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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 r t i c l e
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 ﬁeld and fallow stages that produce food, medicines, and raw materials, and regenerates tall secondary forest. This investigation identiﬁed plants managed by Lacandon to restore soil fertility during fallow. Through interviews, Lacandon identiﬁed 20 plants managed for forest restoration. Leaf litter measurements and soil samples were taken near two of these species, Ochroma pyramidale and Sapium lateriﬂorum. 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 lateriﬂorum
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 ﬁndings indicated an inhibition of degradation that permits accelerated soil organic matter accumulation. Available phosphorus (P) concentrations beneath S. lateriﬂorum 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 fulﬁll 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.
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 magniﬁed 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: firstname.lastname@example.org (J.F. Martin). 0925-8574/$ – see front matter © 2005 Elsevier B.V. All rights reserved. doi:10.1016/j.ecoleng.2005.10.012
2004) and soil organic matter accumulation (Levy and Golicher. 1999). an indigenous group that has met subsistence needs while maintaining both secondary and primary forests for centuries in southern Mexico. in acahual and secondary forest. O. 1980). pyramidale) to soil fertility. economic. 1994. Cecropia obtusifolia Bertol. 2002). whether the milpa stage that may follow in the location would . The milpa. 2004). At all stages of successional development. but in general allow the system to develop without intervention. Tools that offer monetary incentives or sustenance may be important to any long-term solution to rainforest loss and restoration (Nicholson et al. the milpa represents early successional grasses. has been related to greater leaf-litter (Levy. the acahual represents the shrub or early woody stage. Foroughbakhch et al. 1980. permitting forcing functions such as sun. An evaluation of soil chemistry. is a polyculture ﬁeld that includes 20–30 cultivated species. discovered in milpa. and plant community in all ﬁeld stages of the Lacandon system revealed seven plants that correlated positively with improved soil conditions (SAW Diemont. these areas do not return to a mature. Watson. The presence of Ochroma pyramidale Urban. Levy. found in the acahual. The system largely selfdesigns and develops in biodiversity and complexity (Nations and Nigh. 2004). correlated with elevated ammonium and nitrate concentrations. Long and Zhou. Numerous researchers state the importance of recording indigenous knowledge to better understand sustainable land management (Fox et al. Uncontrolled grazing leaves the land compacted and incapable of production.. 1980.. Furthermore. The objectives of this study were to: (1) through interviews. Interviews Interviews were conducted with ﬁve Lacandon farmers to determine what plants they regarded as important for fertility regeneration during the fallow stages of the system. Farmers were asked ﬁrst to identify what plants were good for fertility regeneration in the fallow. 2000. Howard and Homer-Dixon. Previous studies have identiﬁed plants that the Lacandon may be using to restore soil fertility (Levy. swidden practices could contribute to better land management in the tropics (De Clerck and Negreros-Castillo. 1982). 2. These results encouraged additional research into the plants the Lacandon are using during the fallow to restore secondary forest. and then to tall secondary forest. and ecological problem (Nicholson et al. Following crop and cattle production. 2005). even for grazing.B. They seed certain plants during the fallow and eliminate others. These lands have very little productive use and relatively low biodiversity (Miller.. This production does not come at the cost of ecosystem health. and human engineering is supplementary rather than primary. 2004) or shortterm milpa (O’Brien. The Lacandon rely on the regenerative capacity of nature.e. 2004). after a very short time (Garciaoliva et al.. 1995. 