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.     

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.     

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.     

The effect of climate and soil conditions on tree species turnover in a Tropical Montane Cloud Forest in Costa Rica

On a global level, Tropical Montane Cloud Forests constitute important centers of vascular plant diversity. Tree species turnover along environmental gradients plays an important role in larger scale diversity patterns in tropical mountains. This study aims to estimate the magnitude of beta diversity across the Tilardn mountain range in North-Western Costa Rica, and to elucidate the impact of climate and soil conditions on tree species turnover at a local scale. Seven climate stations measuring rainfall, horizontal precipitation (clouds and wind-driven rain) and temperatures were installed along a 2.5km transect ranging from 1200 m.a.s.l. on the Atlantic to 1200 m.a.s.l. on the Pacific slope. The ridge top climate station was located at 1500 m.a.s.l. Climate data were recorded from March through December 2003. Additionally, seven 0.05 ha plots were established. On all plots soil moisture was monitored for one year, furthermore soil type and soil chemistry were assessed. Woody plants with a diameter at breast height (dbh) > or = 5 cm were identified to species. Species' distributions were explored by feeding pairwise Serensen measures between plots into a Principal Component Analysis. Relationships between floristic similarity and environmental variables were analyzed using Mantel tests. Pronounced gradients in horizontal precipitation, temperatures and soil conditions were found across the transect. In total, 483 woody plants were identified, belonging to 132 species. Environmental gradients were paralleled by tree species turnover; the plots could be divided in three distinctive floristic units which reflected different topographic positions on the transect (lower slopes, mid slopes and ridge). Most notably there was a complete species turnover between the ridge and the lower Pacific slope. Floristic similarity was negatively correlated with differences in elevation, horizontal precipitation, temperatures and soil conditions between plots. It is suggested that beta-diversity in the study area is largely driven by species with narrow spatial ranges, due to the interactions between topography, climate and soil formation processes, especially around the wind-exposed and cloud covered ridge area. The findings emphasize the extraordinary conservation value of tropical montane cloud forests in environmentally heterogeneous areas at mid-elevations.     

Landscape-Scale Controls on Aboveground Forest Carbon Stocks on the Osa Peninsula,Costa Rica

Tropical forests store large amounts of carbon in tree biomass, although the environmental controls on forest carbon stocks remain poorly resolved. Emerging airborne remote sensing techniques offer a powerful approach to understand how aboveground carbon density (ACD) varies across tropical landscapes. In this study, we evaluate the accuracy of the Carnegie Airborne Observatory (CAO) Light Detection and Ranging (LiDAR) system to detect top-of-canopy tree height (TCH) and ACD across the Osa Peninsula, Costa Rica. LiDAR and field-estimated TCH and ACD were highly correlated across a wide range of forest ages and types. Top-of-canopy height (TCH) reached 67 m, and ACD surpassed 225 Mg C ha^-1, indicating both that airborne CAO LiDAR-based estimates of ACD are accurate in tall, high-biomass forests and that the Osa Peninsula harbors some of the most carbon-rich forests in the Neotropics. We also examined the relative influence of lithologic, topoedaphic and climatic factors on regional patterns in ACD, which are known to influence ACD by regulating forest productivity and turnover. Analyses revealed a spatially nested set of factors controlling ACD patterns, with geologic variation explaining up to 16% of the mapped ACD variation at the regional scale, while local variation in topographic slope explained an additional 18%. Lithologic and topoedaphic factors also explained more ACD variation at 30-m than at 100-m spatial resolution, suggesting that environmental filtering depends on the spatial scale of terrain variation. Our result indicate that patterns in ACD are partially controlled by spatial variation in geologic history and geomorphic processes underpinning topographic diversity across landscapes. ACD also exhibited spatial autocorrelation, which may reflect biological processes that influence ACD, such as the assembly of species or phenotypes across the landscape, but additional research is needed to resolve how abiotic and biotic factors contribute to ACD variation across high biomass, high diversity tropical landscapes.     

Long-Term Effects of Forest Regrowth and Selective Logging on the Seed Bank of Tropical Forests in NE Costa Rica1.

