Forest Ecology and Management 190 (2004) 359–372
Pure and mixed forest plantations with native species of the dry tropics of Costa Rica: a comparison of growth and productivity Daniel Piottoa, Edgar Vı´quezb, Florencia Montagninic,*, Markku Kanninend aGeotec Consultoria, Rua Estado de Israel 30, Sa˜o Paulo, SP, CEP 04022-000, Brazil bCentro Agrono´mico Tropical de Investigacio´n y Ensen˜anza (CATIE), 7170 Turrialba, Costa Rica cYale University, School of Forestry and Environmental Studies, 370 Prospect Street, New Haven, CT 06511, USA dCenter for International Forestry Research (CIFOR), 16680 Bogor Barat, Indonesia Received 7 April 2003; received in revised form 7 June 2003; accepted 6 November 2003
Abstract
In Costa Rica, most reforestation trials with native species were established in the tropical humid regions. In the dry tropics, research on the performance of native species in forest plantations is incipient and trials comparing pure and mixed designs are limited. This paper presents the results of two experimental plantations with native trees in pure and mixed plots in the dry tropics of Costa Rica. The growth and productivity of 13 native species in pure and mixed plantations was compared with Tectona grandis (L.f.) Lam., an exotic species broadly used in the region. In a plantation of relatively slower growing species, measurements taken at 68 months of age resulted in Samanea saman (Jacq.) Merril. and Dalbergia retusa Hemsl. demonstrating the best growth, followed by Astronium graveolens Jacq. and Swietenia macrophylla King. Measurements in a plantation of relatively faster growing species, at 68 months of age, showed that growth of Schizolobium parahyba (Vell.) Blake was greatest in the pure and mixed plots, followed by Terminalia oblonga (Ruiz & Pav.) Steud., Anarcadium excelsum (Bert. & Balb. ex Kunth) Skeels and Pseudosamanea guachapele (Kunth) Harms. The native species grew better in the mixed plots. The pure plots of T. grandis (L.f.) Lam. were the most productive, compared to all species and the mixture of species. Plantations of T. grandis (L.f.) Lam. seem to be well adapted to the region and are certainly a commercially interesting alternative. Nevertheless, mixed plantations with native species would contribute more to sustainable management, because while single-species plantations do not provide a great range of goods and services when compared to the natural forest, mixed plantations are likely to increase this range of benefits. # 2003 Elsevier B.V. All rights reserved.
Keywords: Costa Rica; Native species; Mixed plantations
1. Introduction cies from timber production to other land uses, and increasing protection of remaining forest areas. Con- Supply of high value timbers worldwide is becom- sequently, over the last few years, interest in establish- ing more limited due to overexploitation of these ing reforestation projects with native species in forest resources, conversion of forests containing these spe- plantations as a means to supplement existing tropical wood markets and as a way to detain the overexploita- * Corresponding author. Tel.: þ1-203-436-4221; tion of natural resources has increased. fax: þ1-203-432-3929. In Costa Rica, most reforestation trials with native E-mail address: [email protected] (F. Montagnini). species began in the 1980s, the majority of which were
0378-1127/$ – see front matter # 2003 Elsevier B.V. All rights reserved. doi:10.1016/j.foreco.2003.11.005 360 D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372 established in the tropical humid regions (Butterfield, compared with Tectona grandis (L.f.) Lam., an exotic 1990; Butterfield and Espinoza, 1995; Gonza´lez and species broadly used in the region. The hypotheses Fisher, 1994; Nichols, 1994; Montagnini et al., 1995; tested were: there is variation in growth and survival Arguedas and Chaverri, 1997; Haggar et al., 1998). In among species; the growth and survival of native spe- the dry tropics, where the forest cover has been cies is higher in mixtures that in pure forest plantations. reduced to a small fraction of the original area (Janzen, 1988), research on the performance of native species in forest plantations is incipient. 2. Materials and methods At the global level, single-species or ‘‘pure’’ plan- tations prevail in the tropics (Evans, 1999). Most forest 2.1. Site description projects use a small group of exotic species that are easy to manage through a variety of propagation The study plots were established in July 1995 in the methods, and are known for easy establishment, fast Peninsula of Nicoya, in the northern pacific region of initial growth and production of multiple products for Costa Rica (coordinates 108010North and established markets (Keenan et al., 1999). It should be 858420West). The land occupied by the study sites noted that many native tropical tree species are also was previously used for cattle ranching and is owned potentially valuable commercially, and that using such by Precious Woods of Costa Rica, a private company species, aside from