better land management in the tropics (De Clerck and Negreros-Castillo, 2000; Fox et al., 2000). The Lacandon Maya, an indigenous group that has met subsistence needs while maintaining both secondary and primary forests for centuries in southern Mexico, combine forest restoration and domestic production (Nations and Nigh, 1980; Levy, 2000; Diemont and Martin, 2005). The Lacandon land management system cycles through three eld stages starting with the milpa, progressing to the acahual (low secondary forest), and then to tall secondary forest, before returning to the milpa (Nations and Nigh, 1980; McGee, 2002). Neighboring primary forest is conserved to maintain a biodiverse seedbank (Quintana-Ascencio et al., 1996). Ecological succession drives the conversion between eld stages (Levy and Aguirre Rivera, in press). From the viewpoint of ecological succession, the milpa represents early successional grasses, the acahual represents the shrub or early woody stage, and the forest is the climax stage. The milpa, or early successional stage, is a polyculture eld that includes 2030 cultivated species. The acahual and forest stages are also productive, offering over 50 plant species used by the Lacandon (Nations and Nigh, 1980). By selecting for certain species and managing the natural succession of the acahual and forest stages, the Lacandon are able to restore soil fertility and regenerate secondary forest following the milpa stage in less than 20 years (Diemont and Martin, 2005). Lacandon Maya agroforestry is ecological engineering as described by Odum et al. (1963), where the technology available from natural systems is dominant, and human engineering is supplementary rather than primary. The Lacandon rely on the regenerative capacity of nature. They seed certain plants during the fallow and eliminate others, but in general allow the system to develop without intervention, permitting forcing functions such as sun, wind, and rain to drive the system (Diemont et al., 2006). Furthermore, the Lacandon system is as Mitsch and Jorgensen (1989) have described ecological

engineeringdesigned for the benet of both humans and the environment. At all stages of successional development, the Lacandon recover harvestable foods, medicines, and raw materials (Nations and Nigh, 1980). This production does not come at the cost of ecosystem health. The system largely selfdesigns and develops in biodiversity and complexity (Nations and Nigh, 1980; Levy, 2000); numerous animals are drawn to the richness of this ecosystem (Nations and Nigh, 1980). Previous studies have identied plants that the Lacandon may be using to restore soil fertility (Levy, 2004; Levy and Golicher, 2004). The presence of Ochroma pyramidale Urban, has been related to greater leaf-litter (Levy, 2004; Levy and Golicher, 2004) and soil organic matter accumulation (Levy and Golicher, 2004), indicating that the Lacandon manage their fallow to accelerate regeneration of soil fertility. An evaluation of soil chemistry, soil nematode populations, and plant community in all eld stages of the Lacandon system revealed seven plants that correlated positively with improved soil conditions (SAW Diemont, JF Martin, and MF Quigley, unpublished data in review). Hampea stipitata S. Watson, found in the acahual, correlated with elevated soil organic matter concentrations. Sapium lateriorum, found in secondary and primary forest, correlated with elevated ammonium and nitrate concentrations. Cecropia obtusifolia Bertol., discovered in milpa, acahual and secondary forest, and Piper auritum H.B.K., in acahual and secondary forest, correlated with higher bacterivorous nematode concentrations. These results encouraged additional research into the plants the Lacandon are using during the fallow to restore secondary forest. The objectives of this study were to: (1) through interviews, determine if the Lacandon have selected for certain species to accelerate the regeneration of soil fertility; (2) better identify the contribution of selected species (i.e. O. pyramidale) to soil fertility; (3) identify the mechanism by which selected species regenerate soil fertility; and (4) evaluate the potential for these methods to be used to restore and conserve tropical rainforests.

2.

Materials and methods

Interviews and soil sampling were conducted in Lacanja Chansayab, Chiapas, Mexico. Lacanja has a population of approximately 400 and is one of three major communities of the Lacandon Maya. Lacanja is located in Chiapas, the southern-most Mexican state, at 16 45 30 N and 91 08 30 W and at an elevation of 400 m. The predominate soil type is Luvisol (INEGI, 1982), with a clayey texture and neutral pH. The surrounding ecosystem is tall moist forest, and annual rainfall is 2500 cm (Guillen-Trujillo, 1998).

2.1.

