United States

Department of

Agriculture

Forest Service

International Institute

of Tropical Forestry

General Technical

Report

IITF-GTR-39

March 2008

The Puerto Rico Gap

Analysis Project

Volume 1: Land Cover, Vertebrate Species

Distributions, and Land Stewardship

William A. Gould, Caryl Alarcón, Brick Fevold, Michael E. Jiménez,

Sebastián Martinuzzi, Gary Potts, Maya Quiñones, Mariano Solórzano,

and Eduardo Ventosa
The Forest Service of the U.S. Department of Agriculture is dedicated to the

principle of multiple use management of the Nation’s forest resources for sustained yields of wood, water, forage, wildlife, and recreation. Through forestry

research, cooperation with the States and private forest owners, and management of the national forests and national grasslands, it strives—as directed by

Congress—to provide increasingly greater service to a growing Nation.

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and where applicable, sex, marital status, familial status, parental status, religion,

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3272 (voice) or (202) 720-6382 (TDD). USDA is an equal opportunity provider and

employer.

Authors

William A. Gould is a research ecologist, Maya Quiñones and Michael E.

Jiménez are geographic information systems specialists, Mariano Solórzano is a

biologist, and Gary Potts is a remote sensing specialist, U.S. Department of

Agriculture, Forest Service, International Institute of Tropical Forestry, Río

Piedras, Puerto Rico 00926-1119. Caryl Alarcón is a geographer, National Park

Service, South Flordia National Resources Center, Homestead, FL 33030. Brick

Fevold is an environmental analysis and review specialist, Bureau of Integrated

Science Services, Wisconsin Department of Natural Resources, Madison, WI 53707.

Sebastián Martinuzzi is an ecologist, University of La Plata—LISEA and the

U.S. Department of Agriculture, Forest Service, International Institute of Tropical

Forestry, Río Piedras, PR 00926-1119. Eduardo Ventosa is an ecologist, Puerto

Rico Department of Natural and Environmental Resources, Río Piedras, Puerto

Rico.

Research performed under cooperative agreement no. 01HQPG0031

(01-IA-111201-002).
i

The Puerto Rico Gap Analysis Project

Volume 1: Land Cover, Vertebrate Species

Distributions, and Land Stewardship

William A. Gould, Caryl Alarcón, Brick Fevold, Michael E. Jiménez,

Sebastián Martinuzzi, Gary Potts, Maya Quiñones, Mariano Solórzano,

and Eduardo Ventosa

U.S. Department of Agriculture Forest Service,

International Institute of Tropical Forestry,

1201 Calle Ceiba, Río Piedras, Puerto Rico 00926-1119

U.S. Geological Survey, Biological Resources Division, Gap Analysis

Program

LIST OF PROJECT AFFILIATES

Puerto Rico Department of Natural and Environmental Resources

North Carolina State University

United States Geological Survey Biological Resources Division
Errata for IITF-GTR-39 

Figure captions on pages 5 and 58 have been reversed: 

Page 5 is the endemic Puerto Rican Tody (Todus mexicanus) and page 58 is the 

endemic Puerto Rican flycatcher (Myarchus antillarum). 

Citations omitted: 

Landcover mapping of Mona Island aided by the work of Cintrón and Rogers 

(1991). 

Landscape units (Figures 7 and 9) build on previous work by Figueroa-Colón 

(1996). 

References: 

Cintrón, B.; Rogers, L. 1991. Plant communities of Mona Island. Acta Científica. 

5:10-64. 

Figueroa-Colón, J.C. 1996. Geoclimatic regions of Puerto Rico (map). San Juan, 

Puerto Rico. U.S. Department of the Interior, Geological Survey Water Resources 

Division. 

Errata for IITF-GTR-39 

Figure captions on pages 5 and 58 have been reversed: 

Page 5 is the endemic Puerto Rican Tody (Todus mexicanus) and page 58 is the 

endemic Puerto Rican flycatcher (Myarchus antillarum). 

Citations omitted: 

Landcover mapping of Mona Island aided by the work of Cintrón and Rogers 

(1991). 

Landscape units (Figures 7 and 9) build on previous work by Figueroa-Colón 

(1996). 

References: 

Cintrón, B.; Rogers, L. 1991. Plant communities of Mona Island. Acta Científica. 

5:10-64. 

Figueroa-Colón, J.C. 1996. Geoclimatic regions of Puerto Rico (map). San Juan, 

Puerto Rico. U.S. Department of the Interior, Geological Survey Water Resources 

Division. 
ii

Abstract

Gould, William A.; Alarcón, Caryl; Fevold, Brick; Jiménez, Michael E.;

Martinuzzi, Sebastián; Potts, Gary; Quiñones, Maya; Solórzano, Mariano;

Ventosa, Eduardo. 2008. The Puerto Rico Gap Analysis Project. Volume 1:

Land cover, vertebrate species distributions, and land stewardship. Gen. Tech.

Rep. IITF-GTR-39. Río Piedras, PR: U.S. Department of Agriculture, Forest

Service, International Institute of Tropical Forestry. 165 p.

Puerto Rico faces a number of problems common to much of the world. Population

is increasing while land area is not, and there are reassessments of land use policy

and practice to accommodate growing populations, shifting economies, and changing public value systems. Puerto Rico shares similarities with the Eastern United

States with its history of agricultural abandonment, relatively high population

density, and abundance of “forested” suburban areas in the wildland-urban interface.

Puerto Rico has affinities with other tropical regions with high biodiversity and an

abundance of rare species. Puerto Rico shares similarities with small islands with

space limitations, an abrupt terrestrial-marine interface, and species diversity

controlled by the isolation of the island as well as climatic, evolutionary, and

historical factors. The Puerto Rico Gap Analysis Project documents Puerto Rico’s

land cover, vertebrate occurrences and natural history information, and land stewardship. The report has four major components: land cover mapping, documentation of vertebrate species distributions, documentation of land stewardship practices

with respect to conservation, and an integrated analysis of these three elements.

Our current reserve system protects a number of important habitats and species and

includes 7.6 percent of Puerto Rico. Expanding this to 15 percent would be more in

line with internationally accepted conservation goals. Abandoned agricultural land

serves as habitat for a number of species and buffers older forests, wetlands, riparian areas, and reserves. These lands have excellent potential for restoration. Recommendations include expanded reserves in the coastal plain, particularly coastal hills

and the matrix of wetland and upland vegetation; regulation of development in the

periphery of existing reserves; and developing viable corridors to connect the

upland and coastal reserves.

Keywords: Biodiversity, conservation, tropical ecology, vertebrate diversity,

land cover, Puerto Rico
iii

Summary

The Puerto Rico Gap Analysis Project (PRGAP) is a comprehensive assemblage of

information on Puerto Rico’s land cover, vertebrate occurrences and natural history

information, and land stewardship. It is based on methods developed by the national

Gap Analysis Program (GAP) to determine the degree to which animal species and

natural communities are represented in the current mix of conservation lands. Those

species or communities not well represented are considered conservation “gaps.”

The PRGAP provides geographic and ecological information on the status of not

only threatened or rare species, but the common species of Puerto Rico.

The PRGAP project has four major components: land cover mapping, documentation of vertebrate species distributions, documentation of land stewardship

practices with respect to conservation, and an integrated analysis of these three

elements.

Land Cover

We developed a land cover map of Puerto Rico using recent (1999–2003) satellite

imagery and information on climate, geology, topography, hydrology, and land use

history. We defined 70 land cover classes in a hierarchical classification scheme

based on whether the cover is natural vegetation, developed, or agricultural and on

whether the natural vegetation is closed forest, woodland, shrubland, or grassland.

Forest and grassland classes are further defined as dry, moist, wet, or flooded. These

units are then differentiated as occurring on soils derived from limestone, alluvial,

serpentine, or noncalcareous substrates. A number of forest types are further

classified as to the forest age (i.e., primary, mature secondary, or young secondary

forests). Wetlands are classified as forested or herbaceous, saline or nonsaline, and

seasonally flooded or emergent. Finally, where information is available, we present

the dominant plant communities and species representative of these land cover units.

We classified 53 percent of Puerto Rico as predominantly woody vegetation, 35

percent as grassland or herbaceous agriculture, 11 percent as developed land, and

about 1 percent each of water and natural barrens. Of the woody areas, low and

mid-elevation moist forests cover 26 percent, upper elevation wet forests cover 18

percent, dry forests cover 7 percent, and flooded mangrove and Pterocarpus forests

cover 1 percent of the island. Coastal wetlands cover less than 4 percent of the

island. Forty-two percent of the wetlands are saline and 58 percent are freshwater.

Mangroves and Pterocarpus swamps cover 1 percent of the island, 67 km2

 and 2.6
iv

km2, respectively. Seventy-four percent of the wetlands are dominated by herbaceous vegetation, and 92 percent of these are seasonally flooded. Of the herbaceous

wetlands, 77 percent are nonsaline and 23 percent are saline.

Vertebrates

Over 470 vertebrate species have been recorded in Puerto Rico and its adjacent

islands including terrestrial and aquatic birds, reptiles, amphibians, and mammals.

Of these, 426 are terrestrial vertebrate species. Many of these are migratory, wintering, accidental or vagrant species that do not breed regularly or at all on the island.

We have developed a database that contains taxonomic information, residence status,

and conservation status of all these species. We predicted the distributions of 98

bird, 47 reptile, 18 amphibian, and 14 mammal species including all native resident

endemic and endangered terrestrial vertebrates and some introduced species. Migratory birds, sea turtles, and marine and freshwater aquatic species will be included in

future analyses.

Species ranges were mapped by using a network of 24-km2

 hexagons that cover

Puerto Rico and its adjacent islands. Each hexagon was attributed with the species

probability of occurrence in one of eight categories. Species probability of occurrence information is derived from published literature, unpublished data sets,

museum records, and expert opinion.

Species distributions were mapped by identifying predicted habitat within the

species range based on literature and expert review. The resulting maps of predicted

species distribution are a result of the integration of information from both the

vertebrate database and land cover mapping. We combined species distribution

information to develop species richness maps. The resulting biodiversity patterns

indicate that forested parts of the landscape are the habitats with the highest predicted species richness. Urban and barren areas are the habitats with the lowest

species richness. Individual taxonomic groups show distinct patterns.

