Effects of forestry practices on vegetation structure and bird community of Kibale National Park, Uganda

I examined the lingering effects of past timber management practices on the vegetation structure and bird community of Kibale National Park, Uganda. I compared four forest treatments: unlogged native forest (UL), two that were selectively logged at low (LL) and high (HL) intensities in the 1960s, and a conifer plantation (PL). Forest-dependent birds were best represented at UL. LL was similar to UL in both vegetation structure and bird community composition, although some forest-dependent bird species were missing from the former. HL had significantly less canopy closure and lower tree density than other plots as a result of the combination of extensive secondary damage and natural disturbance patterns that prevented the reclosure of the forest canopy. Thirty-one percent of the forest-dependent bird species observed during the study were not detected at HL. At PL, bird species richness and bird abundance were about a third of those observed in other plots. There were significant correlations between heterogeneity of tree distribution (horizontal heterogeneity) and abundance and species richness of birds across plots. Abundance and species richness of all, forest-dependent, and forest generalist birds were highest in plots with intermediate measures of horizontal heterogeneity, which were mostly unlogged or lightly logged. If reduced-impact logging practices are not implemented during selective logging operations in tropical forests, consequent long-term changes in vegetation structure may result in significant declines in the populations of some forest-dependent species, as was observed in Kibale National Park.     

Peat swamp forest avifauna of Central Kalimantan, Indonesia: Effects of habitat loss and degradation

Peat swamp forest is a unique wetland ecosystem covering extensive areas in Southeast Asia that has received relatively little scientific attention and is now being lost at a rapid pace. This study examines the effects of anthropogenic degradation on bird communities in disturbed peat swamp forest habitats – namely, intact logged forest, a degraded forest fragment, and non-forest regrowth – in Central Kalimantan, Indonesia. Results show that species richness was significantly higher and species composition significantly different in intact logged forest in comparison to the degraded forest fragment and non-forested area. Nectarivore and tree foliage-gleaning insectivore abundance declined outside the intact forest, while the regrowth was dominated by the yellow vented bulbul, an open country insectivore–frugivore. Surveys reveal that large intact tracts of logged peat swamp forest can harbour threatened and near threatened bird species (36% of records) and thus play a role in their conservation, especially for a few habitat specialists. Given the extent of unmanaged degraded peatlands and continuing pressure to development them, urgent conservation actions are needed to rehabilitate and protect this ecosystem.     

Bird species richness in a Bornean exotic tree plantation: A long-term perspective

To provide sustainable income from forestlands, large areas in the tropics are planted with “agricultural” trees, such as oil palm and rubber, and “industrial” trees, such as Acacia mangium and Gmelina arborea. To examine how native forest birds use such plantations, we surveyed in 2005 the avifauna at Sabah Softwoods, a plantation in southeastern Sabah, Malaysian Borneo. We focused on A. mangium, Albizia (Paraserianthes falcataria), oil palm (Elaeis guineensis), and logged native forest, and compared our results to those of a study conducted at the same plantation in 1982. The number of forest species in the industrial groves did not change dramatically between 1982 and 2005, even though the trees had been cropped several times and the plantation was, by 2005, completely surrounded by cleared land and far removed from primary forest. However, as is common in logged or isolated forests throughout the world, certain primary forest groups (e.g., muscicapine flycatchers) have been extirpated from the entire plantation area. The industrial groves also lacked some larger species of kingfisher, woodpeckers, and canopy frugivores. Nevertheless, numerous primary forest taxa (ca. 50% of species) were found in mature industrial tree groves. Albizia attracted the most species of birds, followed closely by Acacia. Both tree types underpinned relatively complex secondary forests that attracted forest birds. In contrast, younger groves of Acacia and Albizia held mainly open country and scrub species. Oil palm, as a remarkably simple and unusual habitat, attracted few bird species. Sustained occurrence of forest birds in all groves of exotic trees at Sabah Softwoods was substantially enhanced by the relatively rich avifauna of the logged native forest remaining in substantial stands throughout the plantation.     