To investigate long-term effects of land use on the soil seed bank, we compared the abundance/density, species richness, life form distribution, and species composition of seeds stored in the soil of four 15–20 yr-old second-growth stands, two old-growth stands, and two previously selectively-logged stands in the Caribbean lowlands of Costa Rica. Surface soil (10 cm deep, 4.7 cm diameter) was collected at 10 m intervals along three 120–160 m long transects in each stand (44–48 soil cores, 22–24 combined seed bank samples per site). Seed density was highest but variable in second-growth stands (8331–14535 seeds/m²), low and homogeneous in old-growth stands (2258–2659 seeds/m²), and intermediate and highly variable in selectively-logged stands (1165–6854 seeds/m²), which also had contrasting logging intensities. Species richness was strongly dependent on seed density, but showed less variation. Life form distribution did not differ statistically among or within land-use categories. In each stand, herbs-forbs, shrubs, and vines dominated the seed bank (> 75% of the species richness and abundance), whereas trees were a minor component (< 20% of the species richness and < 5% of the abundance) and were predominandy early successional. Shrubs and vines were most abundant in second-growth stands where regrowth vegetation was repeatedly cut before abandonment, whereas grasses and sedges were most abundant in the only forest stand that was completely surrounded by pastures. In terms of species composition, old-growth stands were more similar to selectively-logged stands than to second-growth stands, but across stands, selectively-logged forests were most distinct from the other two forest types. An inventory of the standing woody vegetation in each site showed little representation of the woody taxa found in the seed bank. We discuss these results in the context of the main factors that have been postulated to influence the abundance, life form, and species composition of tropical forest seed banks, and explore the role of the latter during intermediate phases of tropical forest succession and regeneration.     

The genus Neotherina Dognin (Geometridae, Ennominae) in Costa Rica

So far, two species of Neotherina Dognin have been recorded in Costa Rica. Neotherina imperilla (Dognin) occurs primarily at altitudes between 1100 and 1700 meters and Neotherina callas (Druce) which is widely distributed above 1100 meters. A third, new species, Neotherina xanthosa Sullivan and Chacón is described from altitudes above 2400 meters. Heterogeneity of the genus is discussed.     

Postdispersal Seed Fate of Tropical Montane Trees in an Agricultural Landscape, Southern Costa Rica

Variation in postdispersal seed fate is an important factor driving patterns of forest regeneration. Because most previous studies have not tracked final seed fate and have commonly equated seed removal with predation without considering the possibility of secondary dispersal, little is known about individual seed mortality factors in successional and degraded habitats. This study tracked the postdispersal fate of large-seeded tropical montane trees in abandoned pasture, young secondary forests, and small forest fragments during two consecutive years in an agricultural landscape in southern Costa Rica. The incidence of secondary dispersal by animals, scatterhoarding in particular, and the effects of seed burial on germination were measured. Overall, seeds survived through germination more often in secondary forests with high levels of mortality occurring in abandoned pastures and forest fragments. The majority of seed mortality was caused by rodent predation in forest fragments, insects and fungal pathogens in secondary forests, and a combination of desiccation, insects, and fungal pathogens in pastures. Seeds were frequently secondarily dispersed in larger forest fragments, whereas they were only rarely moved in pastures and secondary forests. Burial tended to improve germination in pastures and was important for an often scatterhoarded species, Otoba novogranatensis, in all habitats. The results of this study suggest that: (1) seed mortality factors differ in response to the type of habitat degradation; (2) large-seeded species have high potential for survival when dispersed to young secondary forests; and (3) seed removal is not a reliable proxy for seed predation, particularly in less degraded forest fragments.     

Influence of niche characteristics and forest type on fern species richness,abundance and plant size along an elevational gradient in Costa Rica

An analysis of the fern vegetation on 156 plots along an elevational gradient (45-3400 m) in undisturbed forests in Costa Rica, Central America, showed a hump-shaped pattern of species richness with a maximum of up to 68 species per 400 m2 at mid-elevations. This study documents the contribution of specific habitats (forest types: ridges, ravines) and niches within them (dead wood, rocks, growth zones in trees) to the local fern richness and the relation of species richness to elevation and climatic variables. Forests along ravines showed significantly higher species richness, presumably caused by high environmental humidity. The mean number of individuals of occupied niches per species increased significantly with elevation, suggesting that the niche breadth of species increased and that the differentiation of niches decreased with elevation. Both findings may explain the reduced fern species richness towards and above the upper treeline, but not at low elevations. The key factors for the decreases of species richness at the extremes of the gradient are likely to involve climatic conditions.     