We also looked at the species richness within the network of 24-km2

 hexagons

used to document species occurrences. This analysis indicates that the highest levels

of habitat heterogeneity and resulting biodiversity are in coastal areas with a mix of

wetlands, grassland, and forested coastal hills. The coastal area is also extremely

vulnerable to development, as the topography is less steep, it is close to urban areas

and existing infrastructure, and nonwetlands on the coastal plain and coastal hills

are primarily unprotected. Development is prohibited in the wetlands, but development adjacent to wetlands can destroy the diverse matrix of habitats and affect

hydrologic patterns, altering species composition and biodiversity.
v

Land Stewardship

The national GAP currently uses a scale of 1 to 4 to denote relative degree of maintenance of biodiversity for stewardship areas. A status of “1” denotes the highest,

most permanent level of maintenance, and “4” represents the lowest level of biodiversity management, or unknown status (Crist et al. 1995, Edwards et al. 1995,

Scott et al. 1993).

Although land stewardship, management, and land use are very dynamic, we

have identified 77 stewardship areas that receive some management for conservation

(GAP status 1 through 3). Land ownership of these areas is shared among 20

organizations, with the Puerto Rico Department of Natural and Environmental

Resources (DNER) being the primary landowner. Management of land stewardship

areas is shared among 20 organizations, with the DNER, the U.S. Forest Service,

and the U.S. Fish and Wildlife Service being the primary governmental land

managers and the Conservation Trust of Puerto Rico being the primary nongovernmental land manager.

Of the total land area of Puerto Rico, 7.6 percent receives some management

for conservation (GAP status 1, 2, or 3) with 7.4 percent of the total land area

receiving good management of conservation (GAP status 1 or 2). Fifty-nine percent

of the stewardship areas are managed by Commonwealth agencies, 30 percent by

federal agencies, and 11 percent by nongovernmental or private agencies.

Gap Analyses–Land Cover

Eight of our 70 land cover classes have less than 1 percent of their area represented

in GAP status 1 or 2 conservation areas; they cover 43 percent of the island. They

are primarily subject to human use such as agriculture, housing, and other development. Moist grasslands and pastures cover nearly one quarter of the island and are

primarily active pasture and abandoned agricultural land. Given the resilience of the

natural vegetation in Puerto Rico, this land cover type has potential for management

for reforestation or as natural grasslands and open space.

Twenty-seven land cover units have between 1 and 10 percent of their area

represented in GAP status 1 or 2 conservation areas. They account for 44 percent of

the island. They range from an extent of less than 1 percent to over 6 percent of the

island and include a number of young secondary forest and woodland land cover

classes, as well as artificial and natural barrens, active and abandoned shade coffee

plantations, dry grasslands and pastures, riparian forests, and four mature secondary

forest classes.
vi

Four land unit classes have between 10 and 20 percent of their area represented

in GAP status 1 or 2 conservation areas. They account for 1.7 percent of the island

and include two woodland-shrubland classes that typically occur on abandoned

agricultural land, dryland riparian forest, and palm plantations.

Fourteen land cover classes have between 20 and 50 percent of their area represented in GAP status 1 or 2 conservation areas and account for 6.1 percent of the

island’s total area. They include a number of ecologically important areas including

beaches and shorelines, mature forests, wetlands, mangrove complexes, and Sierra

palm forest.

Seventeen land cover units are over 50 percent protected under GAP status

1 and 2. They account for 5.1 percent of the island. They include important

primary and mature secondary forest types in the Luquillo Mountains, freshwater

Pterocarpus swamps, forests on serpentine substrates, and a number of dryland

habitats unique to Mona Island and the Guánica Biosphere Reserve.

Gap Analyses–Vertebrates

Four species have less than 1 percent of their habitat protected under GAP status

1 or 2. These include two species of Gecko common in urban areas; one bird,

Carduelis cucullata, which is nonnative; and Eleutherodactylus cooki, the guajón

or rock coqui, which has limited habitat, none of which is protected.

One recently discovered species not fully included in the PRGAP analysis, is

the coqui llanero, or plains coqui (Eleutherodactylus juanariveroii) (Ríos-López and

Thomas 2007).

Seventy-seven species have 1 percent to less than 10 percent of their habitat

protected under GAP status 1 or 2. Many are widespread although not necessarily

common and occur in disturbed habitats. A few, such as the blind snake Typhlops

platycephalus, have limited habitat (15 percent of the island), the majority unprotected (98 percent of its habitat).

Thirty-two species have 10 percent to less than 20 percent of their habitat

protected under GAP status 1 or 2. These species are a mix of those with widespread and those with limited habitat extent.

Forty-three species have 20 percent to less than 50 percent of their habitat

protected under GAP status 1 or 2. All of these species have habitat extent limited to

less than 11 percent of the island. A number of endangered species are in this group,

and many are limited to less extensive habitats such as saline and freshwater ponds

and wetlands or high mountain areas.
vii

Twenty-one species have at least 50 percent of their predicted habitat protected

under GAP status 1 or 2. These include a number of species found only on forest

reserves or particular protected satellite islands (Mona and Desecheo). All of these

species have very limited habitat and none exceed 2 percent of the island.

Forty-seven species are listed as either federally threatened or endangered or

given partial status, or are locally listed by the DNER as vulnerable, endangered,

critically endangered, or data deficient. The extent of habitat for 70 percent of these

species is typically below 5 percent of the island’s total area. Eighty-three percent

of the species have a habitat extent below 20 percent of the island’s total area.

Eleutherodactylus cooki, the guajón or rock coqui, is the least protected, with no

protected habitat. Ten species have less than 10 percent of their habitat protected

and 18 species have less than 20 percent of their habitat protected. Five species are

found only in reserves with 100 percent of their current distribution protected.

Distributions for these species could be expanded outside reserves if suitable habitat

is protected or restored and species reintroductions are encouraged.

Puerto Rico is at a crossroads in terms of land use transition, as much of what

was formerly agricultural land is now experiencing more intense, and possibly

irreversible urban-associated development. Our current reserve system is well

located and protects a number of important habitats and species. However, this

system needs to be expanded from 7.6 percent to at least 15 percent of the island’s

area to be more in line with internationally accepted conservation goals. Our

abandoned agricultural land is often a matrix of forested and open green space that

serves as habitat for a number of species and buffers older forests, wetlands, riparian areas, and our current reserves. These lands have excellent potential for restoration. We recommend expanded reserves in the coastal plain, particularly coastal hills

and the matrix of wetland and upland vegetation; better regulation of development

in the periphery of existing reserves to maintain the integrity of hydrologic systems

in wetlands; and protecting viable corridors and buffer zones to connect the upland

and coastal reserves.
viii

Resumen

El Proyecto Gap de Puerto Rico (Puerto Rico Gap Analysis Project—PRGAP) es

una colección comprensiva de información sobre la cubierta del suelo de Puerto

Rico, distribución e historia natural de vertebrados, y áreas de manejo. Está basado

en la metodología desarrollada por el programa nacional GAP de los Estados

Unidos para determinar el grado en el cual especies y sus comunidades naturales

están representadas en los terrenos que están actualmente protegidos. Las especies o

comunidades que no estén bien representadas son consideradas “gaps” o agujeros en

el plan de conservación. El PRGAP provee información geográfica y ecológica en

el estatus de, no solo las especies endémicas o en peligro de extinción, sino de todas

las especies de Puerto Rico, incluyendo las comunes y exóticas.

El PRGAP tiene cuatro componentes principales: mapeo de la cobertura del

terreno, documentación de las distribuciones de especies vertebradas,

documentación de las prácticas de conservación en áreas de manejo, y un análisis

integrando estos tres elementos.

Cobertura del Terreno

Desarrollamos un mapa de cobertura del terreno de Puerto Rico utilizando imágenes

de satélite recientes (1999–2003) e información sobre clima, geología, topografía,

hidrología, e historia del uso del terreno. Definimos 70 clases de cobertura del

terreno en un esquema jerárquico de clasificación basado en las siguientes

coberturas básicas: vegetación natural, desarrollo urbano, o agricultura. La

vegetación natural se clasificó en bosque cerrado, bosque abierto, arbustos o

pastizales. Las clasificaciones de bosque cerrado y pastizales son definidas más a

fondo en seco, húmedo, mojado o inundado.

Estas unidades son entonces diferenciadas entre si ocurren en suelos derivados

de caliza, aluvial, serpentina o substratos no calcáreos. Varios de los tipos de bosque

cerrado fueron clasificados de acuerdo a su edad (por ejemplo primario, secundario

maduro, o joven secundario). Los manglares fueron clasificados en arbolado,

herbáceo, salino o no salino, y temporalmente inundado o emergente. Finalmente,

información sobre comunidades de flora dominante, y especies representativas de la

unidad de cobertura de terreno, fue incorporada cuando ésta estaba disponible.

Clasificamos el 53 por ciento de Puerto Rico como predominantemente

vegetación leñosa, y 35 por ciento como pastizales o agricultura herbácea.

Clasificamos 11 por ciento como terrenos desarrollados y alrededor de 1 por ciento

como agua y terreno descubierto natural. De las 70 unidades de clasificación 49
ix

están dominadas por vegetación leñosa. De éstas, 26 por ciento cubren bosque

húmedo de baja y mediana elevación, 18 por ciento cubre bosque mojado de alta

elevación, 7 por ciento cubre bosque seco, y 1 por ciento cubre manglar inundado y

bosques de Pterocarpus. Los humedales costeros cubren menos del 4 por ciento de

la isla y el 42 por ciento de los mismos son salinos y el 58 por ciento son humedales

de agua dulce. Los humedales costeros-arbóreos cubren 1 por ciento de la isla y

están representados por manglares y pantanos Pterocarpus de agua dulce. Por otro

lado, el 74 por ciento de los humedales están dominados por vegetación herbácea, y

92 por ciento de estos son temporalmente inundados. De los humedales herbáceos,

77 por ciento son no-salinos, y 23 por ciento son salinos.