satisfying economic objectives, that has reforested the majority of their properties with could be more acceptable ecologically and socially pure plantations of T. grandis and Bombacopsis qui- (Ball et al., 1995). nata (Jacq.) Dugand. The planting material was pro- For native species, mixed plantation systems seem duced in the company’s nursery, under a conventional to be the most appropriate for providing a broader black polyethylene nursery bag system. Seeds were range of options, such as production, protection, bio- collected from selected trees in the region. Weeding diversity conservation, and restoration of degraded was the main maintenance activity after field planting areas (Montagnini et al., 1995; Keenan et al., 1995; of trees, and a pruning of secondary apical shoots was Guariguata et al., 1995; Parrotta and Knowles, 1999). conducted in the first year. Mixed plantations can produce more biomass per unit The experimental area is located at an elevation of area because competition among individuals is 10–100 m above sea level and has an annual average reduced and the site is used integrally (Montagnini temperature of 27.5 8C. The mean annual rainfall is et al., 1995). The roots of different species may occupy 2350 mm per year with a 5-month dry period (less than different soil strata allowing more complete utilization 100 mm of precipitation per month). The trial replica- of soil and water resources (Lamb and Lawrence, tions were located in different slope and soil condi- 1993). More solar energy can be captured because tions (Table 1). In general Typic Haplustalf soils different species have different light requirements and prevail in the region, with medium to high fertility. crowns are broadly distributed in the vertical plane (Guariguata et al., 1995). 2.2. Methodology However, the success of the establishment of mixed forest plantations depends on plantation design and an Seven local species well known for slow growth and appropriate definition of the species to be used, taking seven species with relatively faster growth were into consideration ecological and silvicultural aspects planted in single-species plots and in a mixed-planting (Wormald, 1992). There exists very little information design. Table 2 includes the list of species, families, on the growth of tree species native to the dry tropics and range of natural distribution. Species choice was and information on experiences comparing pure and based on growth rate, timber value and availability of mixed forest plantations is limited. planting material. This paper presents the results of two experimental The growth and planting design combinations plantations with native trees in pure and mixed plots in resulted in four different species trials. Each trial the dry tropics of Costa Rica. Thegrowth and survival of has a complete randomized block design, with four 13 native species in pure and mixed plantations was replications, and species as treatments. D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372 361
Table 1 Soil characteristics at 0–20 cm depth in the experimental forest plantations at 68 months of age, in the Garza and Ostional Farms, Nicoya, Costa Rica
Replication
1234
Farm Garza Ostional Ostional Ostional Slope (%) 15 0 45 70 Rockinessa Low Low High Medium Organic matter (%) 5.4 8.1 7.4 4.3 pH (water) 6.4 5.9 7.3 6.6 Total nitrogen (%) 0.28 0.41 0.37 0.24 Phosphorus (mg/l) 3.1 3.5 1.9 1.6 Calcium (cmol/l) 24.51 19.90 35.89 28.27 Potassium (cmol/l) 0.48 0.22 0.20 0.40 Magnesium (cmol/l) 6.09 8.14 2.99 8.60
a Low (1–10%), medium (11–30%), high (>30%).
Every single-species plot trial (fast and slow growth In each plot, diameter at breast height (dbh) and group of species), has 25 trees per plot; while the total height were measured for each tree, including mixed planting design trials have 20 trees per species, border trees, at 16, 46 and 68 months. The averages of randomly distributed within the replication. T. grandis total height, dbh, basal area, volume index and survi- (L.f.) Lam. was only included in a single-species plot val were calculated for each plot. Variance analysis trial. was carried out for all variables for comparison Replications one and two in all of the four trials between species. For the volume index calculation a were established in a terrain with less than 15% slope, form factor of 0.5 was used, as suggested by New- in replication three the slope was 45% and in replica- bould (1967). For productivity variables (basal area tion four 70%. Planting distance was 3 m 3m. and volume index) it was necessary to carry out
Table 2 Names, family and natural distribution of the 14 species used in mixed and pure plantations in the Garza and Ostional Farms, Nicoya, Costa Ricaa
Species Family Natural distribution