Lowland rainforest avifauna and human disturbance: persistence of primary forest birds in selectively logged forests and mixed-rural habitats of southern Peninsular Malaysia

We compared the composition and structure of primary forest avifauna among primary forests, selectively logged forests and mixed-rural areas (e.g. villages and agricultural areas) of Peninsular Malaysia. We found that forests that were selectively logged at least 30 years ago contained only 73-75% of the 159 species of extant primary forest birds, with an increased proportion of dominant species. We estimated that only 28-32% of the primary forest species utilized the mixed-rural habitat, and that the number of species that bred in the agricultural landscapes might be even lower. The microhabitat of different species most affected their vulnerability to disturbance. Most small, arboreal frugivores and omnivores, and insectivores that fed from tree trunks, showed greater persistence in the mixed-rural habitat than ground dwelling bird species, which were affected most by disturbance. Resource abundance and variables that were closely related to forest disturbance such as the density of large trees, density of dead trees, canopy cover density and shrub volume influenced the distribution of the primary forest birds. Large primary forest reserves and a revision of short-cycle logging regimes (ca. 30 years) are needed if we are to conserve the lowland rainforest avifauna of Peninsular Malaysia and other parts of Southeast Asia.     

Comparison of breeding bird and vegetation communities in primary and secondary forests of Great Smoky Mountains National Park

We compared breeding bird communities and vegetation characteristics at paired point locations in primary (undisturbed) and mature secondary forest (70-100 years old) sites in Great Smoky Mountains National Park, USA to understand how sites logged prior to creation of the park compare to undisturbed sites following 70 years of protection from human disturbance. We found that bird and vegetation communities are currently similar, but retain some differences in species composition. Rank abundance curves for primary and secondary forest bird communities showed very similar patterns of species dominance. Species composition was also similar on the two sites which shared 24 of the 25 most frequently recorded species. Nonetheless, comparisons of density estimates derived from distance sampling showed three bird species were more abundant on primary forest sites and that one bird species was significantly more abundant on secondary forest sites. Notably, comparisons based on raw counts (unadjusted for potential differences in detectability) produced somewhat different results. Analyses of vegetation samples for the paired sites also showed relative similarity, but with some differences between primary and secondary forests. Primary forest sites had more large trees (trees greater than 50 cm diameter at breast height) and late successional species. Primary forest sites had a denser tall shrub layer while secondary forest sites had a denser canopy layer. Nonetheless, tree species richness, basal area of live trees and number of standing snags did not differ between primary and secondary forest sites. Results indicate that breeding bird communities on sites within the park that were logged commercially 70 years ago are currently quite similar to bird communities on sites with no history of human disturbance. Similarities between the bird communities on previously disturbed and undisturbed sites in Great Smoky Mountains National Park may exceed those on more fragmented landscapes because large patches of primary forest, adjacent to commercially logged sites, remained in the park when it was established in 1935. These patches of primary forest may have served as source areas for commercially logged sites.     

Bird species abundance–occupancy patterns and sensitivity to forest fragmentation: Implications for conservation in the Brazilian Atlantic forest

Developing a predictive theory for species responses to habitat fragmentation is a large, complex challenge in conservation biology, and meeting this challenge likely requires tailoring predictions to specific habitats and taxa. We evaluate the effects of fragmentation on forest birds living in three distinct forest ecosystems found in Brazilian Atlantic forest: seasonal semi-deciduous forest (SF), mixed rain forest (MF), and dense rain forest (DF). We test the hypotheses that (1) bird species most prevalent in SF (relative to other habitat types) will be least vulnerable to population declines in fragmented SF, and (2) species with stronger affiliations with DF or MF will be relatively more sensitive to fragmentation in SF. Using an exploratory statistical technique called “Rank Occupancy–Abundance Profiles (ROAPs),” we compared distribution and abundance of birds among large “continuous” areas of each forest type, then compared abundances in continuous SF forests with patterns of abundance in small fragments of SF, where edge effects could play a marked role in population dynamics. Overall, 39 species showed substantially lower local abundance, occupancy, or both in SF fragments versus continuous SF. As predicted, a higher proportion of bird species associated with DF appeared sensitive to fragmentation in SF; by contrast, species most abundant in SF and MF were similarly abundant in fragmented SF. Our study demonstrates how quantifying distribution and abundance in diverse habitats may enhance managers’ ability to incorporate species-specific responses to human disturbances in their conservation plans, and points out ways that even small reserves may have significant conservation value.     