2005). Durand and Lazos. at 16◦ 45 30 N and 91◦ 08 30 W and at an elevation of 400 m. the Lacandon system is as Mitsch and Jorgensen (1989) have described ecological engineering—designed for the beneﬁt of both humans and the environment. Mexico. McGee. Levy and Golicher... leading to further deforestation and social conﬂict (Nicholson et al.1. the Lacandon are able to restore soil fertility and regenerate secondary forest following the milpa stage in less than 20 years (Diemont and Martin. Li. 1995). (3) identify the mechanism by which selected species regenerate soil fertility. Fox et al. Hardwick et al. in press). combine forest restoration and domestic production (Nations and Nigh. or early successional stage. The acahual and forest stages are also productive. 2004. 2. before returning to the milpa (Nations and Nigh. Lacandon Maya agroforestry is ecological engineering as described by Odum et al. wind. 2004). numerous animals are drawn to the richness of this ecosystem (Nations and Nigh. JF Martin. Hampea stipitata S. Because indigenous swidden agroforestry systems can be productive (Long and Nair. Durand and Lazos. 2000). progressing to the acahual (low secondary forest). Ecosystem management and restoration in this area of the world is a complex social. and rain to drive the system (Diemont et al. Lands devastated by inappropriate use intensify the demand for new lands. and raw materials (Nations and Nigh. (1963).. the southern-most Mexican state. and (4) evaluate the potential for these methods to be used to restore and conserve tropical rainforests.K.. 1980). whether the plant leaf litter produced good compost. 1996). acahual and secondary forest. The Lacandon Maya. the Lacandon recover harvestable foods. 2006).. and MF Quigley. but to degraded grass and brush vegetation. Materials and methods Interviews and soil sampling were conducted in Lacanja Chansayab. Ecological succession drives the conversion between ﬁeld stages (Levy and Aguirre Rivera. 2004. By selecting for certain species and managing the natural succession of the acahual and forest stages. and the forest is the climax stage. and Piper auritum H. Neighboring primary forest is conserved to maintain a biodiverse seedbank (Quintana-Ascencio et al. 2004). 1980. 1995. where the technology available from natural systems is dominant. 2001. unpublished data in review). 2003). Diemont and Martin. 1998). 1980). 2001. Lacanja is located in Chiapas. soil nematode populations. From the viewpoint of ecological succession. 1999) while maintaining their ecological integrity (Wang and Young.206 e c o l o g i c a l e n g i n e e r i n g 2 8 ( 2 0 0 6 ) 205–212 management of these groups tends to take the form of cattle pasture (Mas and Puig. Chiapas. Sapium lateriﬂorum. Lacanja has a population of approximately 400 and is one of three major communities of the Lacandon Maya. with a clayey texture and neutral pH. 2000. enriched forest. Levy. 2000. and annual rainfall is 2500 cm (Guillen-Trujillo. The Lacandon land management system cycles through three ﬁeld stages starting with the milpa. 1998). 2000). 2001. medicines. determine if the Lacandon have selected for certain species to accelerate the regeneration of soil fertility. offering over 50 plant species used by the Lacandon (Nations and Nigh. correlated with higher bacterivorous nematode concentrations.. The predominate soil type is Luvisol (INEGI. correlated with elevated soil organic matter concentrations.. They were asked if soil that resulted in the area where the plant was located would be good.. Levy and Golicher. The surrounding ecosystem is tall moist forest. 1996). (2) better identify the contribution of selected species (i. indicating that the Lacandon manage their fallow to accelerate regeneration of soil fertility. found in secondary and primary forest.
medicines. The midpoint of lattice edge lines intersected with the canopy edge.e. Extract was stored in 2% formaldehyde.. Four additional points within the lattice were randomly selected. 1934). 2. Of these plants.0(1. and primary forest were used for analysis. The detrital layer was then removed. as in Arancon et al. Uses included food.4) 25 DBH (cm) 26. and omnivore-predator trophic groups according to Parmelee and Alston (1986). (2002) were collected and deposited in the herbarium at El Colegio de la Frontera Sur. arts and crafts. and eight replicate 2. A lattice was laid out beneath the canopy of one tree with sixteen sampling locations at the intersection of lattice lines. Where a difference was present. Dominguez et al. the ratio of dependent and independent factors of lattice and transect points were compared using independent sample Student’s ttests to determine that differences were not present that may skew results. 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.3) were removed. Sampling locations under canopy trees of the same species as the tree under consideration were removed.5 cm diameter cores were taken from 0–20 cm soil depth and pooled. 