Slow growth species P. parviflorum Benth. Fabaceae-Pap S. Mexico to Venezuela S. macrophylla King Meliaceae Mexico to Brazil and Bolivia C. odorata L. Meliaceae Mexico, S. America and Antillas P. pinnatum (Jacq.) Dugand ex Seem. Fabaceae-Pap S. Mexico to Venezuela A. graveolens Jacq. Anarcadiaceae Mexico to Paraguay D. retusa Hemsl. Fabaceae-Pap SE Mexico to Panama S. saman (Jacq.) Merril Fabaceae-Mim. Mexico to Brazil and Paraguay Fast growth species V. lundelli (Standl.) Killip Fabaceae-Pap S. Mexico to Brazil T. oblonga (Ruiz & Pav.) Steud. Combretaceae Honduras to the Amazon P. guachapele (Kunth) Harms Fabaceae-Mim. Mexico to Colombia and Venezuela A. excelsum (Bert. & Balb. ex Kunth) Skeels Anarcadiaceae Honduras to Ecuador and Guyanas P. pinnatum (Jacq.) Dugand ex Seem. Fabaceae-Pap S. Mexico to Venezuela S. apetala (Jacq.) Karst. Sterculiaceae Mexico to N. Brazil and Antillas S. parahyba (Vell.) Blake Fabaceae-Caes. S. Mexico to Brazil T. grandis (L.f.) Lam. Verbenaceae Asia, Malaysia and Indochina
a Source: Jime´nez and Poveda (1997) and Salazar (2000). 362 D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372 covariance tests, using survival as the covariable. This followed by Terminalia oblonga (Ruiz & Pav.) Steud., was done to achieve a better comparison between A. excelsum and P. guachapele, with no statistically species due to relatively high levels of mortality for significant differences (P < 0:05) between pure and most of the species, resulting in high standard devia- mixed plots. tions which diminished the power of the initially The growth in diameter of C. odorata was the applied variance analyses. greatest in the mixed plots, followed by S. saman, S. macrophylla, A. graveolens and D. retusa, with no statistically significant differences (P < 0:05) 3. Results between pure and mixed plots, except for C. odorata which exhibited the highest diametric growth in mixed Measurements in the slow growth plantation, at 68 plantation conditions, with highly significant differ- months of age, showed that Dalbergia retusa Hemsl and ences (P < 0:05) compared to the pure plots. Platy- Samanea saman (Jacq.) Merril. exhibited the highest miscium parviflorum Benth. and P. pinnatum showed rate of survival, followed by Astronium graveolens the slowest growth rates, with no significant differ- Jacq., Schizolobium parahyba King. and Cedrela odor- ences between pure and mixed plantations. S. saman ata L. demonstrated the lowest survival rate. No species and P. pinnatum only exhibited better growth rates in exhibited significant differences (P < 0:05) of survival pure plantation conditions. All other species exhibited between the pure and mixed plots. faster growth under mixed plantation conditions. Pseudosamanea guachapele (Kunth) Harms V. lundelli, P. pinnatum and S. apetala demonstrated showed the highest survival rate in the fast growth the lowest growth rates in height and diameter, with no plantation. All other species exhibited a high rate of significant differences between pure and mixed plan- mortality. Platymiscium pinnatum (Jacq.) Dugand ex tations. Only P. pinnatum showed greater growth Seem, Vatairea lundelli (Standl.) Killip and Sterculia under pure conditions. All other species of the fast apetala (Jacq.) Karst. demonstrated survival rates less growth plantation exhibited greater growth under than 20%. P. pinnatum exhibited better survival under mixed conditions. pure conditions. The other species had better survival Pure plantations of T. grandis were the most pro- rates under mixed conditions. Only Anarcadium excel- ductive, showing significant differences (P < 0:05) in sum (Bert. & Balb. ex Kunth) Skeels demonstrated basal area, compared to all species and the mixture of significant differences (P < 0:05) in survival between species in the two other plantations. The pure planta- pure and mixed plantations. Comparing slow and fast tion of S. parahyba was the only exception (Table 4). growth plantations, in general, species in the slow Productivity of native species was notably influ- growth plantation demonstrated better survival rates. enced by low survival rates, observed in both planta- Notably, the pure plots of T. grandis exhibited high tions, and by high genetic variability within the survival and growth rates, with low variation between species, as opposed to little genetic variation in T. replications (Table 3). grandis. In the slow growth plantation, measurements taken at S. saman, D. retusa and S. macrophylla exhibited 68 months of age resulted in S. saman and D. retusa the greatest productivity among the slow growth spe- demonstrating the best growth in height, followed by cies, when planted in pure stands, with a basal area of A. graveolens and Swietenia macrophylla,withno 7.93, 4.24 and 4.15 m2/ha, respectively (Table 4). statistically significant differences (P < 0:05) between These values were greater than those obtained in pure and mixed plantations. Notably, C. odorata exhib- the plots with a mixture of species (basal area of ited good growth in height in mixed plantations and 3.43 m2/ha, Table 4). The value obtained for S. saman showed the lowest growth in height in pure plantations, was significantly different (P < 0:05) to the rest of with statistically significant differences (P < 0:05) species and the mixture, and it represented a difference between the pure and mixed plots. of about 87% with respect to D. retusa and about Measurements in the fast growth plantation showed 131% with respect to the mix of species. Both Pla- that growth in height and diameter of S. parahyba tymiscium species and C. odorata had the poorest (Vell.) Blake, in the pure and mixed plots was greatest, productivity values (0.77, 0.39 and 0.44 m2/ha, D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372 363
