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Facilitating Regeneration of Secondary Forests with the Use of Mixed and Pure Plantations of Indigenous Tree Species Neâlida J

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Facilitating Regeneration of Secondary Forests with the Use of Mixed and Pure Plantations of Indigenous Tree Species Neâlida J Forest Ecology and Management 163 (2002) 217±227 Facilitating regeneration of secondary forests with the use of mixed and pure plantations of indigenous tree species NeÂlida J. Carnevalea, Florencia Montagninib,* aFaculty of Agricultural Sciences, National University of Rosario, C.C. 14 (S2125-ZAA)Zavalla, Santa Fe, Argentina bManagement and Conservation of Forests and Biodiversity, Centro AgronoÂmico Tropical de InvestigacioÂn y EnsenÄanza (CATIE), 7170 Turrialba, Costa Rica Received 27 December 2000; accepted 28 April 2001 Abstract The establishment of tree plantations on degraded lands can facilitate the regeneration of native species that could not otherwise grow in open micro sites or in competition by herbaceous species. The present research investigated tree regeneration under mixed and pure plantations of native species at La Selva Biological Station in the Atlantic humid lowlands of Costa Rica. The highest abundance of regenerating tree individuals was found in the understory of the mixed plantation (composed of Hieronyma alchorneoides Vochysia ferruginea Balizia elegans Genipa americana) with 10,156 indivi- duals/ha, followed by pure plantation of H. alchorneoides with 7891, V. ferruginea with 5703, B. elegans with 4219, G. americana with 1484, and the natural regeneration control with 703 individuals/ha. The highest mean number of species was found in the understory of the mixed plantation (11 species in 32 m2), followed by pure plantation of V. ferruginea (8.0), H. alchorneoides (7.0), B. elegans (5.0), G. americana (3.0) and control (1.0). Melastomataceae was the most abundant family in the understory of the mixed plantation and in the pure plantation of V. ferruginea, while Rubiaceae was the most abundant under H. alchorneoides. In constrast, Piperaceae was the most abundant family under pure plantations of G. americana and B. elegans. In addition, species of primary forest of the region were found in the understory of the mixed plantation and under the pure plantations of H. alchorneoides and V. ferruginea. These are promising indicators for the use of these plantations as accelerators of natural forest succession in the region. Very few woody species were found in the control, natural regeneration plots. Factors that may impede the establishment of woody species in the control may be the lack of perches for seed dipersers and invasion by herbaceous vegetation which outcompetes the tree seedlings in their growth. # 2002 Elsevier Science B.V. All rights reserved. Keywords: Competition; Dispersers; Litter; Mixed plantations; Regeneration; Shade 1. Introduction excess of soil humidity, high solar radiation, and intra- and inter-speci®c competition (Nepstad et al., 1991). The principal factors limiting tree regeneration in Another limiting factor is seed availability, especially abandoned pastures in tropical humid regions can in sites whose distance to seed sources may limit include scarcity of nutrients, soil compaction, lack or propagule dispersal. Several tree species exist in low densities in the natural forests surrounding the pastures, diminishing the possibilities of dispersal * Corresponding author. Present address: School of Forestry and Environmental Studies, Yale University, 370 Propect St., New of these species. Although the presence of perches Haven, CT 06511, USA. favors seed dispersal, other factors such as seed 0378-1127/02/$ ± see front matter # 2002 Elsevier Science B.V. All rights reserved. PII: S 0378-1127(01)00581-3 218 N.J. Carnevale, F. Montagnini / Forest Ecology and Management 163 (2002)217±227 predation may limit seed survival and germination in especially on the ®rst 2±3 years (Montagnini et al., abandoned pastures (Mc Clanahan and Wolfe, 1993; 1995). Holl, 1998). Trampling by cattle may be a main factor Mixed plantations could promote the regeneration leading to the elimination of seedlings growing in of a greater diversity of species in their understory pastures (Harvey and Haber, 1999). than pure-species plantations by creating a greater The presence of trees, either in plantation, in variability of habitat conditions that may favor seed groups, in lines or in isolated form may contribute to dispersers and germination and growth of tree species the recovery of environmental conditions favorable to (Guariguata et al., 1995; Montagnini et al., 1999). In tree regeneration processes (Parrotta, 1995; Guari- the present research we investigated tree regeneration guata et al., 1995). For example, in pasture sites in under plantations of native species in pure and mixed Monteverde, Costa Rica, seedlings of primary forest