Eucalypt plantations as habitat for birds on previously cleared farmland in south-eastern Australia

In parts of Australia, extensive areas of cleared land are now being planted with commercial plantations of native eucalypts. Questions arise about the extent to which such plantations can rectify previous loss of habitat and contribute to biodiversity conservation. This study assessed abundance of bird species (as one aspect of biodiversity) on 105 sites (25 cleared land, 58 plantations and 22 native forest) in two regions of rural Victoria, south-eastern Australia. Generalised linear modelling was used to assess some of the landscape and habitat variables that contributed to the value of plantation sites for particular groups of bird species. These models demonstrated the importance of on-site habitat variables in explaining the abundance of groups of bird species, with landscape context making small additional contributions.Mean abundance of forest and woodland birds was higher in eucalypt plantations than cleared farmland, and marginally lower than in native forest. Patterns differed between bird guilds. For example, insectivores that forage in the canopy and tall shrub layers were at least as common in plantations as in native forest, with birds in the latter group using young eucalypts as if they were tall shrubs. Birds that forage from open ground among trees were more common in plantations than native forest, and may benefit substantially from the new habitat fortuitously provided for them. This group includes several species that have declined in natural woodland habitats. Nectarivores, carnivores and birds that forage among low shrubs were less common in plantations than in native forest. Insectivores that forage from eucalypt bark made little use of plantations. Different approaches to plantation design and management would be needed to cater for groups such as these. Specific measures include planting of rough-barked eucalypts in addition to smooth-barked species, and provision of artificial hollows. Retention of existing remnants of native forest (e.g. old trees and forest patches) is a priority, to supply habitat elements that would otherwise be missing for long periods.     

Habitat structure and proximity to forest edge affect the abundance and distribution of forest-dependent birds in tropical coastal forests of southeastern Madagascar

Despite the fact that Madagascar is classified a biological `hotspot' due to having both high levels of species endemism and high forest loss, there has been no published research on how Madagascan bird species respond to the creation of a forest edge or to degradation of their habitat. In this study, we examined how forest bird communities and different foraging guilds were affected by patch habitat quality and landscape context (forest core, forest edge and matrix habitat) in the threatened littoral forests of coastal southeastern Madagascar. We quantified habitat use and community composition of birds by conducting 20 point counts in each landscape contextual element in October and November 2002. We found that littoral forest core habitats had significantly (p<0.01) more bird species than forest edge and matrix habitats. Thirty-one (68%) forest dependent species were found to be edge-sensitive. Forest edge sites had fewer species, and a higher representation of common species than forest interior sites. Twenty-nine species were found in the matrix habitat, and the majority of matrix-tolerant forest species had their greatest abundance within littoral forest edge habitats. Guild composition also changed with landscape context. Unlike other tropical studies with which we are familiar, we found that frugivorous species were edge-sensitive while sallying insectivores were edge-preferring. The majority of canopy insectivores (n=15, 88%), including all six endemic vanga species, were edge-sensitive. When habitat quality was assessed, the distributions of nine edge-sensitive species were significantly (p<0.01) affected by changes in habitat complexity and vegetation vertical structure in core or edge point counts. Therefore, we believe that changes in vegetation structure at the edge of littoral forest remnants may be a key indicator of mechanisms involved in edge sensitivity of forest dependent species in these forests. Our findings indicate that habitat fragmentation and degradation affect Madagascan bird communities and that these processes threaten many species. With continued deforestation and habitat degradation in Madagascar, we predict the further decline of many bird species.     

Spatial and temporal variations in the use of rice-paddy dominated landscapes by birds in Japan

Agricultural landscapes with spatial and temporal variations interact with each other to affect the existing biodiversity. Though rice fields provide important habitats for birds all over the world, studies so far have rarely explored the effects of landscape heterogeneity on bird species in rice paddy areas. This study investigated the effects of habitat cover and landscape variables on the species richness and the abundance of birds in rice paddy areas in Japan. Data on bird occurrence and the environment were collected at 32 grid squares (1 × 1 km) in the Tone River basin. The richness and the abundance of agricultural wetland species were particularly high in landscapes with large areas of rice fields in summer, when rice fields were irrigated, but in those with large areas of open water in winter, when rice fields were drained. It is important to maintain a combination of rice fields and open water to satisfy multiple habitat requirements by agricultural wetland species throughout the year. Grassland species were positively associated with a rich diversity of land cover including fallow fields and open water, indicating the importance of a simultaneous existence of multiple landscape elements. Forest cover in landscapes positively affected edge species and woodland species. Since forest cover had a relatively strong correlation with edge density, the responses of bird species to changes in forest cover and edge density need to be explored further. This study illustrates the importance of spatial and temporal landscape complementation for bird species in rice paddy areas.     

