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Logging Impacts on Forest Structure and Seedling Dynamics in a Prioria Copaifera (Fabaceae) Dominated Tropical Rain Forest (Talamanca, Costa Rica)

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Logging Impacts on Forest Structure and Seedling Dynamics in a Prioria Copaifera (Fabaceae) Dominated Tropical Rain Forest (Talamanca, Costa Rica) Logging impacts on forest structure and seedling dynamics in a Prioria copaifera (Fabaceae) dominated tropical rain forest (Talamanca, Costa Rica) Oscar J. Valverde-Barrantes1 & Oscar J. Rocha1 Department of Biological Sciences, Kent State University, PO Box 5190, Kent, Ohio, 44242-0001, USA; [email protected], [email protected] Received 10-IV-2013. Corrected 20-VIII-2013. Accepted 27-IX-2013. Abstract: The factors that determine the existence of tropical forests dominated by a single species (monodomi- nated forests) have been the subject of debate for a long time. It has been hypothesized that the low frequency of disturbances in monodominated forests and the tolerance to shade of the monodominant species are two important factors explaining the prolonged dominance of a single species. We determined the role of these two factors by examining the effects of logging activities on the floristic composition and seedling dynamics in a Prioria copaifera dominated forest in Southeastern Costa Rica. We determined the floristic composition for trees ≥2.5cm DBH and the associated recruitment, survival and mortality of tree canopy seedlings in two sites logged two (L-02) and 12 years (L-12) prior to sampling and an unlogged forest (ULF). Our results showed that L-02 stands had lower species richness (25 species) than the L-12 and ULF stands (49 and 46 species, respectively). As expected, we found significant logging effects on the canopy structure of the altered forests, particularly when comparing the L-02 and the ULF stands. Seedling density was higher in ULF (0.96 seedlings/ m2) than in the L-02and L-12 stands (0.322 and 0.466 seedlings/m2, respectively). However, seedling mortality was higher in the ULF stands (54%) than in the L-02 (26%) and L-12 (15%) stands. P. macroloba in L-02 was the only species with abundant regeneration under P. copaifera in L-02 stand, where it accounted for 35% of the seedlings. Despite the reduction in seedling abundance observed after logging, P. copaifera seems to maintain large seedling populations in these forests, suggesting that this species maintains its dominance after logging disturbances. Our findings challenge the hypothesis that the regeneration of monodominant species is not likely to occur under heavily disturbed canopy conditions. Rev. Biol. Trop. 62 (1): 347-357. Epub 2014 March 01. Key words: Carapa guianensis, floristic composition, forest structure, logging impact, tropical monodomi- nance, tree regeneration, Pentaclethra macroloba, Prioria copaifera. Monodominant forests (stands where more Proctor, & Villela, 1997), whereas in South East than 60% of the basal area is dominated by a Asian forests are dominated forms by members single species) are widespread throughout the of Dipterocarpaceae (Whitmore, 1998). Often tropics. Monodominant forests are scattered these forests thrive next to significantly more across much of the Congo basin in Africa as diverse communities without a clear reason for well as in the lowlands of Central and South their existence (Connell & Lowman, 1989), America (Whitmore, 1998) and Southeastern although several hypotheses had been proposed Asia (Richards, 1996; Yasuda, Matsumoto, & (Janos, 1985; Torti & Coley, 1999; Villela & Osada, 1999). For all Neotropical and Paleo- Proctor, 2002). tropical African forests, member of Caesalpini- Hart, Hart, & Murphy (1989) proposed oideae subfamily (Fabaceae) are the dominant that the main mechanism explaining this tree (Connell & Lowman, 1989; Nascimento, phenomenon is the conjunction of high seed Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 62 (1): 347-357, March 2014 347 production, poor seed dispersal systems, low Arteaga, 1991; Kursar, & Grauel, 2002). Com- light conditions and seedling adaptation to low mercially available stands have been extensive- luminosity. Following studies show that most ly described (Linares, 1987; Condit, Hubell, & monodominant tropical legumes (Caesalpini- Foster, 1993) and heavily exploited for wood oideae subfamily) in South America (Forget, during the last century (Lamb, 1953; Kursar, 1989; Nascimento, & Proctor, 1997; Henkel, & Grauel, 2002). However, there is still a lack Mayor, & Woolley, 2005) and Africa (Hart, of reliable information about the effects of log- 1995; Torti, Coley, & Kursar, 2001) as well ging activities on the structure and diversity as Dipterocarpaceae in South Asia (Herrera, of these forests (Grauel, 2004; Webb, 1997, Jordano, Guitián, & Traveset, 1998;Yasuda et 1999). The aim of this study was to understand al., 1999) follow a mast fruiting pattern, sup- how traditional