Vertebrados

Datos sobre 470 especies de vertebrados han sido recolectados para Puerto Rico y

sus islas adyacentes incluyendo 98 aves terrestres y acuáticas, 47 reptiles, 18

anfibios y 14 mamíferos. De estas, 426 son vertebrados terrestres y muchas son

especies migratorias, accidentales que no se reproducen en la isla, o solo lo hacen en

raras ocasiones. Desarrollamos una base de datos que contiene información

taxonómica, estatus de residencia, y estatus de conservación de todas las especies

registradas. Se crearon predicciones de distribución de un subconjunto de especies

incluyendo todas las especies nativas residentes y endémicas terrestres y algunas

especies exóticas. Esperamos incluir las aves migratorias, las tortugas marinas, las

especies acuáticas marinas o de agua dulce, y los animales domésticos en análisis

futuros.

Las distribuciones conocidas de las especies fueron mapeadas utilizando una red

de hexágonos, de 24 km2 cada uno, que cubre a Puerto Rico e islas adyacentes. Cada

hexágono fue clasificado en una de ocho categorías de probabilidad de presencia.

Información sobre la probabilidad de presencia de una especie es derivada de

literatura publicada, bases de datos no publicadas, registros de museos, y opinión

experta.

Mapeamos las distribuciones de hábitat de las especies estableciendo una

predicción de sus hábitats dentro de su distribución geográfica, basada en la

literatura científica y la revisión de expertos. Por lo tanto, los 177 mapas de

distribuciones predichas de las especies fueron el resultado de la integración de

información de la base de datos de vertebrados y el mapa de cobertura de terreno.

Desarrollamos mapas de riqueza de especies usando esta información para

identificar patrones de biodiversidad. Los patrones de biodiversidad resultantes

indican que las áreas boscosas del paisaje son las más ricas en diversidad de
x

especies, con hasta 73 especies. Las áreas urbanas y las áreas estériles muestran la

menor riqueza con tan pocas como 7 especies. Los distintos grupos taxonómicos

demuestran patrones distintos de distribución.

También examinamos la riqueza de las especies dentro de la misma red de

hexágonos, de 24 km2

 cada uno, usada para documentar la presencia de especies.

Este análisis indica que los niveles más altos de diversidad de hábitat y resultante

biodiversidad se encuentran en áreas costeras que contienen una mezcla de

humedales, pastizales y lomas costeras boscosas. El área costera es extremadamente

vulnerable al desarrollo urbano debido a la topografía menos montañosa del lugar.

El desarrollo urbano está generalmente prohibido en los humedales, pero el

desarrollo en áreas adyacentes puede destruir la matriz de diversidad del hábitat y

afectar los patrones hidrológicos. Esto a su vez altera la composición de especies y

la biodiversidad.

Áreas de Manejo

El GAP Nacional actualmente usa una escala de 1 al 4 para denominar el grado

relativo de mantenimiento de biodiversidad en áreas con planes de manejo

existentes. Un estatus de “1” significa el más alto y permanente nivel de

mantenimiento, y 4 representa el nivel más bajo de manejo de la biodiversidad, o la

falta de información sobre el área.

Aunque la tenencia, el manejo y el uso de los terrenos son bastante dinámicos

en Puerto Rico identificamos un grupo de 77 áreas de manejo que reciben algún tipo

de manejo para la conservación (estatus de GAP del 1 al 3), bajo la tenencia de unas

20 organizaciones con el DRNA siendo el propietario principal. El manejo de estas

áreas es compartido entre 20 organizaciones con el Departamento de Recursos

Naturales y Ambientales, El Servicio Forestal Federal de los Estados Unidos y el

Servicio de Pesca y Vida Silvestre de los Estados Unidos siendo los principales

manejadores gubernamentales, y el Fideicomiso de Conservación de Puerto Rico

siendo el principal manejador no gubernamental. Siete punto seis por ciento de

todas las tierras en Puerto Rico recibe algún tipo de manejo para la conservación

(estatus de GAP 1, 2 o 3), con un 7.4 por ciento recibiendo buen manejo (estatus de

GAP 1 o 2). Cincuenta y nueve por ciento de las áreas de manejo están manejadas

por agencias del Estado Libre Asociado de Puerto Rico, 30 por ciento por agencias

federales, y 11 por ciento por agencias privadas o no gubernamentales.
xi

Análisis “Gap”–Cobertura de Terreno

Ocho de las 70 clases de cobertura de terreno tienen menos de 1 por ciento de su

área representada en tierras con estatus de conservación de GAP 1 o 2; estas cubren

el 43 por ciento de la isla. Dada la perseverancia de la vegetación natural en Puerto

Rico, este tipo de cobertura tiene el potencial de manejo para la reforestación o

como pastos naturales y espacio abierto.

Veinte y siete unidades de cobertura de terreno tienen entre 1 a 10 por ciento de

su área representada en tierras con estatus de conservación GAP 1 o 2. Éstas cubren

44 por ciento de la superficie de la isla y contienen varias clases de bosques

secundarios jóvenes, bosques abiertos terrenos expuestos naturales y artificiales,

plantaciones de café de sombra activas y abandonadas, pastizales secos, bosques

ribereños, y cuatro clases de bosque secundario maduro.

Cuatro unidades de cobertura de terreno tienen entre 10 a 20 por ciento de su

área representada en tierras con estatus de conservación de GAP 1 o 2, y representan

el 1.7 por ciento de la isla. Éstas incluyen dos clases de bosque abierto y arbustos—

los cuales suelen ocurrir en tierras agrícolas abandonadas, bosque ribereño seco, y

plantaciones de palmas.

Catorce clases de cobertura de terreno tienen entre 20 a 50 por ciento de su área

representada por los estatus de conservación de áreas Gap 1 o 2 y representan el 6.1

por ciento del área total de la isla. Estos incluyen un número de áreas

ecológicamente importantes tales como playas, orilla costera, bosques maduros,

humedales, complejos de manglares y bosque de Palma de Sierra. Diecisiete

unidades de cobertura de terreno están sobre el 50 por ciento de protección bajo los

estatus GAP 1 o 2. Estos representan un 5.1 por ciento de la isla. Éstas incluyen

importantes bosques de tipo primario y secundario en la Cordillera de Luquillo,

pantanos de agua de dulce de Pterocarpus, bosques de suelos serpentinos, y varias

áreas de hábitat seco únicos de la Isla de Mona y de la Reserva de la Biosfera de

Guánica.

Análisis “Gap”–Vertebrados

Cuatro especies tienen menos del 1 por ciento de su hábitat protegido bajo el estatus

de conservación de áreas GAP 1 o 2.

Una especie descubierta recientemente, provisionalmente reconocida, y no

incluida todavía en el análisis PRGAP, es el coquí llanero (Eleutherodactylus

juanariveroii). La distribución de este coquí es muy limitada y no se encuentra bajo

ninguna protección.
xii

Setenta y siete especies cuentan con 1 a 10 por ciento de su hábitat protegido

bajo los estatus GAP 1 o 2. Muchas de estas especies tienen una distribución amplia

a lo largo de la isla aunque no son necesariamente comunes y ocurren en hábitat

perturbado.

Treinta y dos especies tienen 10 a 20 por ciento de su hábitat protegido bajo los

estatus de GAP 1 o 2. Estas especies se podrían describir como un balance entre

aquellas de amplia distribución y aquellas de distribución limitada.

Cuarenta y tres especies cuentan con 20 a 50 por ciento de su hábitat protegido

bajo el estatus de conservación de áreas GAP 1 o 2. Muchas están limitadas a

sistemas naturales de menor extensión como charcas de agua dulce o salada,

humedales o áreas montañosas elevadas.

Veintiuna especies tienen al menos 50 por ciento de su hábitat predicho

protegido bajo el estatus de conservación de áreas GAP 1 o 2. Esto incluye varias

especies que se encuentran solamente en reservas forestales o en islas protegidas

(Mona y Desecheo). Todas estas especies cuentan con una extensión de hábitat muy

limitado que no supera en ninguno de los casos el dos por ciento de la isla.

Cuarenta y siete especies están federalmente listadas ya sea como amenazadas,

en peligro o de estatus parcial, o localmente listadas por el DRNA como

vulnerables, amenazadas, en peligro crítico, o con insuficiencia de datos. La

extensión de hábitat para el 70 por ciento de estas especies está típicamente bajo el 5

por ciento del área total de la isla. Eleutherodactylus cooki, el coquí guajón, es la

menos protegida pues no cuenta con hábitat protegido. Diez especies cuentan con

menos del 10 por ciento de su hábitat protegido y 18 tienen menos del 20 por

ciento.

Puerto Rico se encuentra entre la espada y la pared en términos de transición del

uso de terrenos ya que las tierras, que en el pasado estaban destinadas a la

agricultura, hoy en día están experimentando un uso más intenso, posiblemente

irreversible, relacionado al desarrollo urbano. Nuestro sistema de reservas actual

está bien ubicado y protege a variedad de especies y ecosistemas. Sin embargo, este

sistema necesita expandirse de menos de un ocho por ciento a al menos un 15 por

ciento de la superficie de la isla para estar más de acuerdo con las metas de

conservación aceptadas a nivel internacional. Las tierras agrícolas abandonadas se

han ido transformando en una miscelánea de espacios verdes forestados y abiertos

que sirve como hábitat para muchas especies y amortiguan a los bosques maduros,
xiii

humedales, zonas ribereñas, y a las reservas actuales de los efectos antropogénicos.

Estas tierras tienen un potencial magnífico de restauración. Recomendamos la

expansión de reservas en las planicies costeras, particularmente las colinas costeras y

la matriz de humedales y vegetación elevada.