The Effects of the Conversion of a Primary to a Secondary Tropical Lowland Forest on Bullet ant (Paraponera clavata) Foraging Behavior in Costa Rica: A Possible Indicator of Ecosystem Condition

The use of bioindicators of habitat condition can help to better understand the effects of tropical forest degradation and the efficacy of strategies used in the restoration of these lands. The differences in feeding behavior of the ponerine ant Paraponera clavata may serve as such an indicator. The findings from the current study showed that P. clavata in an undisturbed primary forest returned to the nest with prey, nectar, and plant materials, while none of the ants within a 14 year old regenerating secondary forest returned with prey or nectar, few with plant materials, and most of the returns were unsuccessful in their foraging. This suggests a difference in P. clavata feeding behavior and/or food selection is occurring in the disturbed habitat; that P. clavata from the primary forest nest examined in the current study are feeding at a higher trophic level; and that the ants in the primary forest appeared to be more successful and efficient foragers than those in the secondary forest. Future studies should involve more comparisons of P. clavata foraging behavior in secondary to primary forests to determine if the patterns described in this current study are consistent with disturbance in these tropical forests in order to evaluate the possibility of this use of P. clavata as a valuable tool for bioindicators of habitat damage.     

The Disphragis notabilis (Schaus) species-group in Costa Rica (Lepidoptera,Notodontidae)

The four described taxa in the Disphragis notabilis (Schaus) species-group are reviewed, including the types and their dissected genitalia. Disphragis hemicera (Schaus), stat. rev., is elevated to species rank, D. normula (Dognin) is retained as a synonym of D. notabilis, D. sobolis Miller is confirmed as distinct from D. hemicera, and D. bifurcata sp. n., is newly described. Both D. hemicera and D. bifurcata occur in Costa Rica. The known ranges of the other species are outlined. Defining characters of each species are presented and a key to species is provided. Unusual variation in the genitalia is noted.     

Tropical forest restoration in Costa Rica: the effect of several strategies on litter production, accumulation and decomposition

Tropical forest restoration strategies have the potential to accelerate the recovery of the nutrient cycles in degraded lands. Litter production and its decomposition represent the main transfer of organic material and nutrients into the soil substrate. We evaluated litter production, accumulation on the forest floor, and its decomposition under three restoration strategies: plantation (entire area planted with trees), island (trees planted in patches of three different sizes) and control (natural regeneration) plots. We also compared restoration strategies to young secondary forest (7-9 yr). Restoration treatments were established in 50 x 50m plots in June 2004 at six sites in Southern Costa Rica. Planted tree species included two native timber species (Terminalia amazonia and Vochysia guatemalensis) interplanted with two N fixers (Erythrina poeppigiana and Inga edulis). Litter was collected every 15 days between September 2008 and August 2009 in 12 0.25m2 litter traps distributed within each plot; litter that accumulated on the soil surface was collected at four locations (0.25m2 quadrats) within each plot in February and May 2009. Total litter production in plantation (6.3Mg/ha) and secondary forest (7.3Mg/ha) did not differ, but were greater than in islands (3.5Mg/ha) and control (1.4 Mg/ha). Plantation had greatest accumulation of litter on the soil surface (10.6 Mg/ha) as compared to the other treatments (SF = 7.2; I = 6.7; C = 4.9). Secondary forest was the only treatment with a greater annual production of litter than litter accumulation on the soil surface. Carbon storage in litter was similar between plantation and secondary forest, and significantly greater than the other treatments. No differences were found for carbon concentration and storage in the soil among treatments. There was also high variability in the production and accumulation of litter and carbon among sites. Active restoration treatments accelerated the production of litter and carbon storage in comparison to areas under natural recovery. However, the nutrient cycle has not necessarily been restored under these conditions, as high litter accumulation on the soil surface indicates a low decomposition rate, which slows nutrient return to the soil.     