logging practices affect forest porting the idea that high episodes of seed out- structure, diversity and seedling regeneration puts are a key factor in maintaining the stable in P. copaifera-dominated forests. dominance. Moreover, most of these species, Studies in undisturbed forests show that especially those adapted to flood conditions, monodominant species maintain a high seed- have large self-dispersed seeds that tend to cre- ling representation in the understory, which has ate large seedling banks, thus promoting mono- been interpreted as a requirement for continu- dominance (Lopez, 2001; Ter Steege, 1994; Ter ous dominance through time (Hart, 1995, Peh Steege, & Hammond, 2001). et al., 2011). Therefore, if logging activities are Several authors also point out that under- able to disrupt the P. copaifera stand dominance story light conditions in tropical monodominant we would expect: 1) an increase in other tree forests are shadier than that of adjacent mixed- species regeneration under P. copaifera canopy forests (Richards, 1996; Whitmore, 1998). In with respect to undisturbed conditions, 2) an fact, Torti et al. (2001) found that sunlight increase in P. copaifera seedling mortality at levels in the understory were on average lower logged stands in comparison with the unaltered and more homogeneous in a Gilbertiodendron areas (Guariguata, 2000) and 3) a decrease in dewevrei dominated stand than in adjacent seedling recruitments in logged forests. mixed-forests. These data are consistent with the deep crowns and canopies characteristic of MATERIALS AND METHoDS species forming monodominant stands (Rich- ards, 1996). Poor luminosity is considered an Study area: This study was conducted in a important ecological filter maintaining mono- 63ha forest near the Gandoca-Manzanillo Wild- dominance, as the regeneration of the domi- life Refuge, Talamanca, Costa Rica (9º37’83” nant species benefits from the environmental N - 82º38’52” W). The area is classified as a conditions created by the canopy trees (Frelich, tropical humid forest with 2 500-3 100mm of Calcote, Davis, & Pastor, 1993). Therefore, it precipitation annually and temperature ranging has been proposed that intense disturbances from 25 to 27ºC (Herrera, 1985). The forest could reverse monodominance as regeneration thrives on lowland alluvial flood plain soils of light demanding tree species would prolifer- identified as Hydric Psamments (Vázquez, ate under the new light conditions (Hart, 1995; 1979). The dominant floristic component for Peh, Lewis, & Lloyd, 2011). However, this this type of forest is Prioria copaifera Griseb hypothesis has not been formally tested. (Fabaceae-Caesalpinioidae) that represent as Prioria copaifera is one of the Neotropi- much as 60% of the basal area of the stand. cal caesalpinoids trees species known to form The canopy is also interspersed by Carapa monodominant stands. It accounts for 60-90% guianensis Aubl. (Meliaceae) and Pentaclethra of the basal area in the lowlands forest subjected macroloba (Willd.) Kuntze (Fabaceae-Mimo- to periodic flooding distributed from Nicaragua soidae). Simira maxonii (Standl.) Steyerm. to Colombia and Jamaica (Gonzáles, Gómez, & (Rubiaceae) is the dominant understory tree. 348 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 62 (1): 347-357, March 2014 The study site received two episodes of seedlings were recorded as recruits and seed- timber extraction, one in 1988 and the other in lings whose growth exceeded 150cm were 1997. Approximately 60% of all trees ≥60cm considered as saplings. in DBH were harvested in each stand. official Differences in stand structure were ana- records for the logging episode conducted in lyzed using the relative abundance of the 1997 reported an average of 7 trees/ha, for a most common species that were shared among total of 109 trees in the 15ha logged. There stands (P. copaifera, P. macroloba, C. guianen- were no official records available for the log- sis, S. maxonii and Musa textilis; Table 1). For ging operation conducted in 1988, but the the two most common species in all stands (P. wood extraction intensity was likely to be copaifera and P. macroloba), we compared the similar to that of 1997 considering the homo- proportion of individuals in each in diametric geneity of the forest and that extractions were classes for each stand. To capture different made following the same legislation (Quirós aspects of species diversity among stands, & Finegan, 1994). The stands were designated as logged-02 (L-02) and logged-12 (L-12), five indicators of diversity were calculated as according to the time from the extraction recommended by Chazdon, Colwell, Denslow, operation to the beginning of this study. An & Guariguata (1998): 1) species richness, 2) additional 33ha unlogged forest stand was also Simpson’s diversity index (D), 3) Shannon used in this study and it was designated as ULF. Diversity index (H’), 4) the proportion
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