Interviews

Interviews were conducted with ve Lacandon farmers to determine what plants they regarded as important for fertility regeneration during the fallow stages of the system. Farmers were asked rst to identify what plants were good for fertility regeneration in the fallow. They were asked if soil that resulted in the area where the plant was located would be good, whether the plant leaf litter produced good compost, whether the milpa stage that may follow in the location would

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Table 1 Mean (standard deviation) of the age (years) and diameter at breast height (DBH) (cm) of the Sapium lateriorum and Ochroma pyramidale trees sampled Tree
Sapium lateriorum Acahual Secondary Forest Primary forest Total Lattice Ochroma pyramidale Milpa Acahual Secondary forest Total Lattice

n
3 5 1 9 1

Age (year)
3.0(1.0) 16.4(6.2) 40 14.6(12.4) 25

DBH (cm)
26.7(16.3) 78.0 (46.8) 142 68.0 (50.5) 145

2 3 4 9 1

1 3.7 (0.6) 11.5(2.5) 6.6 (5.5) 8

18.0(1.4) 24.0 (6.1) 62.8 (7.1) 39.9 (22.5) 30

produce healthy cultivated species (i.e. maize). Farmers were also asked if the plants had alternative uses aside from fertility regeneration (i.e. construction materials). Plants identied by one farmer were subsequently asked of the following farmers who where interviewed. Species that received more than one negative response to any of these questions from subsequent farmers were discarded from the list of fertility enhancing trees.

Within a 1 m2 circular sampling area at each sampling location, leaf litter depth was measured at three points and averaged. Soil moisture and pH were evaluated using a Kelway soil acidity and moisture meter. The detrital layer was then removed, and eight replicate 2.5 cm diameter cores were taken from 020 cm soil depth and pooled. Each soil sample was analyzed for soil organic matter (SOM) (Walkley and Black, 1934), total nitrogen (N) (semi-microkjeldhal), available phosphorus (P) (Olsen et al., 1954), and texture (Bouyoucos, 1951). Nematodes were extracted from 20 g of the pooled soil from each sampling location for analysis. Soil samples for nematode extraction were kept cool and extracted over 48 h using the Baermann wet funnel technique (McSorley and Welter, 1991). Nematodes were heat xed. Extract was stored in 2% formaldehyde, and all nematodes in the bottom 10 mL of extract were identied to trophic level at 90 magnication (Parmelee and Alston, 1986; Edwards, 1991; Dominguez et al., 2003; Arancon et al., 2003). Nematodes were identied as plant parasite, fungivore, bacterivore, and omnivore-predator trophic groups according to Parmelee and Alston (1986). Omnivore/predator was reported as omnivore, as in Arancon et al. (2003).

2.3.

Analysis

2.2.

Field sampling

Based on interviews and literature review, two trees were chosen for soil sampling and analysis: Ochroma pyramidale and Sapium lateriorum. Soil sampling under Sapium lateriorum was conducted in November 2004 during the wet season. Soil sampling under Ochroma pyramidale was conducted in February 2005 during the intermediate wet-dry season. Trees in milpa, acahual, secondary forest, and primary forest were used for analysis. Three small, medium, and large trees of each species were identied for sampling. Approximate age and diameter at breast height (DBH) of each tree were determined (Table 1). Tree age was determined by a Lacandon farmer familiar with tree size relative to age and the specic history of the individual trees sampled. One transect was laid out through each tree, and sampling was conducted at seven locations along each transect. Sampling locations along transects were evenly spaced so that the sampling location second to the end at each side of each transect was at the canopy edge. The end sampling locations along each transect were outside the tree canopy. The center point of each transect was located next to the tree trunk. A lattice was laid out beneath the canopy of one tree with sixteen sampling locations at the intersection of lattice lines. The midpoint of lattice edge lines intersected with the canopy edge. Four additional points within the lattice were randomly selected. Non-sampled trees with a DBH greater than 15 cm and with a canopy over a sampling location were identied by Lacandon name and cross-referenced for Latin binomials. Voucher specimens for all plants not previously collected and identied in Nations and Nigh (1980) or Levy et al. (2002) were collected and deposited in the herbarium at El Colegio de la Frontera Sur, San Cristobal de Las Casas, Mexico.