Table 3 Growth and survival rates at 68 months of agea
Species Plot Survival (%) Total height (m) dbh (cm)
Slow growth species P. parviflorum Pure 50.0(7.4) bcd 2.96(0.34) de 4.03(0.62) def Mixed 35.0(21.9) cde 3.14(1.08) cde 4.39(1.51) def P. pinnatum Pure 16.0(8.7) de 3.73(1.37) bcde 4.36(1.56) def Mixed 21.3(16.4) de 3.38(1.14) cde 3.60(1.24) ef S. macrophylla Pure 62.0(19.2) abc 4.92(0.64) bcd 7.86(1.13) bcde Mixed 76.2(11.6) ab 5.10(0.50) bcd 7.92(0.55) bcde C. odorata Pure 11.0(6.8) e 1.66(1.29) e 3.03(2.71) f Mixed 12.5(3.3) e 5.44(1.10) bc 10.98(2.49) ab A. graveolens Pure 66.0(19.5) abc 4.71(0.67) bcd 6.48(1.04) cdef Mixed 75.0(7.1) ab 5.35(0.84) bcd 8.01(1.18) bcd D. retusa Pure 97.0(1.9) a 5.40(0.34) bcd 6.99(0.61) bcdef Mixed 88.7(5.6) a 5.87(0.37) b 7.51(0.37) bcde S. saman Pure 90.0(7.4) a 6.09(0.29) b 9.89(0.72) bc Mixed 85.0(11.8) ab 5.47(0.87) bc 9.25(1.96) bc T. grandis Pure 90.0(5.3) a 10.71(0.34) a 14.04(1.05) a Fast growth species V. lundelli Pure 13.0(6.8) ef 2.87(1.25) def 4.17(1.71) de Mixed 18.7(12.5) ef 3.81(1.82) def 4.49(1.86) de T. oblonga Pure 40.0(17.2) cdef 5.51(2.21) def 7.22(2.83) cde Mixed 65.0(20.1) abc 7.89(1.75) bcd 10.67(2.13) bcd P. guachapele Pure 90.0(2.6) ab 5.04(0.28) def 8.09(1.01) bcde Mixed 92.5(3.3) a 6.42(0.50) cde 9.79(0.43) bcd A. excelsum Pure 14.0(5.3) ef 4.11(1.40) def 6.44(2.23) de Mixed 57.5(20.7) bcd 6.90(1.07) cde 10.75(1.87) bcd P. pinnatum Pure 16.0(8.7) ef 3.73(1.37) def 4.36(1.56) de Mixed 12.5(12.5) ef 0.96(0.96) f 1.31(1.31) e S. apetala Pure 7.0(4.8) f 2.13(1.33) ef 5.37(3.32) de Mixed 19.9(15.4) ef 3.39(2.01) def 6.75(4.20) de S. parahyba Pure 28.0(19.1) def 12.15(4.40) ab 14.66(4.93) b Mixed 46.2(6.6) cde 16.38(1.43) a 26.06(2.75) a T. grandis Pure 90.0(5.3) ab 10.71(0.34) bc 14.04(1.05) bc
a For each plantation variables are compared among species and between pure and mixed plots. Means present significant differences when the standard error is followed by different letters in the same column (P < 0:05). respectively). C. odorata values were highly influ- 4. Discussion enced by survival. The basal area and volume values for all species in the slow growth plantation were The tree species of the dry tropics most studied in significantly inferior to the values obtained in pure forest plantations are the Meliaceae C. odorata and S. teak plots (15.33 and 84.69 m3/ha). macrophylla which have existed in plots established Similar results were obtained for the fast growing for genetic improvement in Costa Rica since 1991 species. In this case, S. parahyba and the mix of seven (Mese´n, 1996). In our study C. odorata exhibited the species showed the greatest productivity rates, basal highest mortality, primarily due to its susceptibility to area of 10.51 and 8.04 m2/ha, respectively. P. guacha- long periods of drought which occurred in the estab- pele and T. oblonga exhibited medium values (5.42 lishment phase (5–6 months), and repeated attacks of and 4.06 m2/ha). The other species exhibited very low the shoot borer Hypsipyla grandella (Zeller), both in levels of productivity (Table 4). Again, these values pure and mixed plantations. However, the few remain- were lower than the ones obtained in teak plantations. ing individuals demonstrated an extremely superior 364 D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372
Table 4 documented growth decrease in C. odorata after 20 a Basal Area and volume at 68 months of age years of age (Worbes, 1999). Species Basal area Volume index S. macrophylla seems to be more resistant to pest (m2/ha) (m3/ha) attack and drought, as evidenced by higher levels of Slow growth species survival than C. odorata, and by survival and growth P. parviflorum 0.77(0.27) de 1.51(0.55) c rate results comparable to those reported by Timyan P. pinnatum 0.39(0.16) e 1.05(0.40) c (1996) in Haiti under similar climatic conditions at 10 S. macrophylla 4.15(1.97) c 12.35(6.39) c years of age. In our study, the growth and survival of S. C. odorata 0.44(0.44) e 1.21(1.21) c macrophylla was also greater in mixed plantations. In A. graveolens 2.72(1.38) cde 8.86(5.41) c D. retusa 4.24(0.77) c 12.60(2.83) c other experiences in the region with pure and mixed S. saman 7.93(1.52) b 27.19(6.07) b plantations of S. macrophylla, at 79 months of age Mix of 7 species 3.43(0.70) cd 11.25(2.76) c diametric increment had not surpassed 1 cm per year T. grandis 15.33(1.38) a 84.69(9.34) a (Fonseca et al., 2000). However, growth data for this Fast growth species species in plantations in other regions does show V. lundelli 0.28(0.13) d 0.63(0.37) c promising scenarios (Hazlett