Monitoring coral reefs, seagrasses and mangroves in Costa Rica (CARICOMP)

The coral reefs, seagrasses and mangroves from the Costa Rican Caribbean coast have been monitored since 1999 using the CARICOMP protocol. Live coral cover at Meager Shoal reef bank (7 to 10 m depth) at the Parque Nacional Cahuita (National Park), increased from 13.3% in 1999, to 28.2% in 2003, but decreased during the next 5 years to around 17.5%. Algal cover increased significantly since 2003 from 36.6% to 61.3% in 2008. The density of Diadema antillarum oscillated between 2 and 7ind/m2, while Echinometra viridis decreased significantly from 20 to 0.6ind/m2. Compared to other CARICOMP sites, live coral cover, fish diversity and density, and sea urchin density were low, and algal cover was intermediate. The seagrass site, also in the Parque Nacional Cahuita, is dominated by Thalassia testudinum and showed an intermediate productivity (2.7 +/- 1.15 g/m2/d) and biomass (822.8 +/- 391.84 g/m2) compared to other CARICOMP sites. Coral reefs and seagrasses at the Parque Nacional Cahuita continue to be impacted by high sediment loads from terrestrial origin. The mangrove forest at Gandoca, within the Refugio Nacional de Vida Silvestre Gandoca-Manzanillo (National Wildlife Refuge), surrounds a lagoon and it is dominated by the red mangrove, Rhizophora mangle. Productivity and flower production peak was in July. Biomass (14 kg/m2) and density (9.0 +/- 0.58 trees/100 m2) in Gandoca were relatively low compared to other CARICOMP sites, while productivity in July in Costa Rica (4 g/m2/d) was intermediate, similar to most CARICOMP sites. This mangrove is expanding and has low human impact thus far. Management actions should be taken to protect and preserve these important coastal ecosystems.     

Tropical wetlands for climate change research, water quality management and conservation education on a university campus in Costa Rica

EARTH University is a small agronomic university with a theme of sustainability in eastern Costa Rica. Several natural and constructed wetlands on its campus are used for research, water quality improvement, and higher education. It has become an important location for research and teaching on humid tropical wetland ecology and management. A 112-ha flow-through Raphia taedigera (Arecaceae) forested wetland is being used for climate change research, focusing on carbon sequestration and methane generation. Methane emissions are measured seasonally and are comparable to rates in tropical wetlands published elsewhere. Carbon sequestration by the wetland appears to be substantially higher than similar flow-through temperate zone wetlands. Treatment wetlands are used on campus to improve water quality of effluents from an animal farm, a dairy plant, a landfill, and a banana paper plant. Water quality was substantially improved in all of these wetlands except the landfill leachate wetland. All of these campus wetlands have been integrated into the four-year education program of EARTH University and 22 undergraduate projects have been completed on wetlands over the past 14 years.     

Monitoring of the mangrove forest at Gandoca, Costa Rica (CARICOMP site)

The mangrove forest at Gandoca, Refugio Nacional de Vida Silvestre Gandoca-Manzanillo, Caribbean coast of Costa Rica, has been monitored since 1999, following the CARICOMP protocol. The dominant species was the red mangrove, Rhizophora mangle. The peak of productivity and flowering was in July. The mangrove productivity decline from 2001 to 2004 while the temperature rised. Biomass (14 kg/m2) and density (9 trees/10 m2) in Gandoca were relatively low compared to other CARICOMP sites, while productivity in July in Costa Rica (4 g/m2/day) was intermediate, similar to most CARICOMP sites.     

The Influence of Habitat Variables on Bird Communities in Forest Remnants in Costa Rica

This study examined the effects of forest structure (tree species richness, canopy height, percent canopy cover, understory density, tree density and DBH) and avian species traits (nest type and indicator list status) on the diversity, abundance and dissimilarity of bird communities in forest remnants and reforestation areas adjacent to Costa Rican banana plantations. Bird species richness and abundance were significantly related to tree species richness, canopy height and canopy cover in multiple linear regressions, the latter two forest structure variables being the best statistical predictors. Stratification of analyses by bird species indicator categories improved fits of regressions, because correlations with environmental variables differed in sign for different guilds of birds, a result likely to hold for other avifaunas. Analysis of avifauna dissimilarities among sites demonstrated that the species composition of bird communities was highly correlated with forest structure and tree species composition. Logistic regressions indicated that birds making protected (cavity, burrow, pendant, sphere and covered) nests were 2–6 times more likely to be present in the study avifauna than birds making open (cup, saucer, platform and scrape) nests and indicators of disturbed habitats were 11 times more likely to be present than indicators of primary forest. The forest structure data used were simple and inexpensive to collect, and data on avian traits were drawn from the literature. Thus, these methods could easily be replicated at other locations and would be valuable management aids and biodiversity assessment tools for conservation planning.     