Statistical analysis was conducted using Systat 10.2 computer software. Sampling points from beneath and outside the tree canopy were compared using independent sample Students t-tests. Relationships between factors were determined using Pearson linear regression. Sampling locations under canopy trees of the same species as the tree under consideration were removed. Outliers (|Studentized residuals| > 2.75) and points with large leverage (leverage > 0.3) were removed. Where transect and lattice data were pooled for regressions, the ratio of dependent and independent factors of lattice and transect points were compared using independent sample Students ttests to determine that differences were not present that may skew results. Where a difference was present, results were reported for transect points only.

3.
3.1.

Results
Interviews

Interviews with Lacandon farmers yielded 20 plants that were considered to be important for fertility regeneration (Table 2). Of these plants, 19 are useful to the Lacandon in some way apart from soil restoration. Uses included food, construction materials, furniture, arts and crafts, medicines, canoes, and food for wild birds. Due to uniformly strong interview responses and past research (Levy, 2004; and Levy and Golicher, 2004), Ochroma pyramidale and Sapium lateriorum were chosen for more in-depth assessment. O. pyramidale is a cultivated species that is planted in the milpa, and acahual in dense patches. S. lateriorum is not planted by the Lacandon because seeds are readily disbursed by birds, and an abundant amount of these plants can therefore be found in the secondary forest. Although other species are considered weedy and removed by the Lacandon, S. lateriorum is never removed from an acahual or secondary forest.

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Table 2 Plants identied by the Lacandon that assist soil fertility regeneration and restoration Name Latin binomial
Astrocaryum mexicanum Liebmann ex Martius Belotia mexicana K. Schum. Brosinum sp. Bucida buceras L. Calophyllum brasiliense Camb. var. rekoi Standl. Cedrela sp. Cordia alliodora Oken Dialium guihense Sandw. Guatteria anomala R. E. Fries Hampea stipitata S. Watson Hibiscus sp. Mucuna pruriens L. Ochroma pyramidale Urban Piper auritum H. B. K. Piper aduncum L Sapium lateriorum Hemsl. Simira salvadorensis Standl. Sterculia apetala Jacq. Swietenia macrophylla King Unidentied

Use (Levy 2002; Interviews) Lacandon

ak te taw ba am bax sa puk te baba cedro no (Spanish) bajum wech ek bache tsuk tok jor ka a be chujum jober makarum ucunte chakax anis puna pok te Fruit Construction Fruit Construction and rewood Construction and food for birds Construction and carved art pieces Construction Construction, food, and rewood Food and construction Food and rope Rope for making bags and hammocks Drink Construction and medicine for backpains Eat leaf with sh, wrap tamales Construction Seeds eaten by birds, lumber for construction Medicine for skin cuts Beeds for necklaces Construction, furniture, and canoes Not determined

3.2.

Sapium Lateriorum

3.3.

Ochroma pyramidale

S. Lateriorum was evaluated (Table 1) outside and beneath the tree canopy. The sampling locations at the edge of the tree canopy were considered to be inuenced by the tree canopy and were therefore allocated to beneath the tree canopy; only the transect points completely outside the tree canopy were considered outside the tree canopy. P was 16% greater beneath the tree canopy (10.1 mg/kg) than outside the tree canopy (8.7 mg/kg) (P = 0.027, Fig. 1). To evaluate if this trend was due to S. Lateriorum pioneering P-rich areas, P was regressed as function of DBH. P increased with DBH (Fig. 2). Trends with respect to N, SOM, pH, and nematodes were not discovered.

Pooling all sampling locations for O. pyramidale resulted in no statistically signicant trends (alpha = 0.05) with respect to distance from trees or between parameters (i.e. between total nematodes and soil organic matter). By pooling all trees to evaluate the effect of distance from tree, sampling locations likely incorporated some overlap between those outside the canopy and beneath the canopy. Mean distance to sampling locations outside the canopy was 6.8 m with standard deviation of 1.9. In addition, tree sizes were highly variable; mean DBH was 40 cm with a standard deviation of 21 cm. In order to remove some of this overlap and better homogenize the tree size, trees in the milpa, acahual, and secondary forest were

Fig. 1 Mean available soil phosphorus concentrations beneath and outside the canopy cover of Sapium Lateriorum. Statistical results are for independent sample t-test. Error bars are 1 S.E.