and Montesinos, 1980; T. oblonga 4.06(2.20) cd 19.67(11.34) bc Webb et al., 1984; Geilfus and Serrano, 1991). P. guachapele 5.42(1.34) bcd 15.88(4.32) c A. excelsum 0.97(0.47) d 3.69(2.04) c S. saman achieved good growth in pure and mixed P. pinnatum 0.39(0.16) d 1.05(0.40) c conditions, with increments comparable to those S. apetala 0.59(0.35) d 1.55(0.95) c reported by Herrera and Lanuza (1995) in pure planta- S. parahyba 10.51(7.71) ab 106.96(84.90) a tions in the dry tropics of Nicaragua at 8 years of age Mix of seven species 8.04(2.21) bc 52.72(16.70) abc (height 1.2 m per year and dbh 1.8 cm per year) and by T. grandis 15.33(1.38) a 84.69(9.34) ab Fonseca et al. (2000) in mixed plantations in Liberia in a For each plantation, means present significant differences the northern Pacific region of Costa Rica at 79 months among treatments when the standard error is followed by different (height 0.9 m per year and dbh 1.8 cm per year). P < : letters in the same column ( 0 05). However, the productivity for S. saman plantations found here was less than 5 m3/ha per year, which is much less than the productivity of 25–35 m3/ha per diametric growth rate when in mixed plantations. This year reported by Webb et al. (1984), and 10–15 m3/ha is likely due to the reduction in frequency of H. per year reported by Roshetko (1995) for the same grandella attacks because of the presence of other species. species with different crown characteristics and S. parahyba exhibited excellent growth under because of the lower density of plants per hectare. mixed conditions, with increments in dbh greater than Timyan (1996) reports that under similar climatic those found for T. grandis. Nonetheless, S. parahyba conditions in Haiti the survival of C. odorata was exhibited high mortality rates in pure and mixed 50% and Butterfield (1993) reports survival rates of plantations, and a high variation of tree growth within 62% in mixed plantations in the humid tropics of the same plot. Survival here was much less than the Costa Rica, both much greater than the survival rates 81% reported by Rodr´ıguez (1990) for plantations in found in the dry tropics of Costa Rica. In the region of the same region of Costa Rica at 70 months. Annual San Carlos, Costa Rica, 7-year-old plantations with height increments for the pure plots were greater than low incidence of H. grandella attacks exhibited those reported by Rodr´ıguez (1990), and even higher greater growth when compared to plots that had high in mixed plots. The volumetric increments found for incidence of the pest (ACEFN, 1992). In young plan- this species of 20 m3/ha per year are comparable with tations (1 year old), Geilfus and Serrano (1991) found those reported by Webb et al. (1984), but represent a diametric increments greater than 3 cm per year. high variation of productivity between plots due to However, Hazlett and Montesinos (1980) report marked differences in survival between different repli- diametric increments of 0.64 cm per year for 32- cations. year-old plantations, showing a decrease in growth T. oblonga, P. guachapele and A. excelsum were the with increased age. This is likely due to a previously species with greatest survival rates in the fast growth D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372 365 plantation. T. oblonga exhibited greater levels of survi- Hazlett and Montesinos (1980) found diametric incre- val than those reported by Butterfield (1993) in mixed ments of 0.57 and 0.84 cm per year for A. graveolens plantations in the humid tropics of Costa Rica. The and D. retusa, respectively, in 31-year-old plantations. diametric increments found for T. oblonga are similar to The good form of these species and their high timber those reported by Butterfield (1993), and greater than value could compensate for the low annual increments those reported for 13-year-old plantations in La For- in volume, especially if they are grown in mixed tuna, San Carlos, Costa Rica (ACEFN, 1992) and for plantations. 38-year-old plantations in the Department of Suchite- S. apetala demonstrated high mortality and very pe´quez, Guatemala (Garc´ıa, 1983). In terms of growth, low levels of growth. In other countries, this species is these species have reached annual increments in dia- used in commercial plantations and demonstrates meter greater than 1.5 cm, and consequently figure as high survival rates and greater growth increments promising species for reforestation in the region, taking than those found in this study (Uruen˜a, 1993). For into account the early age of the plantations. V. lundelli, P. pinnatum and P. parviflorum there is no A. graveolens and D. retusa exhibited increments in literature on growth in plantations. The high mortality dbh greater than 1 cm per year, which is greater than rates in all the plots and very low growth levels found values found by Gutie´rrez (2000) for D. retusa in pure here, indicate silvicultural difficulties for plantation and mixed plantations in Guanacaste, Costa Rica. establishment of these species in this region.