También recomendamos una mejor regulación de los desarrollos urbanos en la

periferia de las reservas existentes para mantener la integridad de los sistemas

hidrológicos en los humedales. Finalmente recomendamos el establecimiento y

mantenimiento de corredores viables y zonas de amortiguamiento que conecten las

tierras altas con las reservas costeras.
xiv

Contents

1 Introduction

2 The Gap Analysis Program Mission

4 The Gap Analysis Concept

7 General Limitations

8 Study Area

15 Land Cover Classification and Mapping

17 Land Classification Methods

19 Land Cover Classification Results

24 Land Cover Classification Accuracy Assessment

29 Land Cover Classification Limitations

30 Predicted Vertebrate Distributions and Species Richness

31 Vertebrate Mapping Methods

42 Vertebrate Distribution Results

52 Vertebrate Richness

57 Vertebrate Distribution Accuracy Assessment

59 Vertebrate Data Limitations

59 Land Stewardship

61 Mapping Standards

62 Stewardship Mapping Methods

64 Stewardship Mapping Results

64 Stewardship Map Limitations

75 Analysis Based on Stewardship and Management Status

76 Analysis Methods

76 Analysis Results

98 Analysis Limitations

98 Conclusions and Management Implications

102 Concluciones e Implicaciones de Manejo

105 Product Use and Availability

105 How to Obtain the Products

105 Disclaimer

106 Metadata

106 Appropriate Use of These Data

109 Acknowledgments

111 Agradecimientos
xv

112 English Equivalents

113 Literature Cited

124 Appendix 1: Example GAP Applications

129 Appendix 2: Terrestrial Vertebrate Species of Puerto Rico

143 Appendix 3: Land Cover Vegetation Classification

150 Appendix 4: Puerto Rico Land Cover Descriptions

156 Glossary

161 Glossary of Acronyms

Mariano Solórzano

Tanama River
This Page Left Blank Intentionally
1

The Puerto Rico Gap Analysis Project

Introduction

The Puerto Rico Gap Analysis Project (PRGAP) is a comprehensive assemblage

of information on Puerto Rico’s land cover, vertebrate occurrences and natural

history information, and land stewardship. It is based on methods developed by

the national Gap Analysis Program (GAP) to determine the degree to which animal

species and natural communities are represented in the current mix of conservation

lands. Those species or communities not well represented are considered conservation “gaps.” The PRGAP provides geographic and ecological information on the

status of not only threatened or rare species, but the common species of Puerto

Rico.

The current status of the PRGAP is summarized in a series of four volumes.

Volume 1: Land cover, vertebrate species distributions, and land stewardship,

provides an overview and analysis of the primary data sets used in gap analysis.

Volume 2: Species accounts, provides detailed taxonomic, natural history, conservation status, and bibliographic information on all vertebrate species addressed in this

report.

Volume 3: Species occurrences, provides range maps with 24-km2

 resolution for

each species based on documented occurrences.

Volume 4: Species predicted distributions, provides species distribution maps with

15-m2

 resolution for each species based on documented occurrences and presence

of suitable habitat.

Volume 1 begins with an overview of the Gap Analysis Program (GAP) mission, concept, and limitations. This is followed by a description of the key factors

controlling biodiversity, including the climate, physiography, ecology, and land use

history of Puerto Rico. Three subsequent sections describe the primary data layers

associated with a gap analysis: land cover, vertebrate species distributions, and land

stewardship. Each of these sections describes the methods, results, and discussions

of the related data. A fourth section includes an analysis of the distribution of species and habitats with respect to conservation management (i.e., a “gap” analysis).

Finally, this volume describes the management implications of the analysis results

and provides information on how to acquire and use the data.
2

GENERAL TECHNICAL REPORT IITF-GTR-39

The Gap Analysis Program Mission

The mission of GAP is to prevent conservation crises by providing conservation

assessments of plant communities and native animal species and to facilitate the

application of this information to land management activities. This is accomplished

by meeting the following five objectives:

1. Map current land cover.

2. Map the predicted distribution of selected terrestrial vertebrates.

3. Document the representation of land cover types and animal species in areas

managed for the long-term maintenance of biodiversity.

4. Make GAP project information available to the public and those charged with

land use research, policy, planning, and management.

5. Build institutional cooperation in the application of this information to state and

regional management activities.

To meet these objectives, it is necessary that GAP operate at the state, commonwealth, or regional level while maintaining consistency with national standards.

Participation by a wide variety of cooperators is necessary and can lead to understanding and acceptance of the data and the development of relationships that will

lead to cooperative conservation planning.

State Objectives—

The Puerto Rico GAP had a number of additional objectives. We adapted methodology of the national GAP to meet the needs of Puerto Rico, an archipelago of

tropical islands with a unique social, economic, and ecological environment. Puerto

Rico’s environment includes diverse bioclimatic zones, considerable ecological

variation over small distances, a high degree of development pressure, a small

national heritage database of species occurrences, and a long history of ecological

research but few published descriptions of natural vegetation at the plant community

level. Most land management for conservation is being done by the Puerto Rico

Department of Natural and Environmental Resources (DNER) followed by the U.S.

Department of Agriculture Forest Service (USFS) and the U.S. Fish and Wildlife

Service (USFWS).

Our state-level objectives were to:

• Develop a coverage of 24-km2

 hexagons (fig. 1) for use in occurrence and

range mapping that accommodates the scale of landscape variation in

Puerto Rico (Gould et al. 2008b).
3

The Puerto Rico Gap Analysis Project Figure 1—The Puerto Rico Gap Analysis Project has adapted a hexagon grid network for use in mapping Puerto Rico’s biological diversity. This hexagon grid (PRGAP-HEX) provides a uniform unit of area that can be used to represent the range and occurrence of vertebrate species across a very heterogeneous landscape. The PRGAP-HEX grid consists of 483 individual hexagons with 305 occurring only over land, 161 over coastal areas, and 17 over open marine areas with small reefs and cays. Each hexagon includes 24 km2. The PRGAP-HEX is a modification of the Forest Inventory and Analysis grid developed by the U.S. Forest Service (Gould et al. 2008b).
4

GENERAL TECHNICAL REPORT IITF-GTR-39

• Develop contacts and collaborations with conservation agencies and groups

in Puerto Rico in order to receive input and communicate our findings in

ways useful to immediate conservation concerns.

• Develop databases of species occurrences, scientific literature related to

species habitats and distributions, and species natural history information

that are dynamic and will have a useful life beyond the completion of

PRGAP.

• Compile and develop new information on natural vegetation at the plant

community level, and crosswalk these descriptions with past vegetation

work in Puerto Rico, using federal guidelines (FGDC 1997) and other

descriptive hierarchies.

• Collaborate in all phases of the project with the primary land management

agency in Puerto Rico, the DNER.

• Develop a project that can serve as a foundation for the development of a

Puerto Rico-United States Virgin Islands GAP and Caribbean GAP.

• Develop a bilingual (Spanish and English) project.

The Gap Analysis Concept

The Gap Analysis Program brings together the problem-solving capabilities of

federal, state, and private scientists to tackle the difficult issues of land cover mapping, animal habitat characterization, and biodiversity conservation assessment at

the state, regional, national, and international levels. The program seeks to facilitate

cooperative development and use of information.

Much of the following discussion was taken verbatim from Davis et al. 1995,

Edwards et al. 1995, and Scott et al. 1993. The gap analysis process provides an

overview of the distribution and conservation status of several components of

biodiversity. It uses the distribution of actual vegetation and predicted distribution of terrestrial vertebrates and, when available, invertebrate taxa. Digital map

overlays in a geographic information system (GIS) are used to identify individual

species, species-rich areas, and vegetation types that are unrepresented or underrepresented in existing management areas. It functions as a preliminary step to

the more detailed studies needed to establish actual boundaries for planning and

management of biological resources on the ground. These data and results are then

made available to the public so that institutions as well as individual landowners

and managers may become more effective stewards through more complete knowledge of the management status of these elements of biodiversity. The GAP, by

focusing on higher levels of biological organization, is likely to be both cheaper
5

The Puerto Rico Gap Analysis Project

The endemic Puerto Rican flycatcher Myarchus antillarum is known locally as the Juí de Puerto

Rico.

Tomás A. Carlo
6

GENERAL TECHNICAL REPORT IITF-GTR-39

and more likely to succeed than conservation programs focused on single species

or populations (Scott et al. 1993).

Biodiversity inventories can be visualized as “filters” designed to capture elements of biodiversity at various levels of organization. The filter concept has been

applied by The Nature Conservancy, which established Natural Heritage Programs

in all 50 states and the Commonwealth of Puerto Rico. The Nature Conservancy

employs a fine filter of rare species inventory and protection and a coarse filter of

community inventory and protection (Jenkins 1985, Noss 1987). It is postulated

that 85 to 90 percent of species can be protected by the coarse filter without having

to inventory or plan reserves for those species individually. A fine filter is then

applied to the remaining 15 to 10 percent of species to ensure their protection. Gap

analysis is a coarse-filter method because it can be used to quickly and cheaply

assess the other 85 to 90 percent of species. The GAP is not designed to identify

and aid protection of elements that are rare or of very restricted distribution; rather

it is designed to help “keep common species common” by identifying risk far in

advance of actual population decline.

The intuitively appealing idea of conserving most biodiversity by maintaining

examples of all natural community types is now widely recognized, and numerous

approaches to the spatial identification of biodiversity in association with natural

community type have been described (Kirkpatrick 1983, Margules et al. 1988,

Nicholls and Margules 1993, Pressey and Nicholls 1989). Furthermore, the spatial

scales at which organisms use the environment differ tremendously among species

and relate to body size, food habits, mobility, and other factors. Hence, no coarse

filter will be a complete assessment of biodiversity protection status and needs.

However, species that fall through the pores of the coarse filter, such as narrow

endemics and wide-ranging mammals, can be captured by the safety net of the fine

filter. Community-level (coarse-filter) protection is a complement to, not a substitute for, protection of individual rare species.

Gap analysis is essentially an expanded coarse-filter approach (Noss 1987) to

biodiversity protection. The land cover types mapped in GAP serve directly as a

coarse filter, the goal being to assure adequate representation of all native vegetation community types in biodiversity management areas. Landscapes with great

vegetation diversity often are those with high edaphic (soil or substrate) variety

or topographic relief. When elevational diversity is very great, a nearly complete

spectrum of vegetation types known from a biological region may occur within a

relatively small area. Such areas provide habitat for many species, including those

that depend on multiple habitat types to meet life history needs (Diamond 1986,
7

The Puerto Rico Gap Analysis Project

Noss 1987). By using landscape-sized samples (Forman and Godron 1986) as an

expanded coarse filter, gap analysis searches for and identifies biological regions

where unprotected or underrepresented vegetation types and animal species occur.