Fig. 2 Available soil phosphorus concentrations beneath the tree canopy as a function of diameter at breast height of Sapium Lateriorum.

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Fig. 3 (af) Leaf litter (a), soil organic matter (transect only) (b), total nitrogen (c), total nematodes (d), bacterivore nematodes (e), and omnivore nematodes (f) as a function of distance from Ochroma pyramidale trees in the secondary forest.

analyzed separately (Table 1). For example, in the secondary forest, the distance to the sampling point outside the canopy increased to 7.8, while the standard deviation decreased to 1.0. Mean DBH increased to 63, and standard deviation decreased to 6.1 (Table 1). Considering only trees in the secondary forest revealed several trends (Fig. 3). To compensate for reduced n in examining trees in the secondary forest, transect and lattice sampling locations were pooled (see 2.3 Analysis). Log-transformed leaf litter depth decreased as a function of distance from O. Pyramidale (Fig. 3a). Mean leaf litter at the tree was over 6 cm and

decreased to less than 3 cm outside the tree canopy. SOM (transect only) (Fig. 3b), total N (Fig. 3c), and P (R = 0.047, P = 0.78) did not exhibit any signicant trends. Nematodes displayed trends opposite to that of leaf litter depth. Total nematodes (Fig. 3d) increased from 8 per 20 g soil at the trunk to over 60 at a distance of 14 m. Bacterivorous nematodes (Fig. 3e) increased by an order of magnitude from 4 per 20 g soil at the trunk to nearly 40 outside the tree canopy. Omnivorous (Fig. 3f) nematodes increased from 5 per 20 g soil at the trunk to nearly 15 outside the canopy. Trees in the milpa and acahual did not exhibit any trends with respect to soil nematodes or chemistry.

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4.

Discussion

The numerous plants identied by ve Lacandon farmers as being important for soil fertility (Table 2) and the results of this study lend support to the hypothesis by Levy (2004) and Levy and Golicher (2004) that the Lacandon are actively enhancing the soil of their fallow stages through plant management. Whereas other trees are removed during milpa and fallow stages (Nations and Nigh, 1980; Diemont and Martin, 2005), the trees identied by the Lacandon are either cultivated (e.g. Ochroma pyramidale) or are left in the fallow following aeolian or animal seed disbursal (e.g. Sapium lateriorum and Piper spp.). That most of these plants are also useful (Table 2) suggests that production does not need to come at the cost of fertility regeneration, or vice versa (Nations and Nigh, 1980; Diemont and Martin, 2005). Lacandon farmers are able to restore the rainforest during the fallow, while at the same time producing numerous useful crops (Nations and Nigh, 1980). Levy (2002) recorded over 400 plants that are useful to the Lacandon that occur in the fallow and primary forest. The productivity of secondary vegetation allows the Lacandon to maintain their lands in fallow for longer periods of over twenty years (Diemont and Martin, 2005) and conserve primary forest for medicines, raw materials, food and ecosystem services (Nations and Nigh, 1980; Levy, 2002). In comparison, the fallow duration has been reduced to less than 10 years in many swidden systems in the Yucatan, Mexico (Weisbach et al., 2002). Furthermore, Lacandon plant management to enhance soil fertility may shorten the time needed to restore mature forest to sites previously used for agriculture (Levy and Golicher, 2004). Reducing the time needed to restore soil fertility and enhancing the productivity of fallows are both important ways for protecting primary forest. Determining management techniques that allow simultaneous restoration of soil fertility and productivity, as does the Lacandon system, could reduce deforestation throughout the world. Traditional Lacandon farmers do not use fertilizer (Nations and Nigh, 1980; Levy and Aguirre, 1999; Diemont and Martin, 2005). In a geographic area where phosphorous is typically unavailable, P could become a limiting factor in agricultural production (Bautista-Cruz and del Castillo, 2005). Whereas N can be introduced from the atmosphere by N xing leguminous herbs and trees (Badejo, 1998; Okogun et al., 2000), P can only enter from runoff and plant extraction from the soil or sub-soil. Consequently, the elevated levels of P beneath S. lateriorum (Table 2) and the increasing P concentrations with DBH (Fig. 2) support the conclusion that this plant, which is managed by the Lacandon, serves as a phosphorus pump (Perezllorens et al., 1993; Badejo, 1998). Vejre and Hoppe (1998) describe such a pumping effect of K, P, Ca, and Mg, where minerals in the lower soil proles were brought to surface by deeper tree roots in a forest in Denmark. Similarly, Christanty et al. (1997) found that the nutrient pumping provided by Phyllostachys sp. in an Indonesian agroforestry system was vital to maintaining fertility of the surface soils. The results for Ochroma pyramidale indicate that nematode populations are inhibited by O. pyramidale leaf litter (Fig. 3). Leaf litter concentration decreased with distance from the tree, whereas nematodes were found to increase (Fig. 3).