Fig. 1. dbh (cm) and total height (m) for all species used in the slow growth plantation in pure and mixed plots. 366 D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372
Fig. 1. (Continued ) D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372 367
Fig. 1. (Continued ).
C. odorata, S. apetala, V. lundelli, P. pinnatum and for light was diminished by the variety of crown P. parviflorum had difficulties surviving in the first structures, resulting in more evenly distributed foli- phases of plantation development. At 16 months, high age in the vertical stratum, creating intermediate mortality rates were already documented. These spe- conditions of brightness (Guariguata et al., 1995). cies might demonstrate better results in situations with The interaction between plants of different families more intensive maintenance and management in the with legumes like D. retusa, P. pinnatum, P. parvi- first few years after plantation establishment. florum and S. saman in the slow growth plantation Plots located in the fourth replication showed the and P. guachapele, P. pinnatum, S. parahyba and lowest growth rates, which were significantly different V. lundelli in the fast growth plantation could have than those found in the other replications. This is contributed to a more balanced use of the site likely due to the steep slope of the land used for nutrients in mixed plots (Montagnini et al., 1994; the fourth replication, where the soils were considered Parrotta, 1999). fertile (Table 1). In addition, when we calculated basal area aver- The native species developed better in the mixed age, for example, we found that in the slow growth plots (Figs. 1 and 2). The trees in mixed plots had plantation the seven species in pure plots produced greatest diameter, likely due to reduced intra-specific 2.94 m2/ha, less than the mixture (3.43 m2/ha) and in competition (Montagnini et al., 1995). Competition the fast growth plantation the seven species in pure 368 D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372 plots produced 3.17 m2/ha, less than the mixture S. parahyba exhibited more than 50% of the total (8.04 m2/ha). In other words, if a farmer wants to basal area in the mixed plots (Fig. 3). Some of the produce timber wood of the seven species, mixing trees of this species have diameters greater than the species in the plantation will give better produc- 30 cm and could easily be processed and marketed tivity than if he/she plants seven pure blocks with after the first thinning. This would also open space each one of them. for increased growth and development in the other As total tree height is mainly influenced by site species of the mixed plot that present slower variables (Evans, 1992), significant differences in rhythms of growth. In the slow growth plantation, height were not expected between species in pure S. saman exhibited 30% of the total basal area of the and mixed plots. This was the case for all species mixed plots, followed by D. retusa, S. macrophylla studied, except C. odorata, whose growth in height and A. graveolens (Fig. 4). In that plantation the first was greatly affected by intense attacks of H. grandella thinning probably would not provide many if any in pure plots. commercial products. Mixed plantations could provide commercial In the mixed plots trees were distributed randomly, products after the first thinning, if the plantation limiting a deeper analysis of the interaction between is designed with species of different growth the species. Future research should focus on the rhythms. For example, in the fast growth plantation, identification of conditions and the group of species
Fig. 2. dbh (cm) and total height (m) for all species used in the fast growth plantation in pure and mixed plots. D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372 369
Fig. 2. (Continued ) 370 D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372
Fig. 2. (Continued ).