Effects of selective logging on vascular epiphyte diversity in a moist lowland forest of Eastern Himalaya, India

Effect of selective logging on vascular epiphyte diversity was investigated in a moist lowland forest of Eastern Himalaya. Three epiphytic groups viz. orchids, pteridophytes and non-orchid angiosperm epiphytes were specifically studied in closed, selectively logged and in unlogged forests with treefall gaps. Logging reduced the structural complexity of the forests and altered their microclimate. With logging, there was a general decline in richness and abundance of epiphytes except orchids. The abundance and species composition of pteridophytes and non-orchid angiosperm epiphytes were related to microclimate and substrate features while their richness were correlated only with canopy cover. In contrast, orchid species composition was related to forest structure. A combination of management strategies is required for conservation of all epiphyte groups. A mosaic of logged and unlogged forest patches with undisturbed forests in proximity would maintain the diversity of pteridophytes and other angiosperms. However, for full representation of orchids, it is necessary to maintain the structural diversity of the tree forms.     

Supply of tree-holes limits nest density of cavity-nesting birds in primary and logged subtropical Atlantic forest

Tree cavities are proposed to limit populations and structure communities of cavity-nesting birds. Although the greatest diversity of cavity-nesting birds is found in tropical and subtropical moist forests, little is known about how tropical logging affects the abundance of cavities or cavity-nesting birds. We compared the abundance of cavities and cavity nests between primary and selectively-logged subtropical moist Atlantic forest in Argentina, and conducted the first before-after controlled nest-box addition experiment to determine whether nest sites limit the breeding density of cavity-nesting birds in tropical or subtropical moist forest. Visual inspection of 86 cavities identified through ground-surveys revealed that only 19% were suitable for nesting birds, suggesting that cavity abundance may be overestimated in the literature on tropical forests. Suitable cavities were found in fewer than 1% of 1156 trees <60 cm dbh but 20% of 20 trees >100 cm dbh. Logged forest had half the basal area of primary forest, one third the density of large trees, nine times fewer cavities suitable for nesting birds, and 17 times fewer active nests. When we added nest boxes, nesting density increased on treatment plots but not on control plots in both logged and primary forest, suggesting that cavity supply can limit nest density even in relatively undisturbed forest. This is the first experiment to show how reduced cavity supply in logged tropical forest can limit breeding density of cavity-nesting birds. International initiatives such as forest certification should promote tropical timber management strategies that conserve large live cavity-bearing trees.     

Diversity and abundance of selected animal species in undisturbed forest,selectively logged forest and plantations in East Kalimantan,Indonesia

Animal surveys were conducted in six habitat types within the timber concession of PT International Timber Corporation Indonesia, East Kalimantan. Species diversity was found to be similar in undisturbed forest and in 3- to 5-year-old selectively logged forest, although densities in the latter were considerably less. Both diversity and density declined drastically in forestry plantations, but certain of the species remaining caused significant damage to young trees. Although few animals were observed in areas where there was active road construction and timber felling, their densities in adjacent undisturbed forest areas did not increase, suggesting that the level of local migration actually is low. It is suggested that recolonisation of old logged forest can occur successfully provide that the level of hunting is low and that adjacent areas of undisturbed forest remain to provide a population pool from which colonists can spread.     

Land use and conservation value for forest birds in Central Sulawesi (Indonesia)

