Isthmian-Atlantic moist forests
Covering the lowland Atlantic versant at chiefly below 500 metres elevation in southern Nicaragua, northern Costa Rica, and most of Panama, the Isthmian-Atlantic moist forests represent the epitome of wet, tropical jungle. This forest ecoregion evolved from unique combinations of North American and South American flora and fauna, which came together with the joining of these continents around three million years before present.
The ecoregion is classified to be within the Tropical and Subtropical moist broadleaf forests biome. Currently, much of this ecoregion has been converted to subsistence and commercial agriculture. The Isthmian-Atlantic moist forests exhibit a high level of species richness, with 1021 vertebrate taxa alone having been recorded here, with a particularly vast assortment of amphibians, many of which are endemic or near endemic; moreover, among the amphibians there are representatives of anuran, salamander and caecilian taxa.
Location and general depiction
This ecoregion located at the juncture of Central and South America. Condensation over the warm land produced by moisture-laden air from the Caribbean Sea colliding with the mountains produces constant high humidity and precipitation. Annual rainfall ranges from about 2500 millimetres (mm) in central Panama to over 5000 mm in southern Nicaragua. These forests are distinguished from the cooler subtropical Central American Atlantic moist forest ecoregion to the north by their distinct geologic history and consistent annual temperatures above 24°C.
Until recent geologic times, the isthmus south of central Nicaragua was discontinuous, volcanically active and topographically and environmentally diverse. Basalt bedrock is the parent material of the residual and often unconsolidated soils covering the hilly areas of this ecoregion. Old alluvial terraces form the base of the swamp forests and flat lands in the lowest elevations and near the Caribbean Sea coast. The northern section of this ecoregion is formed of a wide, relatively flat alluvial plain, with a gradual elevation change from sea level to 500 metres in elevation, while south into Panama, steep slopes rise up from the Atlantic Ocean, significantly narrowing the ecoregion to only five to ten kilometres in width.
This ecoregion is characterised by a lush, high canopy tropical evergreen forest of huge buttressed trees reaching 40 metres (m) in height, and an associated rich epiphytic flora. The palm component includes many sub-canopy and understory species. Abundant subcanopy palm species are Amargo Palm (Welfia regia), Walking Palm (Socratea exorrhiza), and in permanently flooded areas, Raphia taedigera. Seasonal swamp forests occur in the lowest and flattest areas in Nicaragua and northern Costa Rica, particularly along the coastal zone, where they grade into mangrove forests. In these swamp forests, Gavilán Tree (Pentaclethra macroloba) dominates the canopy, along with Caobilla (Carapa nicaraguensis). The Almendro (Dipteryx panamensis) and the Monkey-pot Tree (Lecythis ampla) are two notable canopy emergents, although both are diminishing in populations; moreover, they are regional endemic trees of the lowlands, below 250 m in elevation.
While biologically quite diverse, this ecoregion exhibits low levels of endemism. The high species richness is derived in great part from the mixing of North and South American floras and faunas on this land bridge. At least 1021 vertebrates have been observed her. The resident fauna, including butterfly, reptilian, amphibian, avian, and mammalian taxa are generally wide-ranging representative species of a wet tropical forest ecoregion that extends from southern Mexico to northern South America. Strict endemism among fauna is almost non-existent: between 80 to 100 percent of the mammalian taxa found in northern Costa Rica also occur in Panama, Nicaragua, Honduras, and Colombia. However, a number of restricted range bird species are shared with the Central American Atlantic moist forests ecoregion to the north, together forming an Endemic Bird Area. The Caribbean versant is a major migration route; neotropical and altitudinal migrants comprise about thirty percent of the avifauna, particularly against the foothills.
Few large expanses of primary rainforest remain intact, occurring only in large reserves, particularly the Indio-Maiz Biological Reserve (approximately 400,000 hectares) along Nicaragua's coast and in eastern Panama along La Amistad International Park. These blocks retain nearly all vertebrate species of the ecoregion, including most large predators, although increasing isolation threatens their long-term viability. While small in areal size, the 1500 hectare La Selva Biological Station in northeastern Costa Rica hosts permanent populations of large predators such as the Jaguar (Panthera onca) and herbivores like Baird's Tapir (Tapirus bairdii), probably because of its biological corridor connection to the upper montane forests of Braulio Carrillo National Park.