Fig. 3. Percentage of total basal area contributed by each species in Fig. 4. Percentage of total basal area contributed by each species in the mixed plots of the fast growth plantation, at 68 months of age. the mixed plots of the slow growth plantation, at 68 months of age. Averages present significant differences when they are followed by Averages present significant differences when they are followed by different letters (P < 0:05). different letters (P < 0:05). D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372 371 that result in the best growth rates. Mixed plots should Garc´ıa, G., 1983. Estudio silvicultural del volador (Terminalia be designed to allow for analysis of interactions oblonga) en el Departamento de Suchitepequez, Guatemala. Tesis de grado. Facultad de Agronom´ıa, Universidad de San between species (Kelty and Cameron, 1995). Carlos de Guatemala, 76 pp. Geilfus, F., Serrano, M., 1991. Crecimiento inicial de 14 especies maderables. Enda-Caribe. Santo Domingo, Republica Domin- 5. Conclusions icana, 54 pp. Gonza´lez, E., Fisher, R., 1994. Growth of native species planted on abandoned pasture land in Costa Rica. For. Ecol. Manage. 70, Although the plantations are still young and it may 159–167. be too soon to determine the behavior of the species Guariguata, M.R., Rheingans, R., Montagnini, F., 1995. Early studied, it is evident that best growth for these species woody invasion under tree plantations in Costa Rica: implica- was demonstrated in mixed systems. Management tions for forest restoration. Restor. Ecol. 3, 252–260. practices such as pruning and thinning could favor Gutie´rrez, M., 2000. Densidad inicial de siembra de cocobolo. Programa de restauracio´n y silvicultura. A´ rea de Conservacio´n the development of these species in mixed plantations Guanacaste. Reporte interno sin publicar. Liberia, Costa Rica, and provide revenues at earlier ages, when an appro- 5 pp. priate group of species is used. Haggar, J.P., Briscoe, C.B., Butterfield, R.P., 1998. Native species: Plantations of T. grandis seem to be well adapted to a resource for the diversification of forestry production in the the region and are certainly a commercially interesting lowland humid tropics. For. Ecol. Manage. 106, 195–203. Hazlett, D.L., Montesinos, J.L., 1980. El crecimiento de 27 alternative. On the other hand, mixed plantations with especies maderables en plantaciones de Lancetilla. ESNACI- native species would contribute more to sustainable FOR, Siguatepeque, Honduras. Art´ıculo Cient´ıfico 1, 9. management because they provide a greater range of Herrera, Z., Lanuza, B., 1995. Especies para reforestacio´nen goods and services than pure species plantations. Nicaragua. Servicio Forestal Nacional, MARENA. Managua, Nicaragua, 185 pp. Janzen, D.H., 1988. Tropical dry forests: the most endangered major tropical ecosystem. In: Wilson, E.O. (Ed.), Biodiversity. References National Academy Press, Washington, DC, pp. 130–137. Jime´nez, Q., Poveda, L.J., 1997. Lista actualizada de los a´rboles Asociacio´n Costarricense para el Estudio de Especies Forestales maderables de Costa Rica. Aportes al Desarrollo Sostenible. Nativas (ACEFN), 1992. Encuentro Regional sobre Especies Universidad Nacional, Heredia, Costa Rica, 36 pp. Forestales Nativas de la Zona Norte y Atla´ntica de Costa Rica Keenan, R.J., Lamb, D., Sexton, G., 1995. Experience with mixed (2, 1992, Sarapiqu´ı, CR). Memoria. Heredia, Costa Rica, 90 pp. species rainforest plantations in North Queensland. Common- Arguedas, M., Chaverri, P., 1997. Nuevo reporte en siete especies wealth For. Rev. 74 (4), 315–321. forestales nativas en Costa Rica. III Congreso Forestal Keenan, R.J., Lamb, D., Parrotta, J., Kikkawa, J., 1999. Ecosystem Nacional, del 27 al 29 de agosto de 1997. San Jose´, Costa Rica. management in tropical timber plantations: satisfying econom- Ball, J.B., Wormald, T.J., Russo, L., 1995. Experience with mixed ic, conservation, and social objectives. J. Sustain. For. 9 (1/2), and single species plantations. Commonwealth For. Rev. 74 (4), 117–134. 305–310. Kelty, M.J., Cameron, I.R., 1995. Plot designs for the analysis of Butterfield, R., 1990. Native species for reforestation and land species interactions in mixed stands. Commonwealth For. Rev. restoration: a case study from Costa Rica. In: Proceedings of the 74 (4), 322–332. 