Unprecedented deforestation is currently underway in Southeast Asia. Since this trend is likely to continue, it is critical to determine the value of human-modified habitats (e.g., mixed-rural habitat) for conserving the regional native forest avifauna. The impacts of ongoing deforestation on the highly endemic avifauna (33%) of Sulawesi (Indonesia) are poorly understood. We sampled birds in primary and secondary forests in the Lore Lindu National Park in central Sulawesi, as well as the surrounding plantation and mixed-rural habitats. Species richness, species density and population density of forest birds showed a consistent decreasing trend in the following order: primary forests > secondary forests > mixed-rural habitat > plantations. Although primary forests contained the highest proportion (85%) of a total of 34 forest species recorded from our point count surveys, 40-yr old secondary forests and the mixed-rural habitat showed high conservation potential, containing 82% and 76% of the forest species, respectively. Plantations recorded only 32% of the forest bird species. Fifteen forest species had the highest abundance in primary forests, while two species had higher abundance outside primary forests. Our simulations revealed that all forest birds that were sensitive to native tree cover could be found in areas with at least 20% continuous native tree cover. Our study shows that although primary forests have the highest conservation value for forest avifauna, the potential of degraded habitats, such as secondary forests and the mixed-rural habitat, for conserving forest species can be enhanced with appropriate land use and management decisions.     

Density and distribution of the endemic Seram cockatoo Cacatua moluccensis in relation to land use patterns

The endemic Seram cockatoo, Cacatua moluccensis, was placed on Appendix I of CITES in response to declining trade statistics but in the absence of population data. We conducted population surveys and collected data on habitat structure at seven sites on Seram in 1998. Cockatoo densities ranged from 0.93 to 17.25 birds/km² and averaged 7.9 birds/km² across sites. We classified habitat into three types but found that cockatoo densities did not correspond closely to habitat differences across sites. Cockatoo abundance was significantly associated with presence of potential nest trees and strangling figs. Analysis of forest cover and landuse indicates that while most of the island is still covered in lowland forest, only 14% of these forests are protected and almost half the island is classified as logging concession. Conservation recommendations include better information and enforcement of laws protecting Seram cockatoos and resolution of boundary conflicts between parks and logging concessions.     

Agricultural landscape modification increases the abundance of an important food resource: Mistletoes, birds and brigalow

Agricultural landuse alters landscape pattern, with important consequences for native species. Complex responses to changes in landscape pattern may be observed for species with strong inter-specific relationships. We investigate how the abundance of an ecologically important food resource, the grey mistletoe (Amyema quandang), is influenced by the interactive influences of human-modified landscape pattern, bird disperser abundance and host condition. Mistletoe abundance was estimated in brigalow (Acacia harpophylla) dominated remnants in three separate study areas with differing landscape patterns. Negative binomial generalised linear models of mistletoe abundance were tested with combinations of seven explanatory variables: study area, landscape connectivity and edge contrast, patch shape, bird disperser abundance, brigalow foliage cover and basal area of standing dead trees. Ranking of model averaged parameter estimates showed that study area, bird disperser abundance, patch shape, dead trees and connectivity had the strongest influence on A. quandang abundance. Mistletoe abundance apparently increased with landscape modification, particularly in narrow linear patches, but may be limited by the availability of seed dispersers and host condition. Given high landscape modification and ongoing degradation, management should be targeted towards maintaining brigalow remnant condition to ensure bird and mistletoe populations can be supported in the long-term.     

Linking forest structure and composition: avian diversity in successional forests of Chiloé Island, Chile

We show how Chilean forest bird species richness, abundance and guild structure changes as a function of structural properties of forest stands. We surveyed bird assemblages in two old-growth (>200 years), two mid-successional (30-60 years), and two early-successional forest stands (4-20 years), from November 1999 to September 2000, on Chiloé Island, southern Chile (42°S). Birds were grouped into four habitat-use guilds: large-tree users, vertical-profile generalists, understory species, and shrub-users that occasionally use forests. We recorded a total of 24 bird species: 21 in old-growth, 14 in mid-successional and 16 in early-successional stands. Large-tree users and understory birds were most abundant in old-growth stands, vertical-profile generalists were common in both old-growth and mid-successional stands, and shrub-users were only common in early-successional stands. For nine bird species we found significant relationships between their local abundance and forest structural elements. Higher bird densities in old-growth forests were associated with greater availability of canopy emergent trees, snags, logs and understory bamboo cover in this habitat. Accordingly, bird species diversity in forest stands can be predicted by the presence of these structural elements, and forests should be managed to conserve structural elements that create favorable habitat for bird species in order to prevent future species losses due to logging practices.     