More detailed analyses were not part of this project, but are areas of research

that both PRGAP and GAP as a national program is pursuing. For example, a

second filter could combine species distribution information to identify a set of

areas in which all, or nearly all, mapped species are represented. There is a major

difference between identifying the richest areas in a region (many of which are

likely to be neighbors and share essentially the same list of species) and identifying

areas in which all species are represented. The latter task is most efficiently accomplished by selecting areas whose species lists are most different or complementary.

Areas with different environments tend to also have the most different species lists

for a variety of taxa. As a result, a set of areas with complementary sets of species

for one higher taxon (e.g., mammals) often will also do a good job representing

most species of other higher taxa (e.g., trees, butterflies). Species with large home

ranges, such as large carnivores, or species with very local distributions may

require individual attention. Additional data layers can be used for a more holistic

conservation evaluation. These include indicators of stress or risk (e.g., human

population growth, road density, rate of habitat fragmentation, distribution of

pollutants) and the locations of habitat corridors between wildlands that allow for

natural movement of wide-ranging animals and the migration of species in response

to climate change.

General Limitations

The following are general project limitations; specific limitations for the data are

described in the respective sections:

1. The GAP data are derived from remote sensing and modeling to make general

assessments about conservation status. Any decisions based on the data must be

supported by ground-truthing and more detailed analyses.

2. The GAP is not a substitute for threatened and endangered species listing and

recovery efforts. A primary argument in favor of gap analysis is that it is proactive: it seeks to recognize and manage sites of high biodiversity value for the

long-term maintenance of populations of native species and communities before

they become critically rare. Thus, it should help to reduce the rate at which

species require listing as threatened or endangered. Those species that are

already greatly imperiled, however, still require individual efforts to assure

their recovery.
8

GENERAL TECHNICAL REPORT IITF-GTR-39

3. The GAP data products and assessments represent a snapshot in time generally

representing the date of the satellite imagery. Updates are planned on a 5- to

10-year cycle, but users of the data must be aware of the static nature of the

products.

4. The GAP is not a substitute for a thorough national biological inventory. As a

response to rapid habitat loss, gap analysis provides a quick assessment of the

distribution of vegetation and associated species before they are lost, and provides focus and direction for local, regional, and national efforts to maintain

biodiversity. The process of improving knowledge in systematics, taxonomy,

and species distributions is lengthy and expensive. That process must be continued and expedited, however, in order to provide the detailed information needed

for a comprehensive assessment of our Nation’s biodiversity. Vegetation and

species distribution maps developed for GAP can be used to make such surveys

more cost effective by stratifying sampling areas according to expected variation

in biological attributes.

Study Area

A brief description of Puerto Rico—

The Caribbean has a number of large and small islands fostering endemic species

and a tropical climate that promotes high species diversity. It is considered a global

biodiversity hotspot (Myers et al. 2000). Puerto Rico, located at the junction of the

Greater and Lesser Antilles in the Caribbean basin, is in the center of this region.

The archipelago of the Commonwealth of Puerto Rico is made up of the main

island of Puerto Rico (about 160 km long by 50 km wide), the islands of Culebra

and Vieques to the east, the islands of Mona, Monito, and Desecheo to the west, and

a number of smaller cays. Puerto Rico has 9000 km2

 of terrestrial land surface. The

dominant physiographic features of the island include the Central Mountain Range

running east-west, a region of karst hills in the northwest, and the Luquillo Mountains of the northeast. Fifty-three percent of the island is mountainous, 25 percent

plains, 20 percent hills, 1 percent plateau, and 1 percent lakes and rivers (fig. 2).

The topography of the island has a strong control on the climate, with the wetter

regions on the windward, northern side of the mountains, and drier climate in the

leeward rain shadow. Six life zones (sensu Holdridge 1967) have been described

for Puerto Rico (fig. 3) including subtropical dry, lowland moist, subtropical wet,

lower montane wet, subtropical rain, and lower montane rain forest zones (Ewel
9

The Puerto Rico Gap Analysis Project

Figure 2—Physiography of Puerto Rico (Gould et al. 2008a).

Figure 3—Subtropical life zones of Puerto Rico (modified from Ewel and Whitmore 1973).

and Whitmore 1973). Mean annual rainfall ranges from below 900 mm in the

subtropical dry life zone to over 4000 mm in the subtropical wet rain forest. Mean

annual temperatures exhibit a narrow range from 22 to 25 oC with temperatures

decreasing with elevation (Daly et al. 2003). Seasonality in rainfall is most pronounced in the subtropical dry life zone. Peak rainfall throughout the island is found

in April and May and October to December (Daly et al. 2003).
10

GENERAL TECHNICAL REPORT IITF-GTR-39

Puerto Rico is experiencing rapid urban expansion, increasing population

pressures, and dynamic land cover change (Aide and Grau 2004, Birdsey and

Weaver 1982, Chinea 2002, Grau et al. 2003, Helmer 2004, Helmer et al. 2002,

Lopez et al. 2001, Lugo and Helmer 2004, Martinuzzi et al. 2006, Rivera and Aide

1998). In the 19th century, most of Puerto Rico was deforested and converted to

agricultural use, with 6 percent remaining forested and another 6 percent as shade

coffee plantations (Franco et al. 1997). Toward the latter part of the 20 th century,

forested area increased in Puerto Rico to 35 to 42 percent, primarily through a

transition from an agrarian to an industrialized society (Birdsey and Weaver 1987,

Franco et al. 1997, Helmer et al. 2002). Agricultural abandonment and development continue to increase, and our mapping efforts indicate current woody vegetation cover of over 50 percent. Levels of development and urbanization include 11

percent developed land surface with a high level of urban sprawl (figs. 4 and 5),

16 percent urban-use area, and 36 percent densely populated rural or suburban

area (fig. 6) (Martinuzzi et al. 2007a). This dynamic landscape, in which 4 million

people live and in which conservation management decisions must be made, has

parallels to temperate landscapes where urbanization encroaches on regenerated

forests, and tropical landscapes where urbanization is beginning to outpace agricultural deforestation.

Puerto Rico has a range of forest types whose composition is controlled by four

key elements: climatic gradients, substrate differences, topographic patterns, and

human and natural disturbance (Britton and Wilson 1924, Dansereau 1966, Gould

et al. 2006, Lugo 2005, Weaver 1991). Climatic gradients are related to an elevational rise from sea level up to 1340 m on the central cordillera of the island, the

northeasterly trade winds, and the rain shadow effect of the mountains (fig. 7). The

climatic control of dry, moist, and wet forest types in Puerto Rico represents the

Figure 4—Developed or built-up areas of Puerto Rico (in red) include over 11 percent of the area (Martinuzzi et al. 2007a).
11

The Puerto Rico Gap Analysis Project

Figure 5—Developed or built-up areas of Puerto Rico in relative categories of high and low density of population as well as high and

low density of developed area (Martinuzzi et al. 2007a).

Figure 6—Urban and rural land use classes in Puerto Rico (Martinuzzi et al. 2007a).
12

GENERAL TECHNICAL REPORT IITF-GTR-39

highest level of control on species composition and associated attributes such as

diversity and productivity (Lugo 1988). Diverse geologic substrates include volcanic, limestone, and serpentine bedrock and colluvial, alluvial, and marine quaternary deposits (figs. 8 and 9). Eleven of the twelve U.S. Geological Survey (USGS)

soil great groups are found in Puerto Rico (Mount and Lynn 2004). Within these

broad landscape features, plant community composition is controlled by topographic effects on soil moisture and soil development (slope position) and by

disturbance, including land use history, flooding, fire, landslides, and hurricanes,

with each inducing unique effects and subsequent secondary succession.

The resulting landscape in Puerto Rico is a mosaic of (1) primary forests that

have not experienced deforestation—according to Wadsworth (1951) and Birdsey

and Weaver (1982), less than 1 percent of the landscape supported climax vegetation by 1950; (2) mature secondary forests (> 25 years old)—less than 20 percent

of the island; (3) young secondary forests and shrublands (< 25 years old)—more

than 30 percent of the island; (4) agriculture, grasslands, and active and abandoned

pastures—over 30 percent; and (5) developed areas and urban forests—over 15 percent of the island. The mature secondary upland forests are primarily found in the

upland forest reserves, which occupy about 4 percent of the island. These protected

areas harbor our greatest wealth of native, endemic, and endangered plant species

(Figueroa and Woodbury 1996). Lowland flooded forests occupy about 1 percent

of the landscape and are typically protected as wetlands.

Figure 7—There are sixteen geoclimatic zones in Puerto Rico. The eight most extensive include the northern (moist) and southern

quaternary deposits (primarily alluvial) and limestone areas, the moist and wet volcanic areas, and the moist and dry serpentine areas

in the central mountains. Based on Holdridge life zones (Ewel and Whitmore 1973) and geologic terrain units (Bawiec 2001).
13

The Puerto Rico Gap Analysis Project

Figure 8—Topographic positions that are important controls on plant community composition (from Gould et al. 2008d and Martinuzzi

et al. 2007b).

Bromeliads are a common component of the palm forests on the

upper slopes of the Luquillo Mountains in Puerto Rico.