Notes on the ecology of rolled-leaf hispines (Chrysomelidae,Cassidinae) at La Gamba (Costa Rica)

A total of 301 adult hispine beetles of the genera Cephaloleia and Chelobasis were found in rolled leaves of plants of 17 species of Zingiberales (families Costaceae, Heliconiaceae, Maranthaceae, Musaceae, and Zingiberaceae) during a field study at La Gamba, Golfito region, Costa Rica. Of these beetles, Cephaloleia belti was recorded from 12 potential host plant species, C. distincta from 7, C. dilaticollis from 5, C., Chelobasis bicolor, C. championi, and C. histrionica from 3, Chelobasis perplexa and C. instabilis from 2, whereas C. trivittata from only one. Of the plant species, Heliconia latispatha had 7 beetle species in its leaf rolls, Calathea lutea had 5, H. imbricata and H. rostrata had 4, H. stricta and Musa paradisiaca had 3, H. wagneriana had 2, while on H. vaginalis, H. danielsiana, H. densiflora, H. longiflora, Calathea crotalifera, C. platystachya, Goeppertia lasiophylla, Alpinia purpurata, Costus pulverulentus and Costus barbatus, H. densiflora, H. vaginalis, and H. danielsana only hispines of one species were found.Cephaloleia belti occurred together with beetles of six other hispine species, whereas Cephaloleia trivittata never shared a leaf roll with another hispine species. The remaining beetle species aggregated with one to four other hispines. Adults of C. belti and C. championi were frequently seen, occasionally also with C. dilaticollis, C. histrionica, and Chelobasis perplexa, to co-occur with the carabid Calophaena ligata in the same leaf roll without any sign of interspecific aggression.A comparison of host choices and the phylogeny of the hispines and of their host plants revealed no signs that beetles used species level phylogenetic relationships within the Zingiberales to select food plants. Obviously, within this plant order, rolled-leaf hispines choose their plant hosts in a nearly opportunistic manner. Seemingly, they use differences among plants at higher taxonomic levels but within the Zingiberales, the availability of young – rolled – leaves might be the actual decisive factor.     

A food plant specialist in Sparganothini: A new genus and species from Costa Rica (Lepidoptera,Tortricidae)

Sparganocosma docsturnerorum Brown, new genus and new species, is described and illustrated from Área de Conservación (ACG) in northwestern Costa Rica. The new genus shares a long, crescent- or ribbon-shaped signum in the corpus bursae of the female genitalia with Aesiocopa Zeller, 1877, Amorbia Clemens, 1860, Amorbimorpha Kruse, 2011, Coelostathma Clemens, 1860, Lambertiodes Diakonoff, 1959, Paramorbia Powell & Lambert, 1986, Rhynchophyllus Meyrick, 1932, Sparganopseustis Powell & Lambert, 1986, Sparganothina Powell, 1986, and Sparganothoides Lambert & Powell, 1986. Putative autapomorphies for Sparganocosma include the extremely short uncus; the smooth (unspined) transtilla; and the upturned, free, distal rod of the sacculus. Adults of Sparganocosma docsturnerorum have been reared numerous times (>50) from larvae collected feeding on rain forest Asplundia utilis (Oerst.) Harling and Asplundia microphylla (Oerst.) Harling (Cyclanthaceae) at intermediate elevations (375–500 m) in ACG. Whereas most Sparganothini are generalists, typically feeding on two or more plant families, Sparganocosma docsturnerorum appears to be a specialist on Asplundia, at least in ACG. The solitary parasitoid wasp Sphelodon wardae Godoy & Gauld (Ichneumonidae; Banchinae) has been reared only from the larvae of Sparganocosma docsturnerorum.     

Hydroptilidae (Trichoptera) of Costa Rica: the Genus Hydroptila Dalman

Seven new species of Hydroptila (Trichoptera: Hydroptilidae) from Costa Rica are described: H. carara, H. maritza, H. osa, H. paradenza, H. maza, H. rastrilla, H. singri, and one from Panama, H. nusagandia. Ten additional species occurring in Costa Rica are recorded: H. brailovskyi Bueno-Soria, H. constricta Bueno-Soria, H. curvata Bueno-Soria, H. flinti Bueno-Soria, H. icona Mosely, H. meralda Mosely, H. mexicana Mosely, H. misolha Bueno-Soria, H. paschia Mosely, and H. veracruzensis Flint. In addition, illustrations of H. denza Ross and H. grenadensis Flint are included to help clarify the taxonomy of the denza species group. Finally, an illustrated key is provided for males of all species occurring in lower Central America.