2004). S. 2003. Nematodes were heat ﬁxed.5) 30 produce healthy cultivated species (i. Although other species are considered weedy and removed by the Lacandon. construction materials.5) 6.2) 40 14. two trees were chosen for soil sampling and analysis: Ochroma pyramidale and Sapium lateriﬂorum. Arancon et al. S.7 (0. Voucher specimens for all plants not previously collected and identiﬁed in Nations and Nigh (1980) or Levy et al. secondary forest. maize).e c o l o g i c a l e n g i n e e r i n g 2 8 ( 2 0 0 6 ) 205–212 207 Table 1 – Mean (standard deviation) of the age (years) and diameter at breast height (DBH) (cm) of the Sapium lateriﬂorum and Ochroma pyramidale trees sampled Tree Sapium lateriﬂorum 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) 16. 1991. 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. 2003)..6) 11. Soil sampling under Sapium lateriﬂorum was conducted in November 2004 during the wet season. 3. bacterivore. 1986.. Each soil sample was analyzed for soil organic matter (SOM) (Walkley and Black. leaf litter depth was measured at three points and averaged.5) 8 18. Tree age was determined by a Lacandon farmer familiar with tree size relative to age and the speciﬁc history of the individual trees sampled.1) 39. Statistical analysis was conducted using Systat 10. Due to uniformly strong interview responses and past research (Levy. Three small.5) 145 2 3 4 9 1 1 3. Edwards. (2003). pyramidale is a cultivated species that is planted in the milpa. fungivore. 2004. Results Interviews Interviews with Lacandon farmers yielded 20 plants that were considered to be important for fertility regeneration (Table 2). canoes.2.6(12. Analysis 2.4(6. Plants identiﬁed by one farmer were subsequently asked of the following farmers who where interviewed. Field sampling Based on interviews and literature review. available phosphorus (P) (Olsen et al. Sampling points from beneath and outside the tree canopy were compared using independent sample Student’s t-tests. acahual. lateriﬂorum is never removed from an acahual or secondary forest. Outliers (|Studentized residuals| > 2. 3. Where transect and lattice data were pooled for regressions.5(2. medium.3) 78.2 computer software.0 (50. construction materials). lateriﬂorum is not planted by the Lacandon because seeds are readily disbursed by birds.e. total nitrogen (N) (semi-microkjeldhal). Nematodes were identiﬁed as plant parasite. Omnivore/predator was reported as omnivore. San Cristobal de Las Casas. One transect was laid out through each tree. and Levy and Golicher. furniture. Approximate age and diameter at breast height (DBH) of each tree were determined (Table 1).0 (6.0(1.4) 24. Soil moisture and pH were evaluated using a Kelway soil acidity and moisture meter. Relationships between factors were determined using Pearson linear regression. and food for wild birds. The end sampling locations along each transect were outside the tree canopy.3. Trees in milpa.75) and points with large leverage (leverage > 0. . and all nematodes in the bottom 10 mL of extract were identiﬁed to trophic level at 90× magniﬁcation (Parmelee and Alston. 1954).9 (22. 19 are useful to the Lacandon in some way apart from soil restoration. 1951). 1991). results were reported for transect points only. Mexico. Ochroma pyramidale and Sapium lateriﬂorum were chosen for more in-depth assessment.0 (46.1) 62.1. O. and an abundant amount of these plants can therefore be found in the secondary forest. The center point of each transect was located next to the tree trunk. and large trees of each species were identiﬁed for sampling.8) 142 68.8 (7. and sampling was conducted at seven locations along each transect. Nematodes were extracted from 20 g of the pooled soil from each sampling location for analysis. Farmers were also asked if the plants had alternative uses aside from fertility regeneration (i.7(16. Soil samples for nematode extraction were kept cool and extracted over 48 h using the Baermann wet funnel technique (McSorley and Welter. and acahual in dense patches. Non-sampled trees with a DBH greater than 15 cm and with a canopy over a sampling location were identiﬁed by Lacandon name and cross-referenced for Latin binomials. and texture (Bouyoucos.6 (5. Soil sampling under Ochroma pyramidale was conducted in February 2005 during the intermediate wet-dry season.