In fact, this connection represents the last Central American intact gradient of primary forest from near sea level to 2900 metres elevation. Tortuguero National Park, along the Caribbean coast of northern Costa Rica, acts as an isolated refuge to many species (as does Barra de Colorado Wildlife Refuge); however, enforcement of protection remains a challenge in both areas. Logging and clearing of remaining forests threaten many species of slow-growing trees, such as the Almendro and Monkey-pot Tree. The Atlantic lowlands and middle elevations contain some of the rarest butterfly species in Central America and some of the world's highest butterfly species richness. There are numerous special status taxa that are found in the Isthmian-Atlantic moist forests, denoted as Near Threatened (NT), Vulnerable (VU), Endangered (EN), or Critically Endangered (CR).
A considerable number of amphibian taxa occur in the ecoregion. Endemic anurans to the Isthmian-Atlantic moist forests include the Misfit Leaf Frog (Agalychnis saltator), which breeds in swamps, but lives mostly in the tree canopy; the Tilaran Robber Frog (Craugastor mimus); Diasporus tigrillo and the Cross-banded Treefrog (Smilisca puma), found only on the Caribbean versant of Costa Rica and Nicaragua. A further endemic frog to the ecoregion is the Rio Changena Robber Frog (Craugastor jota), narrowly limited to Río Changena, Provincia Bocas del Toro, Panamá. Other anuran species found here are: Veragua Robber Frog (Craugastor rugosus), a nocturnal anuran whose ova are laid in leaf litter; Agua Buena Robber Frog (Diasporus vocator), whose breeding occurs in bromeliads; Noble's Robber Frog (Craugastor noblei); Pygmy Robber Frog (Pristimantis ridens), who deposits eggs in leaf litter; La Loma Robber Frog (Pristimantis caryophyllaceus), whose eggs are deposited on leaf surfaces where the female broods them; Limon Robber Frog (Pristimantis cerasinus); Almirante Robber Frog (Craugastor talamancae), a small sexually dimorphic frog found in leaf litter; Leopard Robber Frog (Pristimantis pardalis), a nocturnal frog found only in Panama and Costa Rica; Atlantic Robber Frog (Craugastor andi); Evergreen Robber Frog (Craugastor gollmeri); Mountain Robber Frog (Pristimantis altae), found only in Costa Rica and Panama among bromeliads and leaf litter; Isla Bonita Robber Frog (Craugastor crassidigitus); White-spotted Cochran Frog (Cochranella albomaculata); Spiny Cochran Frog (Cochranella spinosa), usually found in shrubs overhanging streams where ova are laid on leaves; Paramba Cochran Frog (Centrolene prosoblepon), who breeds in streams within primary forests; Sabinal Frog (Leptodactylus melanonotus), who breeds in foam nests within burrows; Splendid Leaf Frog (Cruziohyla calcarifer), whose ova are laid in water filled tree crevices; Lemur Leaf Frog (Hylomantis lemur), a species who lays eggs on plants overhanging water; Spurrell's Leaf Frog (Agalychnis spurrelli), whose extensive webbing aids in arboreal gliding; Blue-sided Leaf Frog (Agalychnis annae EN), a limited range taxon found only in northern Cordillera de Talamanca, Cordillera de Tilarán and Cordillera Central, Costa Rica and possibly western Panama; Bob's Robber Frog (Craugastor punctariolus), a west-central Panama endemic.
