Litterfall Dynamics Under Different Tropical Forest Restoration Strategies in Costa Rica

In degraded tropical pastures, active restoration strategies have the potential to facilitate forest regrowth at rates that are faster than natural recovery, enhancing litterfall, and nutrient inputs to the forest floor. We evaluated litterfall and nutrient dynamics under four treatments: plantation (entire area planted), tree islands (planting in six patches of three sizes), control (same age natural regeneration), and young secondary forest (7–9‐yr‐old natural regeneration). Treatments were established in plots of 50 × 50 m at six replicate sites in southern Costa Rica and the annual litterfall production was measured 5 yr after treatment establishment. Planted species included two native timber‐producing hardwoods (Terminalia amazonia and Vochysia guatemalensis) interplanted with two N‐fixing species (Inga edulis and Erythrina poeppigiana). Litter production was highest in secondary forests (7.3 Mg/ha/yr) and plantations (6.3), intermediate in islands (3.5), and lowest in controls (1.4). Secondary forests had higher input of all nutrients except N when compared with the plantation plots. Inga contributed 70 percent of leaffall in the plantations, demonstrating the influence that one species can have on litter quantity and quality. Although tree islands had lower litterfall rates, they were similar to plantations in inputs of Mg, K, P, Zn, and Mn. Tree islands increased litter production and nutrient inputs more quickly than natural regeneration. In addition to being less resource intensive than conventional plantations, this planting design promotes a more rapid increase in litter diversity and more spatial heterogeneity, which can accelerate the rate of nutrient cycling and facilitate forest recovery.     

Avian Habitat Preference in Tropical Forest Restoration in Southern Costa Rica

An important question for tropical forest restoration is whether degraded lands can be actively managed to attract birds. We censused birds and measured vegetation structure at 27 stations in young (6–9‐yr old) actively and passively restored pasture and old growth forest at Las Cruces Biological Station in southern Costa Rica. During 481 10‐min point counts, we detected a high diversity—186 species—of birds using the restoration area. Surprisingly, species richness and detection frequency did not differ among habitats, and proportional similarity of bird assemblages to old growth forest did not differ between restoration treatments. Bird detection frequency was instead explained by exotic grass cover and understory stem density—vegetation structures that were not strongly impacted by active restoration. The similarity of bird assemblages in actively and passively restored forest may be attributed to differential habitat preferences within and among feeding guilds, low structural contrast between treatments, or the effect of nucleation from actively restored plots into passively restored areas. Rapid recovery of vegetation in this recently restored site is likely due to its proximity to old growth forest and the lack of barriers to effective seed dispersal. Previous restoration studies in highly binary environments (i.e., open pasture vs. tree plantation) have found strong differences in bird abundance and richness. Our data contradict this trend, and suggest that tropical restoration ecologists should carefully consider: (1) when the benefits of active restoration outweigh the cost of implementation; and (2) which avian guilds should be used to measure restoration success given differential responses to habitat structure.     

Tropical Montane Forest Restoration in Costa Rica: Overcoming Barriers to Dispersal and Establishment

Tropical forests are being cleared at an alarming rate although our understanding of their ecology is limited. It is therefore essential to design restoration experiments that both further our basic knowledge of tropical ecology and inform management strategies to facilitate recovery of these ecosystems. Here we synthesize the results of research on tropical montane forest recovery in abandoned pasture in Costa Rica to address the following questions: (1) What factors limit tropical forest recovery in abandoned pasture? and (2) How can we use this information to design strategies to facilitate ecosystem recovery? Our results indicate that a number of factors impede tropical forest recovery in abandoned pasture land. The most important barriers are lack of dispersal of forest seeds and seedling competition with pasture grasses. High seed predation, low seed germination, lack of nutrients, high light intensity, and rabbit herbivory also affect recovery. Successful strategies to facilitate recovery in abandoned pastures must simultaneously overcome numerous obstacles. Our research shows that establishment of woody species, either native tree seedlings or early‐successional shrubs, can be successful in facilitating recovery, by enhancing seed dispersal and shading out pasture grasses. On the contrary, bird perching structures alone are not an effective strategy, because they only serve to enhance seed dispersal but do not reduce grass cover. Remnant pasture trees can serve as foci of natural recovery and may enhance growth of planted seedlings. Our results highlight the importance of: (1) understanding the basic biology of an ecosystem to design effective restoration strategies; (2) comparing results across a range of sites to determine which restoration strategies are most generally useful; and (3) considering where best to allocate efforts in large‐scale restoration projects.     