2.e. Fries Hampea stipitata S. and nematodes were not discovered.3. Mucuna pruriens L. Sterculia apetala Jacq.8 m with standard deviation of 1. lumber for construction Medicine for skin cuts Beeds for necklaces Construction. and secondary forest were Fig. Fig. Piper aduncum L Sapium lateriﬂorum Hemsl. Trends with respect to N. In order to remove some of this overlap and better homogenize the tree size. 2 – Available soil phosphorus concentrations beneath the tree canopy as a function of diameter at breast height of Sapium Lateriﬂorum. To evaluate if this trend was due to S.9. Interviews) Lacandon ak te taw ba am bax sa puk te baba cedro ﬁno (Spanish) bajum wech’ ek bache ts’uk tok jor ka a be chujum jo’ber makarum u’cunte chak’ax anis puna pok te Fruit Construction Fruit Construction and ﬁrewood Construction and food for birds Construction and carved art pieces Construction Construction. between total nematodes and soil organic matter). By pooling all trees to evaluate the effect of distance from tree.1 mg/kg) than outside the tree canopy (8. 1 – Mean available soil phosphorus concentrations beneath and outside the canopy cover of Sapium Lateriﬂorum. Ochroma pyramidale S. food. Pooling all sampling locations for O. Schum. The sampling locations at the edge of the tree canopy were considered to be inﬂuenced by the tree canopy and were therefore allocated to beneath the tree canopy. Cedrela sp. var. P increased with DBH (Fig. Mean distance to sampling locations outside the canopy was 6. Statistical results are for independent sample t-test. Sapium Lateriﬂorum 3.027. SOM. Ochroma pyramidale Urban Piper auritum H. Simira salvadorensis Standl. Lateriﬂorum was evaluated (Table 1) outside and beneath the tree canopy. tree sizes were highly variable. Cordia alliodora Oken Dialium guihense Sandw. trees in the milpa. .208 e c o l o g i c a l e n g i n e e r i n g 2 8 ( 2 0 0 6 ) 205–212 Table 2 – Plants identiﬁed by the Lacandon that assist soil fertility regeneration and restoration Name Latin binomial Astrocaryum mexicanum Liebmann ex Martius Belotia mexicana K. Lateriﬂorum pioneering P-rich areas. Brosinum sp. Fig. In addition. Bucida buceras L. 1). acahual. furniture. rekoi Standl. Swietenia macrophylla King Unidentiﬁed Use (Levy 2002. Calophyllum brasiliense Camb. only the transect points completely outside the tree canopy were considered outside the tree canopy. K. sampling locations likely incorporated some overlap between those outside the canopy and beneath the canopy.05) with respect to distance from trees or between parameters (i. P was regressed as function of DBH. B.E. E. 2). pyramidale resulted in no statistically signiﬁcant trends (alpha = 0. and ﬁrewood Food and construction Food and rope Rope for making bags and hammocks Drink Construction and medicine for backpains Eat leaf with ﬁsh.7 mg/kg) (P = 0. Error bars are ± 1 S. Guatteria anomala R. mean DBH was 40 cm with a standard deviation of 21 cm. wrap tamales Construction Seeds eaten by birds. Watson Hibiscus sp. and canoes Not determined 3. pH. P was 16% greater beneath the tree canopy (10.
3e) increased by an order of magnitude from 4 per 20 g soil at the trunk to nearly 40 outside the tree canopy.047. in the secondary forest. 3 – (a–f) Leaf litter (a). Mean leaf litter at the tree was over 6 cm and decreased to less than 3 cm outside the tree canopy. 3b). P = 0. 3f) nematodes increased from 5 per 20 g soil at the trunk to nearly 15 outside the canopy. Log-transformed leaf litter depth decreased as a function of distance from O. 3c). soil organic matter (transect only) (b). SOM (transect only) (Fig. total nematodes (d).8. Pyramidale (Fig. analyzed separately (Table 1). Nematodes displayed trends opposite to that of leaf litter depth. 3a).1 (Table 1). 3d) increased from 8 per 20 g soil at the trunk to over 60 at a distance of 14 m. and P (R = 0. and standard deviation decreased to 6.0. transect and lattice sampling locations were pooled (see 2. Total nematodes (Fig.e c o l o g i c a l e n g i n e e r i n g 2 8 ( 2 0 0 6 ) 205–212 209 Fig. Considering only trees in the secondary forest revealed several trends (Fig. 3). For example.78) did not exhibit any signiﬁcant trends. Bacterivorous nematodes (Fig. and omnivore nematodes (f) as a function of distance from Ochroma pyramidale trees in the secondary forest. . the distance to the sampling point outside the canopy increased to 7. Omnivorous (Fig. total nitrogen (c). while the standard deviation decreased to 1. To compensate for reduced n in examining trees in the secondary forest.3 Analysis). Trees in the milpa and acahual did not exhibit any trends with respect to soil nematodes or chemistry. Mean DBH increased to 63. total N (Fig. bacterivore nematodes (e).