Conservation of seed-dispersing migrant birds in Mediterranean habitats: Shedding light on patterns to preserve processes

Migratory frugivorous birds disperse the seeds of many plant species, forming mutualistic associations that render frugivores a priority for conservation in many habitats worldwide. We analysed the distribution of seed-dispersing frugivorous passerines in southern Spain, which is an important area for the conservation of European birds during winter. Frugivorous birds showed similar regional abundance and richness during four winters, although fruit availability changed among years. However, the spatial distribution of frugivorous birds in the area changed among years. These changes were principally determined by annual variation in the distribution of fruits in the area, revealing a clear ability of birds to track the distribution of fruits. The unpredictable distribution of fruits each year suggests that regional fruit crops, rather than selected habitat patches, need to be protected for the long-term conservation of frugivorous bird populations in wintering grounds. Remarkably, the distribution of frugivores was independent of forest development or general cover of shrubs, which helps to reconcile the protection of fruiting shrubs with forest cleaning, an usual management to prevent devastating summer fires that is destroying fleshy-fruited plant communities in many areas of southern Spain. Thus, leaving a part of the fruiting shrubs untouched when cleaning forest undergrowth will allow the settlement of frugivorous birds. Interestingly, both abundance and richness of frugivores decreased with elevation, probably as a consequence of impaired climatic conditions at high altitude, revealing the importance of lowland shrublands as wintering grounds for frugivorous birds. These habitats deserve special conservation efforts, as they are seriously threatened by the ongoing encroachment of agricultural and urban areas along the Mediterranean coasts.     

Changes in bird species composition on a remote and well-forested Wallacean Island, South-East Asia

In South-East Asia patterns of forest loss can predict the number of threatened birds. On the oceanic islands of Wallacea, small-scale agriculture, rather than commercial logging, is the main cause of forest loss and conversion, but few studies have directly examined their responses to land use. In 2001, I studied the birds of primary forest and anthropogenic gardens on the well-forested, and remote, Damar Island (198 km²) in the Banda Sea. Furthermore, I examined broad changes to the islands bird fauna by comparing baseline bird lists obtained in the 19th century with 2001 data. The conversion of primary forest to garden resulted in substantial changes in avifaunal composition, and to the abundance of individual bird species, particularly to frugivorous and globally restricted-range species. Of 15 common birds, four were significantly more abundant in primary forest (blue-tailed imperial pigeon Ducula concinna, northern fantail Rhipidura rufiventris, golden whistler Pachycephala pectoralis and red-chested flowerpecker Dicaeum maugei) and one (scaly-breasted honeyeater Lichmera squamata) was more abundant in garden habitat. Incidental observations provided further evidence that many forest specialists (including the island endemic Damar flycatcher Ficedula henrici) rarely, if ever, used garden habitats notwithstanding its relative proximity. The number of resident birds recorded in 2001 (39) was similar to the 1890s (38), but six forest-dependent passerines were extirpated between samples. These losses are associated with the conversion of 25% of the primary forest to modified habitat since the 1890s, but given the sparse species and environmental change data available it is impossible to definitively pinpoint this as the only cause underlying the species losses. This study demonstrates that remote and relatively well-forested islands are not immune to the threatening processes impacting bird populations elsewhere in South-East Asia.     

The importance of gallery forests in the tropical lowlands of Costa Rica for understorey forest birds

Rivers and streams lined by narrow forest strips are common in the lowland countryside of south-western Costa Rica. We studied the importance of these gallery forests for understorey birds, especially forest species. Using mist-nets, we captured 1110 birds belonging to 90 species between June and September 2007 at 16 sites spread equally over four habitat types: forest interior, forest margin, gallery forest connected to closed forest, and isolated gallery forest. Though isolated gallery forests had the greatest number of expected species in total, they supported the lowest number of forest-specialist species, lower even than connected gallery forests. Granivorous birds showed an increase from forest to isolated gallery forest, all other feeding guilds however showed no change. The studied habitats differed from each other in their faunal composition. Assemblages of three species groups categorised by their habitat preferences (forest specialists, forest generalists, non-forest species) showed nested distribution patterns across the four habitats. There was no significant difference in the proportion of birds with brood patches or of recaptures across the studied habitat types, leading to the assumption that birds not only use gallery forests for movement and foraging but also for reproduction. Though of limited conservation value for most forest understorey birds, at least for a small fraction of forest species gallery forests constitute an important secondary habitat. More significantly, they can form corridors or stepping stones that allow movements within the matrix of human-dominated habitats, and represent an important landscape component benefiting total bird species richness in the Costa Rican countryside.