Patch Size and Tree Species Influence the Number and Duration of Bird Visits in Forest Restoration Plots in Southern Costa Rica

Efforts to restore tropical forest in abandoned pasture are likely to be more successful when bird visitation is promoted because birds disperse seeds and eat herbivorous arthropods that damage leaves. Thus, it is critical to understand bird behavior in relation to different restoration strategies. We measured the likelihood of visitation, number of visits, and duration of visits for all birds and for Cherrie's Tanager ( Ramphocelus costaricensis ), a common seed disperser, in five sizes of forest restoration patches planted with four tree species in southern Costa Rica. We predicted that the largest patches, and the tree species with the greatest canopy cover, would be visited most frequently and have the longest visits because we assumed that these patch types had the greatest food resources and the lowest predation risk. We found that birds were more likely to visit large patches and the tree species with the highest canopy cover ( Inga edulis ). Birds visited Inga trees more often and stayed in Inga and Erythrina poeppigiana trees for longer periods of time than in other tree species. We found similar results for Cherrie's Tanagers. Thus, we identified two factors, tree species and patch size, which may be manipulated in restoration projects to increase bird visitation.     

Contrasting Cloud Forest Restoration Potential Between Plantations of Different Exotic Tree Species

The role of exotic tree plantations for biodiversity conservation is contested. Such plantations nevertheless offer various ecosystem service benefits, which include carbon storage and facilitation of indigenous tree species regeneration. To assess forest restoration potential in tropical exotic tree plantations, we assessed native cloud forest tree regeneration in 166 plots in ca. 50‐year‐old plantations of five timber species that are widely used in tropical plantations (Pinus patula, Eucalyptus saligna, Cupressus lusitanica, Grevillea robusta and Acacia mearnsii). Differences in species abundance, diversity and composition were compared among plantations, and between plantations and disturbed and undisturbed indigenous Afromontane cloud forest (southeast Kenya) relicts after controlling for environmental variation between plots (i.e. altitude, distance to indigenous forest, soil depth, slope, aspect) and for environmental and stand structural variation (i.e. dominant tree height and basal area). Regenerating trees were mostly early‐successional species. Indigenous tree species regeneration was significantly higher in Grevillea plantations, where the seedling community also included late‐successional tree species. Regeneration under Eucalyptus was particularly poor. Acacia had a strong invasive nature, reducing its potential role and usefulness in indigenous forest restoration. Our study underlined that exotic tree plantations have differential effects on native tree species regeneration, with high potential for Grevillea plantations and low potential for invasive exotic species.     

Tree Regeneration and Understory Woody Plants Show Diverse Responses to Forest–Pasture Edges in Costa Rica

ABSTRACT Edge effects along tropical forest–pasture margins are thought to cause a shift toward early successional characteristics of the understory forest vegetation. We tested this idea by sampling vegetation at five forest sites in northeast Costa Rica each of which had edges that were established over 20 yr earlier. Four of these sites had been selectively logged. We sampled woody plants >0.2 and ≤1.3 m height in 54 m² within 0.2 ha plots at edges (N=14), and at 150 m (N=11) and 300 m from edges (N=9). Composition and diversity did not vary with edge distance. Abundance of tree regeneration, mainly of canopy and emergent species, increased at edges. Abundance of lianas and slow‐growing tree species did not differ significantly across the sampling locations. Weighted mean wood density varied little, with a reduction at edges for canopy species. Palms were less abundant at edges, but not less species rich. At edges, these plant assemblages maintain many of the characteristics of forest interior vegetation, though the changes observed may indicate ongoing functional change. Degradation of forest–pasture edges is not a universal feature of tropical forest fragmentation, and forests with high rates of natural turnover might have a high capacity to maintain themselves within forest edges alongside pasture.     

Allometric Models for Tree Volume and Total Aboveground Biomass in a Tropical Humid Forest in Costa Rica1

Allometric equations for the estimation of tree volume and aboveground biomass in a tropical humid forest were developed based on direct measurements of 19 individuals of seven tree species in Northern Costa Rica. The volume and the biomass of the stems represented about two‐thirds of the total volume and total aboveground biomass, respectively. The average stem volume varied between 4 and 11 Mg/tree and the average total aboveground biomass ranged from 4 to 10 mg/tree. The mean specific gravity of the sampled trees was 0.62 ± 0.06 (g/cm³). The average biomass expansion factor was 1.6 ± 0.2. The best‐fit equations for stem and total volume were of logarithmic form, with diameter at breast height (R²= 0.66 ? 0.81) as an independent variable. The best‐fit equations for total aboveground biomass that were based on combinations of diameter at breast height, and total and commercial height as independent variables had R² values between 0.77 and 0.87. Models recommended for estimating total aboveground biomass are based on diameter at breast height, because the simplicity of these models is advantageous. This variable is easy to measure accurately in the field and is the most common variable recorded in forest inventories. Two widely used models in literature tend to underestimate aboveground biomass in large trees. In contrast, the models developed in this study accurately estimate the total aboveground biomass in these trees.     