Nematodes would generally decrease in number with distance from the tree along with leaf litter depth (Fig. 3a) if an inhibitory effect were not present. Ferris and Matute (2003) and Matlack (2001) found that the abundance of bacterivores were elevated with the addition of organic matter. Bjornlund and Christensen (2005) found that leaf litter addition resulted in bacterivorous nematode spikes followed by fungivorous nematode dominance as the leaf litter was broken down. In other successional swidden systems, similar to the Lacandon system, fallow length, and thus greater leaf litter accumulation, appeared to increase the abundance of nematodes (Pate et al., 2000; Villenave et al., 2001). The correlation between nematodes, litter, and distance from trees was not noted in younger trees (e.g. trees less than four years old), but only in trees found in the secondary forest; this result indicates that the inhibition may be a cumulative effect. Chemical analysis of O. pyramidale showed an abundance of soluble saccharides compared to starch (Marenco et al., 2001). ODowd (1979) found that O. pyramidale produces nectar on the leaves of young trees. The abundance of foliar nectar or other secondary compounds deposited over the course of early succession may inhibit nematode populations beneath the canopy. In addition, Molofsky and Augspurger (1992) found that establishment of other species of tree seedlings underneath O. pyramidale was limited during seed germination, not seed deposition, which may indicate a possible allelopathic effect. Leaf litter and thus organic matter beneath O. pyramidale are abundant (Levy, 2004; Levy and Golicher, 2004), an unusual situation in tropical rainforests where conditions favor rapid bacterial and fungal decomposition (Attignon et al., 2004). Dense planting of O. pyramidale may allow Lacandon farmers to accumulate and store leaf litter and organic matter for future use. O. pyramidale is an early succession species, which would permit the next successional stage to have an abundance of soil organic matter available at the outset. That no trend was noted for soil organic matter as a function of distance from the tree despite the abundance of leaf litter closer to the tree further supports this idea. Levy and Golicher (2004) discovered higher soil organic matter near O. pyramidale trees compared to where O. pyramidale was not present, but this difference was patchy and inconsistent between plots. Other species represented on the list of fertility-enhancing trees (Table 2) may contribute to the accumulation of leaf litter during the fallow. Piper auritum contains Safrol in its leaves, which has been shown to have high cytotoxic effects, and Piperacae in general are notoriously allelopathic (Anaya-Lang, 1979, Gupta et al., 1996, Xuan et al., 2004). The Lacandon favor O. pyramidale in their fallows for fertility enhancement, but farmers stated in the interviews that these trees will only grow in a quarter of their fallow lands. Therefore, a mosaic of other trees that have similar functions may be important for managing restoration.

5.

Conclusions

Lacandon Maya indigenous agroforesty uses numerous trees for soil restoration during fallow. These trees also produce food, medicines, and raw materials. Results indicate that one

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of these trees, Sapium laterijlorum, acts as a phosphorus pump, restoring this nutrient to the soil during fallow. Ochroma pyramidale appears to be slowing organic matter degradation and allowing storage of organic matter in leaf litter. Future research should verify these results with experimental testing and examine the use of other trees identied by the Lacandon for soil fertility regeneration. Lacandon land management appears to offer new tools for rainforest restoration and soil fertility enhancement.

Acknowledgements
We wish to thank Manuel Castellanos, Kin Bor, Adolfo Chan Kin, Poncho Kin, Jorge Paneagua, Vicente Paneagua, Enrique Paneagua, and Kin Paneagua. Financial support from the Ohio Agricultural Research and Development Center, the Fulbright Program, and NSF Grant OISE-0431230 is gratefully acknowledged.

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