Mariano Solórzano
14

GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 9—A classification of 69 landscape units of Puerto Rico using climate, substrate, and topography, the primary controls on natural

vegetation cover (Gould et al. 2008c).
15

The Puerto Rico Gap Analysis Project

The fauna of Puerto Rico includes 436 species (app. 2). Three hundred twentyeight are birds, 57 reptiles, 27 mammals, and 24 amphibians. Of these, 387 are

terrestrial, 69 endemic, and 27 threatened or endangered. The birds include 43

introduced species with about 34 of these reproducing in the wild (Delannoy 2005)

and 98 resident species, including 19 endemic species. The reptiles are important

ecologically and serve as both prey and predator on a number of organisms and in

most habitats on the islands. Joglar (2005) indicated that nearly one quarter of the

native nonmarine reptiles are threatened or endangered with extinction. The majority of the terrestrial mammals, particularly all of the native species, are bats, which

include 13 species in Puerto Rico (Gannon et al. 2005). The amphibians are important ecologically and culturally in Puerto Rico. The tree frogs, or coquís, are

a national symbol as well as one of the most important nocturnal predators on the

island because of their abundance. Puerto Rican amphibians have high species

diversity but a low number of families and genera, a high degree of endemism,

and a number of threatened and endangered species (Joglar 2005). The Puerto

Rico GAP has analyzed 177 of the terrestrial vertebrates, including all of the

resident, endangered, and endemic species. Excluded species can be incorporated

in the future. These include migratory birds, and recently established exotics.

Land Cover Classification and Mapping

The GAP commits a high level of effort to mapping natural land cover. Generally,

land cover mapping is done by adopting or developing a land cover classification

system, delineating areas of relative homogeneity, then labeling these areas using

categories defined by the classification system. More detailed attributes of the

individual areas are added as more information becomes available, and a process

of validating both spatial pattern and labels is applied for editing and revising the

map. This is done in an iterative fashion, with the results from one step causing

reevaluation of results from another step. Finally, an assessment of the overall

accuracy of the data is conducted. The final assessment of accuracy will show

where improvements should be made in future updates (Stoms 1994).

In its “coarse filter” approach to conservation biology (e.g., Jenkins 1985, Noss

1987), gap analysis relies on maps of dominant natural land cover types as the most

fundamental spatial component of the analysis (Scott et al. 1993) for terrestrial

environments. For the purposes of the GAP, most of the land surface of interest

(natural) can be characterized by its dominant vegetation.
16

GENERAL TECHNICAL REPORT IITF-GTR-39

Vegetation patterns are an integrated reflection of the physical and chemical

factors that shape the environment of a given land area (Whittaker 1965). They also

are determinants for overall biological diversity patterns (Franklin 1993, Levin

1981, Noss 1990), and they can be used as a currency for habitat types in conservation evaluations (Austin 1991, Specht 1975). As such, dominant vegetation types

need to be recognized over their entire ranges of distribution (Bourgeron et al.

1994) for beta-scale analysis (sensu Whittaker 1960, 1977). These patterns cannot

be acceptably mapped from any single source of remotely sensed imagery. Reliance

on ancillary data, previous maps, and field surveys is necessary. The central concept

is that the physiognomic and floristic characteristics of vegetation across the land

surface can be used to define biologically meaningful biogeographic patterns. There

may be considerable variation in the floristics of subcanopy vegetation layers

(community association) that are not resolved when mapping at the level of dominant canopy vegetation types (alliance), and there is a need to address this part of

the diversity of nature. As information accumulates from field studies on patterns

of variation in understory layers, it can be attributed to the mapped units of vegetation alliances.

Land cover classifications rely on specified attributes, such as the structural

features of plants, their floristic composition, or environmental conditions, to

consistently differentiate categories (Küchler and Zonneveld 1988). The criteria

for a land cover classification system for GAP are:

• Ability to distinguish areas of different dominant vegetation.

• Utility for modeling animal species habitats.

• Suitability for use within and among biogeographic regions.

• Applicability to Landsat Enhanced Thematic Mapper plus (ETM+)

imagery both for rendering a base map and to extract basic patterns (GAP

relies on a wide array of information sources; ETM+ offers a convenient

meso-scale base map in addition to being one source of actual land cover

information).

• A framework that can interface with classification systems used by other

organizations and nations to the greatest extent possible.

• Capability to fit, both categorically and spatially, with classifications of

other themes such as agricultural and built environments.

For GAP in general, the system that fits best is referred to as the National

Vegetation Classification System (NVCS) (FGDC 1997). The origin of this system

was referred to as the UNESCO/TNC system (Lins and Kleckner 1996) because it
17

The Puerto Rico Gap Analysis Project

is based on the structural characteristics of vegetation derived by Mueller-Dombois

and Ellenberg (1974), adopted by the United Nations Educational, Scientific, and

Cultural Organization (UNESCO 1973) and later modified for application to the

United States by Driscoll et al. (1983, 1984). The Nature Conservancy and the

Natural Heritage Network (Grossman et al. 1994) have been using this system in

recent years with partial funding supplied by GAP. The basic assumptions and

definitions for this system have been described by Jennings (1993).

Classification of the PRGAP land cover follows a hierarchical structure separating vegetated from nonvegetated areas; forests, woodlands, and shrublands from

grasslands; and differentiating the composition of these vegetated areas based on

climate, substrate, topographic position, and age of forest types. These units are

then related to described formations, alliances, and associations to the extent

descriptions are available in the literature. We have been careful not to attribute

names of alliances or associations where there is uncertainty in the source or reliability of these names, and the degree of floristic information derived from field

studies. Alliance and association names are given only where there is supporting

documentation of the composition of these syntaxa in the literature. Community

descriptions with a full suite of species presence and abundance are not common

for Puerto Rico, and the dynamic and diverse nature of the vegetation make these

difficult to extrapolate. Alliance descriptions are more widely available, but primarily for the mature forest types. Information on the composition and variation in

herbaceous vegetation (excepting Dansereau 1966) and young secondary forests has

not been systematically done for the island.

Land Cover Classification Methods

The Puerto Rico Gap land cover mapping is based on two types of information:

(1) spectral and spatial information from satellite imagery and (2) ancillary or

additional data on the kinds of environmental factors that control vegetation composition, i.e., climate, substrate, topography, and disturbance. The spectral information was derived from Landsat 7 ETM+ imagery, and our initial task was to

develop a cloud-free data set of this imagery (Martinuzzi et al. 2006). Clouds are

a common feature of visible and infrared remotely sensed images collected from

Puerto Rico, as from many other tropical, humid, and coastal regions of the world.

Cloud-free data acquisition for a single date is extremely difficult for the entire

island, particularly for higher elevations where a high percentage of the protected
18

GENERAL TECHNICAL REPORT IITF-GTR-39

forest resources are located. Up to now, the continued presence of clouds in imagery has forced researchers to conduct most projects only when low cloud cover

images are available (years 1985, 1991, and 1992). We have developed simple

semi-automated procedures to generate cloud and cloud-shadow masks in tropical

regions for use in the creation of cloud-free composite Landsat 7 ETM+ imagery

using multiple recent images acquired at different times over the same region. Our

goals in developing these methods were to make use of spectral and geographic

information to facilitate the identification of cloud cover, separate cloud and topographic shadows, and minimize the masked areas needed to eliminate these features

from a scene. Ultimately, we developed mosaics of the four Landsat ETM+ pathrows that cover Puerto Rico, using the best imagery for each area.

In addition to the cloud and cloud-shadow masking method, we have incorporated processes for the creation of the final set of images. We sought to minimize

differences in reflectance owing to atmospheric conditions by correcting for

Rayleigh (atmospheric) scatter and differences in seasonality of vegetation by

using, whenever possible, imagery acquired during the same season. We also improved our future ability to separate subpixel features (some urban or mangrove

areas) by enhancing the spatial resolution, incorporating the higher (15-m2

) resolution of the panchromatic band of Landsat 7 ETM+.

Mapping standards and data sources—

Mapping was based on Landsat 7 ETM+ imagery acquired between 1999 and 2003,

and pan sharpened to 15-m2

 resolution. Most of the imagery used was from 2001

and 2003. A 15-m2

 pixel resolution is the minimum mapping unit (MMU) of the

PRGAP land cover. We developed a 96.5-percent cloud-free data set by compositing 18 images from four Landsat 7 ETM+ path-rows (4-47, 4-48, 5-47, and 5-48)

(Martinuzzi et al. 2006). For our analysis, we developed individual mosaics of the

area covered by each path-row using the best imagery for each area. We used

IKONOS imagery from 2001 and 2002 to manually interpret the remaining cloudcovered areas.

Land cover map development—

Our land cover layer includes spectral information from 1999-2003 Landsat

7 ETM+ imagery corrected for atmospheric distortion, clouds, and cloud shadows,

and pan-sharpened to a 15-m2

 resolution. Most of the imagery used was from the

years 2001-2003. Pixel classification was performed on the four mosaics (northeast,

northwest, southeast, and southwest Puerto Rico) by using ERDAS Imagine 8.7

(imagery processing software) unsupervised classification to distinguish the major
19

The Puerto Rico Gap Analysis Project

spectral classes. These were then mosaicked and stratified by information on the

primary factors controlling vegetation composition: i.e., climate, substrate, topography, and disturbance. Additionally, we incorporated information from site visits,

visual interpretation of aerial photography and IKONOS (Earth observation satellite) imagery, and comparison of classification results with published information

on vegetation for known sites. Image stratification was based on information that

allowed us to delimit sharp ecological boundaries with strong controls on vegetation composition, including geologic substrates (Bawiec 2001), and wetland classes.

Wetlands were delimited as forested or nonforested, seasonally or permanently

flooded, and saline or nonsaline by interpretation of the National Wetland Inventory

(NWI) digital database (USFWS 2005). Climatic boundaries include a combination

of the Holdridge life zones (Ewel and Whitmore 1973) and modeling using aspect,

which is related to rainfall and soil moisture patterns. We developed a landforms

model to delimit ridge, slope, valley, plains, and depressions in Puerto Rico and

used that to identify topographic positions and related variation in vegetation composition (Gould et al. 2006, Martinuzzi et al. 2007b). We also assessed information

from previous vegetation mapping efforts (Cintrón 1991, Helmer et al. 2002,

Ramos-González 2001, Ramos-González and Lugo 1994) as an aid in interpreting

classes. Our forest age class was determined by comparing our forested areas with

those considered forest or nonforest in the 1997 island-wide vegetation map

(Ramos-González and Lugo 1994). Areas considered pasture in 1997 and forest

in our mapping are designated as young (< 25-year-old) forests. Mona Island, an

uninhabited natural reserve, was mapped using different methods than the other

islands. We first stratified the island into its primary central limestone plateau and

a small area of coastal plain. Within these areas we used the variation in spectral

signal from recent Landsat imagery (15-m2

 resolution), variation in the normalized

difference of vegetation index (NDVI) (Goward et al. 1985), previous mapping

efforts (Cintrón 1991), and an ongoing effort to map the forested sinkholes in the

central plateau to develop a map of the island. Finally, we used IKONOS image

interpretation to refine the delimitation of small features including mangrove and

Pterocarpus swamps, agricultural lands such as row crops and palm plantations,

barrens, and some riparian vegetation.