Sustainable ecotourism in Costa Rica: the Monteverde Cloud Forest Preserve

The paper discusses the sustainability of the Monteverde Cloud Forest Preserve in the context of Costa Rican ecotourism. While the history of the Preserve is somewhat unique, the analysis of visitation, financial, ecological and economic factors provides a convincing case that tourism at the Preserve is sustainable. The experience of the Preserve is also put in the context of Costa Rican ecotourism, particularly to the national parks. The paper concludes that the Preserve has played a very important role in the development of Costa Rica as an ecotourism destination. Nonetheless, the failure of experience at the Preserve to inform recent changes in national park pricing policy reveal that Costa Rica has yet to fully capitalize on the experience gained and lessons learned at the Preserve.     

Rain Forest Structure at Forest-Pasture Edges in Northeastern Costa Rica

Land-use change in the Sarapiquí region of Costa Rica has resulted in a fragmented forest landscape with abrupt edges between forest and pasture. Forest responses to edge effects vary widely and can significantly affect ecosystem integrity. Our objective was to examine forest structure at 20+ yr old forest-pasture edges in Sarapiquí. Three transects with 0.095-ha plots at seven distances from forest edges were established in each of six forest patches. Stem density, basal area, and aboveground biomass in trees and palms ≥ 10-cm diameter at breast height were measured in all plots. In addition, hemispherical photographs were taken to determine leaf area index, understory light availability, and percent canopy openness. Linear mixed-effects models showed significantly higher tree stem density at forest edges, relative to interiors, a pattern reflected by increased stem density, basal area, and aboveground biomass in small diameter trees (≤ 20 cm) growing near edges. No differences in total tree basal area, aboveground biomass, or hemispherical photograph-derived parameters were detected across the forest edge to interior gradient. The recruitment of small diameter trees following edge creation has contributed to the development of dense vegetation at the forest edge and has aided in the maintenance of similar tree basal area and aboveground biomass between edge and interior environments. These data reflect on the robustness of forest edges in Sarapiquí, a characteristic that will likely minimize future detrimental edge effects and promote a number of high-value environmental services in these forests.     

Tropical Rain Forest Structure,Tree Growth and Dynamics along a 2700-m Elevational Transect in Costa Rica

Rapid biological changes are expected to occur on tropical elevational gradients as species migrate upslope or go extinct in the face of global warming. We established a series of 9 1-ha plots in old-growth tropical rainforest in Costa Rica along a 2700 m relief elevational gradient to carry out long-term monitoring of tropical rain forest structure, dynamics and tree growth. Within each plot we mapped, identified, and annually measured diameter for all woody individuals with stem diameters >10 cm for periods of 3-10 years. Wood species diversity peaked at 400-600 m and decreased substantially at higher elevations. Basal area and stem number varied by less than two-fold, with the exception of the 2800 m cloud forest summit, where basal area and stem number were approximately double that of lower sites. Canopy gaps extending to the forest floor accounted for <3% of microsites at all elevations. Height of highest crowns and the coefficient of variation of crown height both decreased with increasing elevation. Rates of turnover of individuals and of stand basal area decreased with elevation, but rates of diameter growth and stand basal area showed no simple relation to elevation. We discuss issues encountered in the design and implementation of this network of plots, including biased sampling, missing key meteorological and biomass data, and strategies for improving species-level research. Taking full advantage of the major research potential of tropical forest elevational transects will require sustaining and extending ground based studies, incorporation of new remotely-sensed data and data-acquisition platforms, and new funding models to support decadal research on these rapidly-changing systems.     