pyramidale showed an abundance of soluble saccharides compared to starch (Marenco et al. 1979. 2005). pyramidale leaf litter (Fig. Levy and Golicher. this result indicates that the inhibition may be a cumulative effect. fallow length. pyramidale produces nectar on the leaves of young trees. pyramidale was limited during seed germination. appeared to increase the abundance of nematodes (Pate et al. 2005). 1980. Mexico (Weisbach et al. 2004. P. raw materials... Diemont and Martin. 1998). 1993. 1980. pyramidale are abundant (Levy. not seed deposition. 2000). 2004). In comparison. Leaf litter and thus organic matter beneath O. which may indicate a possible allelopathic effect.. and raw materials... P can only enter from runoff and plant extraction from the soil or sub-soil. Diemont and Martin. 2004). Levy and Golicher (2004) discovered higher soil organic matter near O. but only in trees found in the secondary forest. Diemont and Martin. The productivity of secondary vegetation allows the Lacandon to maintain their lands in fallow for longer periods of over twenty years (Diemont and Martin. could reduce deforestation throughout the world. Okogun et al. Results indicate that one . Gupta et al. Christanty et al. Vejre and Hoppe (1998) describe such a pumping effect of K. P could become a limiting factor in agricultural production (Bautista-Cruz and del Castillo. the trees identiﬁed by the Lacandon are either cultivated (e. Xuan et al.g. but this difference was patchy and inconsistent between plots. Traditional Lacandon farmers do not use fertilizer (Nations and Nigh. in an Indonesian agroforestry system was vital to maintaining fertility of the surface soils. Reducing the time needed to restore soil fertility and enhancing the productivity of fallows are both important ways for protecting primary forest. Lacandon farmers are able to restore the rainforest during the fallow. litter. 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. Consequently. O. Levy (2002) recorded over 400 plants that are useful to the Lacandon that occur in the fallow and primary forest. and Piperacae in general are notoriously allelopathic (Anaya-Lang. 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. but farmers stated in the interviews that these trees will only grow in a quarter of their fallow lands. Piper auritum contains Safrol in its leaves. Furthermore. pyramidale is an early succession species. while at the same time producing numerous useful crops (Nations and Nigh. Chemical analysis of O. and distance from trees was not noted in younger trees (e. the fallow duration has been reduced to less than 10 years in many swidden systems in the Yucatan. Conclusions Lacandon Maya indigenous agroforesty uses numerous trees for soil restoration during fallow. Similarly. These trees also produce food. 2005) and conserve primary forest for medicines. and thus greater leaf litter accumulation. 1998. 2001). whereas nematodes were found to increase (Fig. 3a) if an inhibitory effect were not present.g. similar to the Lacandon system. Badejo.. Villenave et al. pyramidale may allow Lacandon farmers to accumulate and store leaf litter and organic matter for future use. Whereas N can be introduced from the atmosphere by N ﬁxing leguminous herbs and trees (Badejo. a mosaic of other trees that have similar functions may be important for managing restoration. Therefore. food and ecosystem services (Nations and Nigh. 3). The results for Ochroma pyramidale indicate that nematode populations are inhibited by O. lateriﬂorum (Table 2) and the increasing P concentrations with DBH (Fig. trees less than four years old). Ferris and Matute (2003) and Matlack (2001) found that the abundance of bacterivores were elevated with the addition of organic matter. In a geographic area where phosphorous is typically unavailable. In other successional swidden systems. and Mg. as does the Lacandon system.. Determining management techniques that allow simultaneous restoration of soil fertility and productivity. 2002). The Lacandon favor O. Discussion The numerous plants identiﬁed by ﬁve 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. Levy. 1996. where minerals in the lower soil proﬁles were brought to surface by deeper tree roots in a forest in Denmark. Leaf litter concentration decreased with distance from the tree. 2004). pyramidale in their fallows for fertility enhancement.210 e c o l o g i c a l e n g i n e e r i n g 2 8 ( 2 0 0 6 ) 205–212 4. 2001). 2) support the conclusion that this plant. or vice versa (Nations and Nigh. Dense planting of O. 3). 2000. Other species represented on the list of fertility-enhancing trees (Table 2) may contribute to the accumulation of leaf litter during the fallow. pyramidale trees compared to where O. Nematodes would generally decrease in number with distance from the tree along with leaf litter depth (Fig. 1999.. That most of these plants are also useful (Table 2) suggests that production does not need to come at the cost of fertility regeneration. In addition. 1980. which would permit the next successional stage to have an abundance of soil organic matter available at the outset. serves as a “phosphorus pump” (Perezllorens et al. pyramidale was not present. Sapium lateriﬂorum and Piper spp. 2004). an unusual situation in tropical rainforests where conditions favor rapid bacterial and fungal decomposition (Attignon et al.. the elevated levels of P beneath S. 2005). 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. The correlation between nematodes. Levy and Aguirre. The abundance of foliar nectar or other secondary compounds deposited over the course of early succession may inhibit nematode populations beneath the canopy. Molofsky and Augspurger (1992) found that establishment of other species of tree seedlings underneath O. medicines. (1997) found that the nutrient pumping provided by Phyllostachys sp. 2005).g. Ca.). 1980). Whereas other trees are removed during milpa and fallow stages (Nations and Nigh. Ochroma pyramidale) or are left in the fallow following aeolian or animal seed disbursal (e. 5. 1980. which has been shown to have high cytotoxic effects. 2002). O’Dowd (1979) found that O. which is managed by the Lacandon.
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