Land Cover Classification Results

The PRGAP land cover of Puerto Rico includes 70 land cover units (table 1, app. 3

and 4). Puerto Rico, Vieques, Culebra, Mona, and the smaller islands have a

terrestrial extent (including lakes, rivers, and saline lagoons) of 8949 km2.
20

GENERAL TECHNICAL REPORT IITF-GTR-39

Table 1—The Puerto Rico Gap Analysis Project land cover classes and their area and percentage of

Puerto Rico’s total area

Land cover classes Area

Hectares Percent

Puerto Rico terrestrial extent 894 913 100.0

Forest, woodland, and shrubland 471 866 52.7

Forest 353 892 39.6

Mature forest 177 036 19.8

Young secondary forest 176 856 19.8

Woodland and shrubland 117 974 13.2

Dry 67 523 7.6

Dry forest 35 407 4.0

Mature dry forest 22 918 2.6

Young secondary forest 12 489 1.4

Woodland and shrubland 32 117 3.6

Alluvial deposits 10 130 1.1

Mature secondary lowland dry alluvial semideciduous forest 1548 0.2

Young secondary lowland dry alluvial semideciduous forest 2881 0.3

Lowland dry alluvial shrubland and woodland 4472 0.5

Lowland dry riparian forest 806 0.1

Lowland dry riparian shrubland and woodland 423 0.1

Calcareous substrates 28 741 3.2

Mature secondary lowland dry limestone evergreen forest 982 0.1

Mature secondary lowland dry limestone semideciduous forest 10 669 1.2

Young secondary lowland dry limestone semideciduous forest 3922 0.4

Lowland dry limestone woodland and shrubland 8181 0.9

Lowland dry limestone shrubland 4694 0.52

Lowland dry cactus shrubland 47 < 0.1

Coastal dwarf woodland and shrubland 103 < 0.1

Lowland dry limestone cliffside semideciduous forest 12 < 0.1

Lowland dry limestone cliffside shrubland and woodland 49 < 0.1

Abandoned dry forest plantation 81 < 0.1

Noncalcareous substrates 24 013 2.7

Mature secondary lowland dry noncalcareous semideciduous forest 7062 0.8

Young secondary lowland dry noncalcareous semideciduous forest 3645 0.4

Lowland dry noncalcareous shrubland and woodland 13 306 1.5

Ultramafic serpentine substrates 4640 0.5

Mature secondary dry and moist serpentine semideciduous forest 1839 0.2

Young secondary dry and moist serpentine semideciduous forest 1960 0.2

Dry and moist serpentine woodland and shrubland 841 0.1

Moist 236 768 26.5

Moist forest 176 258 19.7

Mature moist forest 90 041 10.1

Young secondary moist forest 86 217 9.6

Moist woodland and shrubland 60 510 6.8
21

The Puerto Rico Gap Analysis Project

Table 1—The Puerto Rico Gap Analysis Project land cover classes and their area and percentage of

Puerto Rico’s total area (continued)

Land cover classes Area

Hectares Percent

Alluvial deposits 15 497 1.7

Mature secondary lowland moist alluvial evergreen forest 2256 0.3

Young secondary lowland moist alluvial evergreen forest 6675 0.8

Lowland moist alluvium shrubland and woodland 5141 0.6

Lowland moist riparian forest 916 0.1

Lowland moist riparian shrubland and woodland 508 0.1

Calcareous substrates 72 255 8.1

Mature secondary moist limestone evergreen and semideciduous forest 46 423 5.2

Young secondary moist limestone evergreen and semideciduous forest 15 492 1.7

Moist limestone shrubland and woodland 10 338 1.2

Noncalcareous substrates 149 017 16.7

Mature secondary lowland moist noncalcareous evergreen forest 40 446 4.5

Young secondary lowland moist noncalcareous evergreen forest 52 981 6.0

Lowland moist noncalcareous shrubland and woodland 44 523 5.0

Lowland moist abandoned and active coffee plantations 11 068 1.2

Wet 158 614 17.7

Wet forest 133 266 14.9

Mature wet forest 55 116 6.2

Young secondary wet forest 78 150 8.7

Wet woodland and shrubland 25 348 2.8

Alluvial deposits 2501 0.3

Mature secondary montane wet alluvial evergreen forest 613 0.1

Young secondary montane wet alluvial evergreen forest 988 0.1

Montane wet alluvial shrubland and woodland 900 0.1

Noncalcareous substrates 151 724 17.0

Mature secondary montane wet noncalcareous evergreen forest 25 207 2.8

Montane wet evergreen abandoned and active coffee plantation 54 859 6.1

Mature primary and secondary montane wet noncalcareous evergreen

tabonuco forest 8715 1.0

Mature primary and secondary montane wet noncalcareous evergreen

palo colorado cloud forest 3712 0.4

Mature primary and secondary montane wet noncalcareous evergreen

sierra palm forest 11 953 1.3

Mature primary and secondary montane wet noncalcareous evergreen

elfin woodland cloud forest 1537 0.2

Young secondary montane wet noncalcareous evergreen forest 21 651 2.4

Montane wet evergreen noncalcareous shrubland and woodland 24 090 2.7

Ultramafic serpentine substrates 4389 0.5

Mature secondary montane wet serpentine evergreen forest 3379 0.4

Young secondary montane wet serpentine evergreen forest 651 0.1

Wet serpentine shrubland and woodland 358 < 0.1

Flooded forests 8960 1.0

Mangrove forest and shrubland 8700 1.0

Freshwater Pterocarpus swamp 261 < 0.1
22

GENERAL TECHNICAL REPORT IITF-GTR-39

Forest, woodland, and shrubland—

Fifty-three percent (4719 km2

) of Puerto Rico is covered predominantly by woody

vegetation, and 49 land cover units are described as dominated by woody vegetation. Of these, low- and mid-elevation moist woody vegetation covers 27 percent

(2368 km2

), upper elevation wet woody vegetation covers 18 percent (1586 km2),

dry woody vegetation covers 8 percent (675 km2

), and flooded mangrove and

Pterocarpus forests cover 1 percent (90 km2

) of the islands. We have classified 40

percent (3539 km2

) of the woody vegetation as forest (> 60 percent tree cover), and

Table 1—The Puerto Rico Gap Analysis Project land cover classes and their area and percentage of

Puerto Rico’s total area (continued)

Land cover classes Area

Hectares Percent

Grasslands 286 407 32.0

Dry 42 389 4.7

Dry grasslands and pastures 42 176 4.7

Dry cactus grassland and shrubland 213 < 0.1

Moist 218 945 24.5

Moist grasslands and pastures 218 945 24.5

Wet 23 090 2.6

Seasonally flooded herbaceous nonsaline wetlands 18 225 2.0

Seasonally flooded herbaceous saline wetlands 4865 0.6

Flooded 1984 0.2

Emergent herbaceous nonsaline wetlands 1102 0.1

Emergent herbaceous saline wetlands 882 0.1

Agriculture 26 257 2.9

Hay and row crops 25 764 2.9

Woody agriculture and plantations 492 0.1

Natural barrens 3576 0.4

Rocky cliffs and shelves 397 < 0.1

Gravel beaches and stony shoreline 82 < 0.1

Fine to coarse sandy beaches, mixed sand and gravel beaches 1155 0.1

Riparian and other natural barrens 449 0.1

Salt and mudflats 1493 0.2

Artificial barrens 8694 1.0

Artificial barrens 8674 1.0

Salt production 20 < 0.1

Developed areas 89 573 10.0

High-density urban development 52 335 5.9

Low-density urban development 37 237 4.2

Water 8540 1.0

Freshwater 4434 0.5

Saline water 4005 0.5

Aquaculture 101 < 0.1
23

The Puerto Rico Gap Analysis Project

13 percent (1180 km2) as woodland (< 60 percent tree cover) or shrubland (> 25

percent cover small trees or shrubs). Mature (> 25-year-old) and young secondary

(< 25-year-old) forests each cover 20 percent of the islands. Dry, moist, wet, and

flooded forests cover 4 percent, 20 percent, 15 percent, and 1 percent of the islands,

respectively. Mature dry, moist, wet, and flooded forests cover 3 percent, 10 percent, 6 percent, and 1 percent of the islands, respectively. The most abundant

forest types are the montane wet evergreen secondary forest, including active

and abandoned coffee plantations, and young secondary lowland moist forest on

noncalcareous substrates. These each cover 6 percent of the islands (549 km2

 and

530 km2

, respectively).

Grassland—

Thirty-five percent (3127 km2

) of Puerto Rico has been classified as grassland

(32 percent) or herbaceous agriculture (3 percent). Dry, moist, wet, and flooded

grasslands make up 5 percent, 25 percent, 3 percent, and less than 1 percent, respectively, of the area. Nearly all of the moist and dry grasslands are maintained by

disturbance. They may have continuous or intermittent cattle grazing and may burn

frequently, particularly the dry grasslands. Natural barrens make up less than 1 percent of the area. These are a small but important component of the landscape, both

for human use and as wildlife habitat. Natural barrens include the stony and sandy

beaches, rocky cliffs and shelves, active riparian flood plains, and salt- and

mudflats.

Wetlands—

Coastal wetlands cover 4 percent of the island (340 km2

). Forty-two percent of these

are saline wetlands and 58 percent freshwater wetlands. Seventy-four percent of the

wetlands are dominated by herbaceous vegetation (251 km2), and 92 percent of

these (230 km2

) are seasonally flooded. Of the herbaceous wetlands, 77 percent are

nonsaline (193 km2

) and 23 percent are saline (58 km2). Forested coastal wetlands

cover 1 percent of the island; 67 km2

 are mangroves and 3 km2 are freshwater

Pterocarpus swamps. About 1 percent of the area has been classified as either fresh

or saline water.