A Model to Assess Restoration of Abandoned Pasture in Costa Rica Based on Soil Hydrologic Features and Forest Structure

An ecological functional assessment (EFA) was used on 10 southwest Costa Rica sites representing a chronosequence of formerly pastured lands to undisturbed tropical wet forest. Ecological functional assessment is a tool designed to assess wetland functions in the United States that was adapted to upland forests. Models to indicate characteristic soil hydrologic features and soil structure and aboveground spatial structure of habitat were used to examine the degree to which selected sites within the chronosequence approach the undisturbed condition of the natural forest. An index of the functional model for the maintenance of characteristic soil hydrologic features (such as infiltration, bulk density, etc.) showed that the 20‐year‐old secondary forest was at approximately 60% of the condition of the undisturbed sites, whereas active pasture was evaluated at approximately 20% of the reference undisturbed forest; 4‐ and 10‐year‐old sites were intermediate. The spatial structure of habitat model showed that 20‐year‐old secondary forest was approximately 50% of reference forest, whereas active pasture was approximately 10% of the condition of undisturbed forest; 4‐year‐old sites were evaluated at approximately 20% and 10‐year‐old sites at approximately 60% of the reference state. Overall the functional assessment process indicated that degraded tropical wet forest sites have recovered almost 60% of their functional qualities 10 years following pasture abandonment. These results indicate that EFA can be a useful technique for monitoring restoration programs in the tropics.     

Nurse Trees as a Forest Restoration Tool for Mixed Plantations: Effects on Competing Vegetation and Performance in Target Tree Species

Multi‐species mixed plantations can be designed to meet social, economic, and environmental objectives during forest restoration. This paper reports results from an experiment in southern Sweden concerning the influence of three different fast growing nurse tree species on the cover of herbaceous vegetation and on the performance of several target tree species. After 10 years, the nurse trees had reduced the competing herbaceous vegetation but the effect was weak and it may take more than a decade to achieve effective vegetation control. The nurse tree species Betula pendula and Larix x eurolepis did improve stem form in some target tree species, but had a minor effect on survival and growth. The open conditions before crown closure of nurse trees strongly influence seedling performance and so delayed planting of target tree species may provide a means to avoid those conditions. Survival and growth differed greatly among the tree species. Besides the two nurse tree species mentioned above, high survival was found in Picea abies and Quercus robur and intermediate survival in Fagus sylvatica, Tilia cordata, and in the N‐fixing nurse tree Alnus glutinosa. Survival was low in the target tree species Fraxinus excelsior L. and Prunus avium. For restoration practitioners, our results illustrate the potential of using nurse trees for rapidly building a new forest structure and simultaneously increase productivity, which might be a cost‐effective strategy for forest restoration.     

The Landscape Ecology of Tropical Secondary Forest in Montane Costa Rica

Multinomial logistic models of land use/land cover in montane Costa Rica and landscape pattern analysis showed that relative to agriculture, secondary forest occurred closer to old-growth forest, further from roads, in forest reserves, and at higher elevations. Collinearity between explanatory variables yielded simple multivariate models; proportion of surrounding old growth predicted secondary forest most accurately. An old-growth matrix [mean patch size (MPS) 24.5 ha], located mainly within protected areas, dominated elevations greater than 2500 m. A matrix of agriculture (MPS 23.5 ha), with smaller patches (approximately 9 ha) of secondary forest and old growth, dominated elevations from 1500 to 2500 m. Combining secondary forest with old growth decreased forest patch number and increased MPS from 7.3 to 37.1 ha. I concluded that: (a) secondary forest pattern is nonrandom, so ancillary data will aid its mapping with satellite imagery. The variables elevation, agriculture distance, road distance, and population density distinguished secondary forest from old growth with 74% accuracy; (b) socioeconomic and biological forces probably interact to create these secondary forest patterns; and (c) the strong association between secondary forest and old growth supports the concept that tropical forest recovery depends on the landscape structure of remnant forest. Received 16 February 1999; accepted 20 August 1999.     

Planting Seedlings in Tree Islands Versus Plantations as a Large‐Scale Tropical Forest Restoration Strategy

Planting tree seedlings in small patches (islands) has been proposed as a method to facilitate forest recovery that is less expensive than planting large areas and better simulates the nucleation process of recovery. We planted seedlings of four tree species at 12 formerly agricultural sites in southern Costa Rica in two designs: plantation (entire 50 × 50 m area planted) and island (six patches of three sizes). We monitored seedling survival, height, and canopy area over 3 years. To elucidate mechanisms influencing survival and growth, we measured soil and foliar nutrients, soil compaction, and photosynthesis. Survival of all species was similar in the two planting designs. Seedling height and canopy area were greater in plantations than islands at most sites, and more seedlings in islands decreased in height due to damage incurred during plot maintenance. Survival, height, and canopy area were both site‐ and species‐specific with the two N‐fixing species (Inga edulis and Erythrina poeppigiana) greater than the other species (Terminalia amazonia and Vochysia guatemalensis). Foliar N was higher in Terminalia and Vochysia in sites where Inga growth was greater. Soil nutrients, however, explained a small amount of the large differences in growth across sites. Leaf mass per area was higher in islands, and P use efficiency was higher in plantations. Our results show advantages (good seedling survival, cheaper) and disadvantages (more seedling damage, slightly lower growth) to the island planting design. Our study highlights the importance of replicating restoration strategies at several sites to make widespread management recommendations.     