Other anurans found here are: Smoky Jungle Frog (Leptodactylus pentadactylus), often found near swamps and other lotic waters; Peralta Frog (Lithobates taylori), whose eggs attach to aquatic vegetation; Boquete Rocket Frog (Silverstoneia nubicola), found from a single locale in southwestern Costa Rica and adjacent western Panama into eastern Panama in lowland and premontane zones to the Pacific coast of western Colombia, south to Valle de Cauca; Talamanca Rocket Frog (Allobates talamancae), usually found near fast flowing streams; Rainforest Rocket Frog (Silverstoneia flotator); Pratt's Rocket Frog (Colostethus pratti) and Pristimantis museosus.
Treefrogs found in the ecoregion include: Yellow Cricket Treefrog (Dendropsophus microcephalus), associated with marshes and lagoons; Zetek's Treefrog (Isthmohyla zeteki), whose larvae develop in bromeliads; Volcan Barba Treefrog (Isthmohyla picadoi), whose larvae develop in bromeliads; Veined Treefrog (Trachycephalus venulosus), a canopy dweller; Coronated Treefrog (Anotheca spinosa), who lays eggs in bromeliads and water cavities of trees; San Carlos Treefrog (Dendropsophus phlebodes), that breeds in shallow ephemeral grassy pools; Rosenberg's Treefrog (Hypsiboas rosenbergi ), who breeds in a shallow depression constructed by the male in the riparian zone; Sipurio Snouted Treefrog (Scinax elaeochrous), an arboreal frog often found at the canopy top; Red-eyed Treefrog (Agalychnis callidryas); Veragua Cross-banded Frog (Smilisca sordida), found in bromeliads or rocky streams; New Granada Cross-banded Treefrog (Smilisca phaeota); Panama Cross-banded Treefrog (Smilisca sila); Mahogany Treefrog (Tlalocohyla loquax); Lancaster's Treefrog (Isthmohyla lancasteri), whose ova are deposited on leaves overhanging surface waters; Cope's Brown Treefrog (Ecnomiohyla miliaria); Hourglass Treefrog (Dendropsophus ebraccatus); La Loma Treefrog (Hyloscirtus colymba), who breeds in swift creeks and whose larvae cling to rocks by a sucking oral disk; Boulenger's Snouted Treefrog (Scinax boulengeri), occurring from the Caribbean versant of Nicaragua, Costa Rica, and Panama to the Pacific versant from Costa Rica to Ecuador; and Mexican Treefrog (Smilisca baudinii).
There are many Robber Frogs within the Isthmian-Atlantic moist forests, including: Fleischmann's Robber Frog (Craugastor fleischmanni), a Costa Rican endemic; Bransford's Robber Frog (Craugastor bransfordii), whose range is restricted within eastern Nicaragua to central Costa Rica; Fort Randolph Robber Frog (Pristimantis gaigei), often found in caves or along rocky riparian zones; Canal Zone Treefrog (Hypsiboas rufitelus), found along the Atlantic versant from eastern Nicaragua to the Panama Canal Zone; Caretta Robber Frog (Diasporus diastema).
Further anurans found in the ecoregion are: Central American Rainfrog (Craugastor rugulosus); Cerro Utyum Robber Frog (Craugastor podiciferus), found on both versants of the cordilleras of Costa Rica and adjacent western Panama; Warszewitsch's Frog (Lithobates warszewitschii); Turbo White-lipped Frog (Leptodactylus poecilochilus), an adaptive taxon even found in urban zones; Tungara Frog (Engystomops pustulosus), whose males call at night while floating; Grainy Cochran Frog (Cochranella granulosa), who deposits eggs in plants overhanging streams; Mitred Toad (Rhinella margaritifera); Blackbelly Toad (Rhaebo haematiticus), who breeds in rocky pools; Veragoa Stubfoot Toad (Atelopus varius), a Panama endemic, virtually extirpated in Costa Rica; Cerro Utyum Toad (Crepidophryne epiotica), found only in Costa Rica and Panama; Gulf Coast Toad (Incilius valliceps); Southern Roundgland Toad (Incilius coccifer); Evergreen Toad (Incilius coniferus); Wet Forest Toad (Incilius melanochlorus), who breeds in sizable streams; Cane Toad (Rhinella marina); Granular Toad (Rhinella granulosa); Nicaragua