designs, at La Selva Biological Station in the Atlantic species were observed under the canopy of isolated humid lowlands of Costa Rica. In previous research trees, suggesting that seeds can germinate under the with other species in pure and mixed plantations, also shade of parental trees (Harvey and Haber, 1999). In at La Selva, tree regeneration was more successful other research in southern Costa Rica, germination of under the forest plantations than in abandoned seeds of tree species dispersed in pastures was similar pastures (Guariguata et al., 1995; Montagnini et al., to that in forests, although germination rates were 1999). In the present research the following hypoth- considerably lower in the pasture during the dry eses were tested. (1) Tree regeneration of native season (Holl, 1999). species is more abundant under the canopy of the The establishment of tree plantations in degraded plantations than in adjacent areas without trees areas may facilitate regeneration of native species (control); (2) tree regeneration is more diverse under that could not otherwise establish in open microsites the mixed plantations than under the pure-species or in competition by herbaceous species (Lugo, plantations; (3) tree regeneration is greater under the 1992). Several authors report on the role of tree plantation species whose litter decomposes more plantations as catalizers of natural succession in rapidly, because nutrients released from decomposi- tropical and subtropical sites (Parrotta, 1992, 1995; tion will favor seedling growth; (4) tree regeneration is Jussi et al., 1995; Parrotta et al., 1997; Keenan et al., less abundant in areas with higher litter accumulation, 1999; Parrotta, 1999; Otsamo, 2000). For example, since this may impede seed germination or seedling in southeast Asia, Jussi et al. (1995) reported on the growth; and (5) the majority of the tree species spontaneous and fast growth of indigenous tree regenerating in the understory of plantations are species under plantations of exotic trees. On the animal-dispersed. other hand, in north Queensland, Australia, a greater diversity of species was found in the understory of plantations of native species than in plantations 2. Methods of exotic species (Keenan et al., 1999). In Puerto Rico, in the understory of plantations of Albizia 2.1. Site description lebbek, 22 species of trees and shrubs were found, in comparison with just one species in control plots The plantations were at La Selva Biological Station, without trees (Parrotta, 1992). At La Selva Biolo- Costa Rica (108260N, 868590W, 50 m mean altitude, gical Station, Costa Rica, results of some studies also 24 8C mean annual temperature, 4000 mm mean suggest that tree plantations have a good potential for annual rainfall). Soils are Fluventic Dystropepts accelerating the processes leading to recovery of derived from volcanic alluvium. They are deep, well biodiversity in degraded soils (Guariguata et al., drained, stone-free, acid (pH in water <5.0), with 1995; Powers et al., 1997; Montagnini et al., 1999). low or medium organic matter prior to planting On the other hand, high establishment and main- (2.5±4.5%), cation exchange capacity 10±14 mol/kg, tenance costs are potential disadvantages of the use 10±15% base saturation, and moderately heavy texture of plantations for accelerating natural regeneration, (50±60% sand, 5±15% silt and 25±45% clay) (Sancho given the intensive management that is needed and Mata, 1987). Prior to plantation establishment, N.J. Carnevale, F. Montagnini / Forest Ecology and Management 163 (2002)217±227 219 soil conditions were too poor for cultivation of Plantation plots were in randomized blocks, with bananas or other commercial crops commonly grown four replicates of six treatments: pure plantation plots in the region (Bertsch, 1986; Sancho and Mata, 1987; of each species, mixed-species plots (with all four Montagnini, 1994). species), and plots that were not planted and left for The experimental plantations were part of a larger natural forest regeneration as control. Each plantation project to compare growth and nutrient dynamics in plot was 32 m  32 m. Initial planting distance was mixed and pure stands with indigenous tree species relatively tight (2 m  2 m) to speed canopy closure (Montagnini et al., 1995). The plantations were and obtain early impacts on soils. The three planta- established on land that was cleared in the mid- tions were thinned at 3 and 6 years after planting. With 1950s, grazed until 1981, then abandoned. In 1990, at thinning, the initial 2 m  2 m planting distance (2500 the time of clearing for the plantations, the site was trees/ha) was widened to 2 m  4 m and then to covered with shrubs and early successional trees, 4m 4 m. Within each mixed-tree plot, trees of the interspersed with patches of grass and ferns. The four species were planted with two species per row. experimental site was surrounded
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