14th IUFRO World Congress, vol. 2. Montreal, Canada, pp. 3–14. Lamb, D., Lawrence, P., 1993. Mixed species plantations using Butterfield, R., 1993. Tropical timber species growth in the Atlantic high value rainforest trees in Australia. In: Lieth, H., Lohmann, lowlands of Costa Rica and wood variation of two native M. (Eds.), Restoration of Tropical Forest Ecosystems. Kluwer species. Thesis Ph.D. North Carolina State University, Raleigh, Academic Publishers, Holanda, pp. 101–108. NC, 76 pp. Mese´n, F., 1996. Mejoramiento de especies forestales nativas de Butterfield, R., Espinoza, M., 1995. Screening trial of 14 tropical Costa Rica. En: Asociacio´n Costarricense para el Estudio de hardwoods with an emphasis on species native to Costa Rica: Especies Forestales Nativas (ACEFN). Foro ‘‘Especies fourth year results. New For. 9, 135–145. forestales nativas, una opcio´n para la reforestacio´nsusten- Evans, J., 1992. Plantation Forestry in the Tropics, 2nd ed. table en Costa Rica’’. Memoria. Moravia, Costa Rica, pp. Clarendon Press, Oxford, 403 pp. 17–34. Evans, J., 1999. Planted forests of the wet and dry tropics: their Montagnini, F., Fanzeres, A., da Vinha, S., 1994. Studies on variety, nature, and significance. New For. 17, 25–36. restoration ecology in the Atlantic forest region of Bahia, Fonseca, W., Chinchilla, O., Gutie´rrez, M., 2000. Cen´ızaro Brazil. Interciencia 19 (6), 323–330. (Samanea saman). Estacio´n Experimental Forestal Horizontes. Montagnini, F., Gonza´lez, E., Rheingans, R., Porras, C., 1995. Reporte interno sin publicar. Liberia, Costa Rica, 10 pp. Mixed and pure forest plantations in the humid neotropics: a 372 D. Piotto et al. / Forest Ecology and Management 190 (2004) 359–372
comparison of early growth, pest damage and establishment Production: A Field Manual. Winrock International. Morrilton, costs. Commonwealth For. Rev. 74 (4), 306–314. Arkansas, USA, 1996, 125 pp. Newbould, P.J., 1967. Methods for Estimating the Primary Salazar, R., 2000. Manejo de semillas de 100 especies forestales de Production of Forest. IBP Handbook 2. Blackwell Scientific Ame´rica Latina. Proyecto de Semillas Forestales/Danida Forest Publications, Oxford, UK, 62 pp. Seed Centre. CATIE, Turrialba, Costa Rica (Serie te´cnica. Nichols, D., 1994. Terminalia amazonia (Gmel.) Exell.: develop- Manual te´cnico no. 41), 204 pp. ment of native species for reforestation and agroforestry. Timyan, J., 1996. Bwa yo: Important trees of Haiti. In: Proceedings Commonwealth For. Rev. 73 (1), 9–13. of the South-east Consortium for International Development, Parrotta, J.A., Knowles, O.H., 1999. Restoration of tropical moist Washington, DC, 420 pp. forests on bauxite-mined lands in the Brazilian Amazon. Uruen˜a, H., 1993. Evaluacio´n de un ensayo de progenie/pro- Restor. Ecol. 7, 103–116. cedencia de Sterculia apetala de cinco an˜os de edad. Informe Parrotta, J.A., 1999. Productivity, nutrient cycling, and succession de Investigacio´n no. 22. Monterrey Forestal, Cartagena, in single- and mixed-species plantations of Casuarina equise- Colombia, 3 pp. tifolia, Eucalyptus robusta, and Leucaena leucocephala in Webb, D.B., Wood, P.J., Smith, J.P., Sian Henman, G., 1984. A Puerto Rico. For. Ecol. Manage. 124, 45–77. guide to species selection for tropical and sub-tropical Rodr´ıguez, E., 1990. Comportamiento inicial de cinco especies plantations. Tropical Forestry Papers no. 15. Commonwealth forestales para la produccio´n de len˜a en la alta pen´ınsula de Forestry Institute, Oxford, UK, 256 pp. Nicoya, Costa Rica. Informe Te´cnico Interno CATIE 04/ Worbes, M., 1999. Annual growth rings, rainfall-dependent growth Exp.092. CATIE, Turrialba, Costa Rica, 13 pp. and long-term growth patterns of tropical trees from the Caparo Roshetko, J.M., 1995. Albizia saman: pasture improvement, shade, Forest Reserve in Venezuela. J. Ecol. 87, 391–403. timber and more. NFT-Highlights no 95-02. In: Roshetko, J.M., Wormald, T.J., 1992. Mixed and Pure Forest Plantations in the Gutteridge, R.C. (Eds.), Nitrogen Fixing Trees for Fodder Tropics and Subtropics, no. 103. FAO, 166 pp.