Narrowmouth Toad (Gastrophryne pictiventris), an ecoregion endemic; Green-striped Glass Frog (Hyalinobatrachium talamancae), an endemic to this ecoregion; Suretka Glass Frog (Hyalinobatrachium chirripoi), whose eggs are laid on leaves overhanging streams; Plantation Glass Frog (Hyalinobatrachium colymbiphyllum); La Palma Glass Frog (Hyalinobatrachium valerioi), whose eggs are laid on vegetation overhanging waters; Fleischmann's Glass Frog (Hyalinobatrachium fleischmanni), who breeds chiefly along forest streams; Chiriqui Glass Frog (Cochranella pulverata), occurring widely from Honduras to Colombia; Fitzinger's Robber Frog (Craugastor fitzingeri); Chiriqui Robber Frog (Pristimantis cruentus), narrowly found from northern Costa Rica to western Panama; Rusty Robber Frog (Strabomantis bufoniformis), seen only in climax forest; Common Rocket Frog (Colostethus inguinalis); Lovely Poison Frog (Phyllobates lugubris); Confusing Poison Frog (Ameerega maculata), a rare species seen only in Panama; Strawberry Poison-dart Frog (Oophaga pumilio), whose males are strongly territorial; Rufous-eyed Brook Frog (Duellmanohyla rufioculis); Savage's Brook Frog (Duellmanohyla lythrodes EN), known only to the Atlantic versant of eastern Costa Rica and western Panama; Vaillant's Frog (Lithobates vaillanti); Mexican White-lipped Frog (Leptodactylus labialis); Green and Black Poison Dart Frog (Dendrobates auratus), whose tadpoles devour protozoans and rotifers; Granulated Poison Dart Frog (Oophaga granulifera), whose tadpoles are transported by the female to finish metamorphosis in bromeliads; San Miguel Island Frog (Leptodactylus insularum); Horned Marsupial Frog (Gastrotheca cornuta EN), who has the largest eggs of any amphibian, carried by the female on a dorsal pouch until hatched directly to froglets; Costa Rica Brook Frog (Duellmanohyla uranochroa CR), found on both versants of the cordilleras of Costa Rica and western Panama; Costa Rica Nelson Frog (Nelsonophryne aterrima);
fossorial species; Ringtail Salamander (Bolitoglossa robusta), seen only in Costa Rica and western Panama; Cukra Climbing Salamander (Bolitoglossa striatula), a taxon usually found foraging near ponds; False Cienega Colorado Worm Salamander (Oedipina pseudouniformis EN), usually found in swamps and deep forest; Gamboa Worm Salamander (Oedipina complex), who breed by direct development and is not dependent on surface waters; Limon Worm Salamander (Oedipina alfaroi VU); the ecoregion endemic Long-tailed Worm Salamander (Oedipina gracilis EN); La Loma Salamander (Bolitoglossa colonnea), found only in Panama and Costa Rica and uses a form of camoulflage making itself look like a twig.The rare Los Diamantes Worm Salamander (Oedipina carablanca EN) is an endemic salamander to the ecoregion, and is known from only four sites on the Atlantic versant of Costa Rica. Other salamanders found in the ecoregion are: Tapanti Mushroom-tongue Salamander (Bolitoglossa epimela), endemic to the Atlantic versant of central Costa Rica; Rio Quiri Salamander (Bolitoglossa gracilis VU); Cocle Mushroom-tongue Salamander (Bolitoglossa schizodactyla), found chiefly on the Atlantic versant of Panama (with a single specimen observed in Costa Rica); Alvarado's Salamander (Bolitoglossa alvaradoi), an arboreal salamander that is a bromeliad specialist; Colombian Worm Salamander (Oedipina parvipes), usually found amid leaf litter; Costa Rica Worm Salamander (Oedipina cyclocauda), a semi-
The La Loma Caecilian (Dermophis parviceps) is a subterranean amphibian species found within the Isthmian-Atlantic moist forests in saturated soils and sometimes under fallen logs; the Mexican Caecilian (Dermophis mexicanus VU) is also found in the ecoregion. The Varagua Caecilian (Gymnopis multiplicata), found in the ecoregion, is a viviparous organism that can attain a length of up to one half metre.