Microhabitat distribution of protostelids in a Tropical Wet Forest in Costa Rica

A microhabitat study of protostelids was carried out in a Tropical Wet Forest at the La Selva Biological Station in Costa Rica. Nine species were recorded from sterile wheat straws placed out and then re-collected over a period of six weeks from two different litter microhabitats in an area of primary forest. All nine species were present on straws placed in the aerial litter microhabitat, but only six species were present on straws placed in the forest floor litter microhabitat. Total colonies, percent of straws colonized, and mean number of species per straw increased significantly over time. One species (Schizoplasmodiopsis pseudoendospora) typical of temperate litter was the overwhelming dominant on the forest floor litter, while Echinostelium bisporum, a species rare in temperate litter microhabitats, was the single most abundant species in the aerial litter microhabitat. Both of these species had significantly increased frequencies over time. Two species abundant in temperate aerial litter microhabitats and one species abundant in temperate forest floor litter were rare at La Selva. Our data conform to those obtained in an earlier study carried out in tropical forests in the mountains of Puerto Rico and provide additional support towards developing a model of microhabitat distribution of protostelids in terrestrial ecosystems.     

Phenology, Lignotubers, and Water Relations of Cochlospermum vitifolium, a Pioneer Tropical Dry Forest Tree in Costa Rica

We examined structural and physiological traits relevant to the phenology of the tropical dry forest (TDF) pioneer tree Cochlospermum vitifolium . Despite marked seasonality in rainfall, meristem activity occurred throughout the year. Leaves were produced almost continuously during the rainy season, while leaf shedding started early during drought, before changes in soil water content were observed. Phenological activity under drought included flowering and fruiting of leafless trees; bud break and shoot extension took place before the end of the dry season. Low wood density of C. vitifolium stems (0.17 g/cm³) and lignotubers (0.14 g/cm³) provided water and starch storage needed to support phenological events such as branch extension, leaf flushing, and reproduction during the dry season, and probably also contributed to survival following mechanical damage and fire, typical of early TDF successional stages. Lignotuber water and starch contents showed substantial seasonal variation, declining from the beginning of the dry season to their lowest levels at the time of reproduction and dry-season flushing. Stems progressively replaced lignotubers as main storage organs as tree size increased. Evidence for a role of water stores in buffering daily water deficits was weak. Leaf water potentials remained above −1.2 MPa and stomatal conductance below 350 mmol/m²/s, suggesting that gas exchange during the rainy season was limited to prevent xylem cavitation. Leaf shedding occurred when early-morning and mid-day Ψ_L converged at the rainy–dry season transition, without changes in lignotuber or soil water content, suggesting that leaves of C. vitifolium are closely tuned to atmospheric drought.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp .     

Effects of Tree Guards and Weed Mats on the Establishment of Native Tree Seedlings: Implications for Forest Restoration in Hong Kong, China

The effects of tree guards and weed mats on establishment and growth of native tree seedlings, Thick‐leaved oak (Cyclobalanopsis edithiae (Skan) Schott., Fagaceae), planted in an exposed hillside grassland in Hong Kong, were investigated. The natural regeneration of C. edithiae is poor due to a lack of seed dispersal agents and high seed predation, and therefore, this species is often targeted for forest restoration. The experiment lasted for 3.5 years during which the height, basal diameter, and crown diameter of individual seedlings were measured and survivorship recorded. The use of weed mats alone did not have a significant effect, but a combination of tree guards and weed mats led to a significant improvement in establishment, survivorship, and growth of the seedlings during the experimental period. Initially, the guards promoted rapid height growth of the seedlings, although lateral growth and secondary stem thickening were compromised. After the seedlings grew over the tree guards, the basal diameter and crown diameter increased at a notably faster rate. The combined effect of the tree guard and weed mat on the seedling growth pattern was found to be beneficial and contributed to the high survivorship of the seedlings. Comparing the survivorship data and the costs of various treatments, the use of tree guards in combination with the weed mats was demonstrated to be more cost‐effective than planting the seedlings without tree guards or weed mats. The potential for applying the technique in afforestation programs with native tree species for forest restoration in Hong Kong and other tropical regions is discussed.