An endemic reptile found in the Costa Rican part of the ecoregion is the Viquez's Tropical Ground Snake (Trimetopon viquezi). Four taxa of marine turtles are found in the ecoregion's coastal zones, including the Green Turtle (Chelonia mydas EN), who may take almost six decades to reach sexual maturity; the Hawksbill Sea Turtle (Eretmochelys imbricata CR) is another marine species found here. In addition a number of freshwater turtles are found here such as the Brown Wood Turtle (Rhinoclemmys annulata LR/NT). Other reptiles found in the ecoregion include the Spectacled Caiman (Caiman crocodilus LR/NT); Cienega Colorado Worm Salamander (Oedipina uniformis NT), a limited range amphibian found only in Costa Rica along slopes surrounding the Meseta Central;
A wide gamut of tropical bird species occur in the ecoregion. Scarlet Macaw (Ara macao) and Great Green Macaw (A. ambiguus) nest in the lowland forests; moreover, the Great Green Macaw seasonally moves to higher elevations within the ecoregion. Other endangered restricted range species that migrate seasonally to this ecoregion are the Three-wattled Bellbird (Procnias tricarunculata) and the Bare-necked Umbrella Bird (Cephalopterus glabricollis).
Although a few large blocks of intact habitat are extant, the once vast Atlantic lowland forests have been severely fragmented in the latter twentieth and early twenty-first century. The tropical evergreen forests are among the least well represented of Costa Rica's protected areas, although large reserves exist in southern Nicaragua and eastern Panama. Only about 130,000 hectares in the lowland Atlantic zone are currently protected, and difficult economic conditions offer scant likelihood that the area under protection will be significantly expanded.
The lack of protection of the Atlantic lowlands and the heavy bias toward deforestation at elevations of less than 1000 metres contribute to the fragmentation and elimination of these forests. With gradual slopes and relatively good access, much of Costa Rica's remaining Atlantic versant forest has been intervened or exists in small fragments. Nicaragua's lack of access and the until-recently inaccessible steeper slopes of western Panama's Atlantic lowlands and foothills have left these areas relatively forested.
Ecological threat profile
Relatively level areas with alluvial soils are under banana cultivation, while the less fertile hilly basaltic soils have more recently been logged and converted to cattle pasture. New access roads, malaria control, and incentives for migration to these areas in the last few decades have encouraged settlement and resource extraction from the area. The last remaining intact forests in this ecoregion are currently under tremendous logging pressure and are being felled, rapidly. Squatting and other absence of clear property rights have resulted in unregulated destruction in many areas, despite laws in place to protect forests. Clearcutting has also been illegally conducted within many parks, including Tortuguero National Park, which now provides ready access for poachers to the once isolated second largest Green Turtle (Chelonia midas) nesting beach in the world.
While future threats in this ecoregion vary among the three countries, destruction of forest habitat through logging and conversion to cattle pasture are the most widespread and significant. Illegal logging and squatting is making inroads to the remaining large forest blocks in Nicaragua, while in the already fragmented Costa Rican forests; the principal threat is clearing for cattle production. In Panama, the Atlantic slopes are steeper and less likely to be converted to agriculture; however, new roads in this region are accelerating human settlement and cutting of accessible forests. The Mesoamerican Biological Corridor project, which enhances connectivity among the Atlantic slope habitats, offers support for new or expanded protected areas, or for payments for environmental services provided by private lands.
Justification of ecoregion delineation
The delineations for this ecoregion were derived from two separate national vegetation maps. In Costa Rica we followed the delineations of life zones by Tosi. In this case we lumped the following ecoregion of the Pacific slope catchment, south of the Central Valley: Tropical Wet Forest, Premontane Wet Forest, Tropical Moist Forest (TMF), TMF Perhumid Province Transition, and TMF Premontane Transition Belt. The western delineation is marked by the continental divide, or where it does not abut to montane forests at higher elevations. In Panama reliance is placed on the UNDP vegetation map, again, lumping the following lowland and premontane moist forest components on the southern portion of the continental divide: Tropical Moist Forest, Premontane Rainforest, and Tropical Wet Forest. The northern delineation, which separates this ecoregion from the Central American Atlantic moist forests to the north, was determined by expert opinion due to distinct species associations. This ecoregion hosts many endemic species.
- Asociación Nacional Pro Medio Ambiente y Recursos Naturales (ProAmbiente). 1998. Evaluación Ecológica Rápida: Corredor Biológico Talamanca-Caribe. The Nature Conservancy.
- J.R. Barborak, A. F. Carr III, and L. D. Harris. 1994. Recomendaciones para la consolidación territorial y conectividad de las áreas protegidas de Costa Rica. A. Vega, editor, Corredores Conservacionistas en la Región Centroamericana: Memorias de una Conferencia Regional auspiciada por el Proyecto Paseo Pantera. Florida: Tropical Research and Development, Inc.
- M.A. Boza. 1996. El Corredor Biológico Mesoamericano. In Informe de la reunión de la comisión técnica sectorial binacional de recursos naturales, Proyecto Binacional de Manejo y Conservación de la Reserva de la Biósfera La Amistad Costa Rica-Panamá. Boquete, Panama.
- L. Cardenal Sevilla. 1994. Informe de País Nicaragua: Diversidad y Prioridades. In A. Vega, editor. Corredores Conservacionistas en la Región Centroamericana: Memorias de una Conferencia Regional auspiciada por el Proyecto Paseo Pantera. Florida: Tropical Research and Development, Inc.
- S.D. Davis, V.H. Heywood, O. Herrera MacBryde, J. Villa-Lobos and A.C. Hamilton, editors. 1997. Centres of Plant Diversity. A Guide and Strategy for their Conservation. Volume 3. The Americas. IUCN Publications Unit, Cambridge, U.K. 562 pp.
- F. Delgado. 1985. Present situation of the forest birds of Panama. In A. W. Diamond, and T.E. Lovejoy, editors, Conservation of tropical forest birds. ICBP Technical Publication No. 4. UK: International Council for Bird Preservation. ISBN: 0946888051
- DeVries, Philip. 1987. The butterflies of Costa Rica and their natural history. Volumes 1 and 2. Princeton: Princeton University Press.
- Dinerstein, E., et al. 1995. An Evaluation on the status of conservation of terrestrial ecoregions of Latin America and the Caribbean. DC: World Wildlife Fund-US.
- Emmons, L. 1990. Neotropical Rainforest Mammals. Chicago: University of Chicago Press. ISBN: 0226207218
- Gómez, L. D., editor. 1986. La vegetación de Costa Rica (map series). Vol. I, In Vegetación y clima de Costa Rica. Ed. UNED. Costa Rica. cited in R.García, 1997. Biología de la conservación y áreas silvestres protegidas: situación actual y perspectivas en Costa Rica. INBio. Costa Rica.
- Guyer, C. 1990. The Herpetofauna of La Selva, Costa Rica. In A. H. Gentry, editor, Four neotropical rainforests. Yale Univesity Press. ISBN: 0300054483
- Hartshorn, G. 1983. Plants Introduction. In D. H. Janzen, editor, Costa Rican natural history. Chicago: University of Chicago Press. ISBN: 0226393348
- C. Michael Hogan. 2012. Craugastor talamancae (Dunn, 1831) Almirante Robber Frog . GlobalTwitcher
- Holdridge, L.R. 1967. Life zone ecology. Costa Rica: Tropical Science Center.
- Lieberman, D., M. Lieberman, R. Peralta, and G. S. Hartshorn. 1996. Tropical forest structure and composition on a large-scale altitudinal gradient in Costa Rica. Journal of Ecology 84: 137-152.
- Mendez, E. 1994. Estado de la Conservación de Biodiversidad en Panamá. In A. Vega editor, Corredores Conservacionistas en la Región Centroamericana: Memorias de una Conferencia Regional auspiciada por el Proyecto Paseo Pantera. Florida: Tropical Research and Development, Inc.
- Powell, G. 1999. Personal communications regarding linework for ecoregional deliniations.
- Powell, G., J. H. Rappole, and S. A. Sader. 1992. Neotropical migrant landbird use of lowland Atlantic habitats in Costa Rica: a test of remote sensing for identification of habitat. In J.M. Hagan III, and D.W. Johnston, editors. Ecology and conservation of Neotropical migrant landbirds. DC: Smithsonian Institution Press.
- Palminteri, S., G. Powell, A. Fernandez, and D. Tovar. 1999. Talamanca Montane-Isthmian Pacific Ecoregion-Based conservation plan: Preliminary reconnaissance phase. Report to WWF-Central America.
- Raven, P. H. 1985. Research, Tropical ecology, and the future of Panama. In W. G. D'Arcy and M. D. Correa, editors. The botany and natural history of Panama. Saint Louis: Missouri Botanical Garden. ISBN: 0915279037
- Reid, F. 1997. A field guide to the mammals of Central America and Southeast Mexico. New York: Oxford University Press. ISBN: 0195064011
- Rich, P. V. and T. H. Rich. 1983. The Central American dispersal route: biotic history and paleogeography. In D. H. Janzen, editor, Costa Rican natural history. Chicago: University of Chicago Press. ISBN: 0226393348
- Ridgely, R. 1976. A guide to the birds of Panama. Princeton, New Jersey: Princeton University Press.
- Sánchez-Azofeifa, G. A., C. Quesada-Mateo, P. Gonzalez-Quesada, S. Dayanandan, and K. S. Bawa. 1999. Protected areas and conservation of biodiversity in the tropics. Conservation Biology 13: 407-411.
- Stattersfield, A.J., M.J. Crosby, A.J. Long, and D.C. Wege. 1998. Endemic bird areas of the world: Priorities for biodiversity conservation. Birdlife Conservation. Series No. 7. Birdlife International, Cambridge, UK.
- Stiles, F.G. 1985. Conservation of forest birds in Costa Rica: problems and perspectives. In A. W. Diamond, and T. E. Lovejoy, editors. Conservation of tropical forest birds. ICBP Technical Publication No. 4. UK: International Council for Bird Preservation. ISBN: 0946888051
- Stiles, F.G., and A. F. Skutch. 1989. A guide to the birds of Costa Rica. New York: Cornell University Press.
- Stiles, F.G., and D.A. Clark. 1989. Conservation of tropical rain forest birds: a case study from Costa Rica. In American birds. Vol. 43. No. 3. UK: International Council for Bird Preservation.
- Tosi Jr., J.A. 1969. Republica de Costa Rica: mapa ecológico. Map 1:750,000. Tropical Science Center, San Jose, Costa Rica.
- UNDP. 1970. Mapa ecólogico de Panama. Map 1:5,000,000. Programa de las Naciones Unidas para el Desarrollo, Panama City, Panama
- Vásquez Morera, A. 1983. Soils. In D. H. Janzen, editor. Costa Rican natural history. Chicago: University of Chicago Press. ISBN: 0226393348
- Wille, C. 1999. Personal communication.
- Wilson, D. E. 1990. Mammals of La Selva, Costa Rica. In A. H. Gentry, editor, Four neotropical rainforests. New Haven: Yale Univesity Press. ISBN: 0300054483
- World Wildlife Fund-Central America. 1999. Concept Paper for a reconnaissance of the Talamancan Montane Forest Ecoregion in Costa Rica and Panama.
Disclaimer: This article contains some information that was originally published by the World Wildlife Fund. Topic editors and authors for the Encyclopedia of Earth have edited its content and added new information. The use of information from the World Wildlife Fund should not be construed as support for or endorsement by that organization for any new information added by EoE personnel, or for any editing of the original content.