Developed areas—

We classified 11 percent (983 km2

) of Puerto Rico as developed (10 percent) or

artificially barren (1 percent). This is a very dynamic component of the landscape

and has likely increased since the imagery for mapping was acquired. Development

does not occur equally around the region and is concentrated in the coastal plain and

lower hills (Martinuzzi et al. 2007a, Pares et al. in press).
24

GENERAL TECHNICAL REPORT IITF-GTR-39

Land Cover Classification Accuracy Assessment

The purpose of accuracy assessment is to allow a potential user to determine the

map’s “fitness for use” for their application. It is impossible for the original

cartographer to anticipate all future applications of a land cover map, so the

assessment should provide enough information for the user to evaluate fitness for

their unique purpose. This can be described as the degree to which the data quality

characteristics collectively suit an intended application. The information reported

includes details on the database’s spatial, thematic, and temporal characteristics and

their accuracy.

Assessment data are valuable for purposes beyond their immediate application

to estimating accuracy of a land cover map. The reference data are therefore made

available to other agencies and organizations for use in their own land cover

characterization and map accuracy assessments (see “Data Availability” for access

information). The data set will also serve as an important training data source for

later updates.

Even though we have reached an endpoint in the mapping process where

products are made available to others, the gap analysis process should be considered

dynamic. We envision that maps will be refined and updated on a regular schedule.

The assessment data will be used to refine GAP maps iteratively by identifying

where the land cover map is inaccurate and where more effort is required to bring

the maps up to accuracy standards. In addition, the field sampling may identify new

classes that were not identified at all during the initial mapping process.

Methods—

We used island-wide 1-m2

-resolution color IKONOS imagery from 2001-2002,

including Vieques, Culebra, Mona, and the smaller cays to evaluate the thematic

accuracy of the 2003 land cover map of Puerto Rico and the surrounding islands,

derived from 18 Landsat 7 ETM+ scenes collected from 1999 to 2003.

A feasibility appraisal was conducted prior to the accuracy assessment, which

concluded that the accuracy assessment should be conducted on the six original

classes obtained through the unsupervised classification. The final 70 PRGAP land

cover units were created through modeling of the original classes in combination

with geological, climatological, and other auxiliary data. Therefore, the 70-unit

classification was simplified to six classes. The six classes were chosen as they

represented the main classes originally separated spectrally through the 18 Landsat

7 ETM+ scenes prior to the extensive modeling process. Furthermore, the recoded
25

The Puerto Rico Gap Analysis Project

six land cover classes simplified the accuracy assessment process and helped to

reduce image interpretation errors when using the reference IKONOS imagery

(table 2).

The use of alternative methods for the accuracy assessment of GAP land cover

products has previously been recommended by the GAP where the collection of

ground data has been considered impractical. Lin and Laporte (2003) assessed

IKONOS and aerial videography for use in accuracy assessment, and the National

Aeronautics and Space Administration (NASA) aerial photos have been found

suitable for the accuracy assessment where large study areas and staff resources

made reliance on field data impractical (Helmer et al. 2002, Zhu et al. 2000).

A remote-sensing specialist, GIS specialist, and a research ecologist with an

indepth knowledge of the vegetation of Puerto Rico were available for image

interpretation of the IKONOS imagery, helping to reduce inaccuracy. Additionally,

tone, shape, size, pattern, texture, shadow, and association as well as true- and

false-color composites were all used as criteria for visual interpretation. Image

interpretation training using the IKONOS imagery included creating small reference subsets of each class that could be used as a base for image interpretation.

Sample size—

Prior to conducting the accuracy assessment, we determined the minimum number

of sample points required so that our calculated classification accuracy would have

an allowable error of 5 percent at the 95-percent confidence interval. An assumed

error range was used to calculate the minimum number of samples required to

achieve the specified allowable error. For this calculation, we assumed that the

overall accuracy of the land cover map was between 70 and 85 percent. The number

of sample points was calculated using the following equation based on binomial

probability theory (Fitzpatrick-Lins 1981):

Where

N = number of samples

p = expected or calculated accuracy ( percent)

q = 100 – p

E = allowable error

Z = standard normal deviate for the 95-percent two-tail confidence level (1.96).

2

2

E

Z pq N =
26

GENERAL TECHNICAL REPORT IITF-GTR-39

For the lowest expected map accuracy of 70 percent with an allowable error

of 5 percent, 323 sample points were required, whereas for a map accuracy of

85 percent with an allowable error of 5 percent, 196 sample points were required

(table 3).

We then decided to use the lowest expected land cover map accuracy (70 percent) in determining the minimum number of sample points (~ 323) for the accuracy assessments. A stratified random sampling technique was considered the most

appropriate for the land cover accuracy assessment. Table 4 identifies the allocation

of the sample points to the different land cover categories. However, some classes

were not appointed an adequate number of sample points because the area covered

by some of the smaller land cover classes is negligible compared to the rest of the

classes. Therefore, the minimum sample size was set to 20 for these classes (Maingi

et al. 2002, van Genderen and Lock 1977), therefore increasing our total number

of sample points from 323 to 358.

Three hundred and fifty-eight sample points were randomly allocated to the

PRGAP land cover classes as shown above. To avoid bias, image interpreters did

not know what classifications had been assigned to the sample points in the PRGAP

land cover. The corresponding reference sample points were then analyzed in the

IKONOS imagery and allocated to one of the six classes. Some of the sample

points fell in areas of cloud cover within the reference IKONOS imagery; these

points were removed and new reference points were selected through stratified

sampling until the allocated sample number for each class was attained. An ERDAS

imagine 9.0 was used to generate an error matrix, accuracy totals, and kappa

statistics.

Table 2–Absolute and relative areas of the Puerto Rico

Gap Analysis Project land cover classes

Land cover Area

Hectares Percent

Forest (except mangrove) 345 132.0 39

Woodland and shrubland 117 974.4 13

Mangrove 8 699.5 1

Grassland, pasture, agriculture 312 663.9 35

Urban and barren 101 844.5 11

Water 8 540.3 1

Total 894 913.0 100
27

The Puerto Rico Gap Analysis Project

Land cover accuracy assessment results—

The results of the IKONOS-based 2003 PRGAP land cover accuracy assessment

(tables 5 and 6) show an overall accuracy of 84.92 percent and a kappa value of

0.8, which indicates substantial agreement (Landis and Koch 1977). However,

there is significant variability in the producer’s and user’s accuracy. The producer’s

accuracy (PA) relates to the probability that a reference sample (IKONOS-interpreted land cover class) will be correctly mapped and measures the errors of omission, whereas the user’s accuracy (UA) indicates the probability that a sample from

the land cover map matches the reference data and measures the error of commission. The producer’s accuracy ranges from 52.54 percent to 100 percent and the

user’s accuracy ranges from 72.09 percent to 95 percent (table 6). Overall, accuracy

Table 3—Minimum number of sample points

(N1) required to achieve an allowable error

of 5 percent (E1) at the 95-percent confidence

interval

N1 p q

323 70 30

288 75 25

246 80 20

196 85 15

Note: Z = 1.96 (standard normal deviate for the 95-

percent two-tailed confidence interval), p = expected

percentage accurracy, q = 100 - p , E1

2

 = 25 (square

of allowable error).

Table 4—Minimum number of sample points per land cover class stratified by area

Estimated Final

number number

Land cover Area of samples of samples

Hectares Percent

Forest (except mangrove) 345 132 39 125 125

Woodland and shrubland 117 974 13 43 43

Mangrove 8 700 1 3 20

Grassland, pasture, agriculture 312 664 35 113 113

Urban and barren 101 845 11 37 37

Water 8 540 1 3 20

Total 894 913 100 323 358
28

GENERAL TECHNICAL REPORT IITF-GTR-39

assessment for five of the six recoded classes tended to fall in a similar range: from

87 percent to 100 percent for the producer’s accuracy and from 82 percent to 95

percent for the user’s accuracy. However, for the open forest and shrubland class,

the PA fell to 52 percent and the UA to 72, indicating a degree of misclassification.

With any land cover classification produced from satellite imagery, misclassification

often results from subpixel spatial variability and spatial and spectral resolution

limitations.

Table 5—Error matrix of IKONOS-based accuracy assessment of the Puerto Rico

Gap Analysis Project major land cover classes

Total number of

Land cover class Error matrix pixels

(1) (2) (3) (4) (5) (6)

(1) Forest (except mangrove) 108 9 0 620 125

(2) Woodland and shrubland 8 31 0 310 43

(3) Mangrove 0 0 19 001 20

(4) Grassland, pasture, agriculture 2 16 0 93 2 0 113

(5) Urban and barren 0 2 0 1 34 0 37

(6) Water 0 1 0 0 0 19 20

Total 118 59 19 103 39 20 358

Note: The reference data are from IKONOS 2001–2002 imagery. The number of correctly identified

pixels are in the diagonal portion of the matrix and misidentified pixels are in the row or column of

the land cover type in which they occur in the IKONOS imagery.

Table 6—Accuracy of land cover classifications of the Puerto Rico Gap Analysis

Project

Land cover class RT CT NC PA UA Kappa

– – – Percent – – –

Forest (except mangrove) 118 125 108 91.53 86.40 0.7971

Woodland and shrubland 59 43 31 52.54 72.09 0.6659

Mangrove 19 20 19 100.00 95.00 0.9472

Grassland, pasture, agriculture 103 113 93 90.29 82.30 0.7515

Urban and barren 39 37 34 87.18 91.89 0.9090

Water 20 20 19 95.00 95.00 0.9470

Total 358 358 304

Overall Kappa statistics (KHAT value) 0.8007

Overall accuracy (percent) 84.92

Note: RT = reference pixels, CT = classified pixels, NC = number pixels correctly classified, PA =

producer’s accuracy (samples correctly mapped), US = user’s accuracy (mapped point matches data).
29

The Puerto Rico Gap Analysis Project

Puerto Rico provides a diverse, dynamic, fragmented, and significantly

heterogeneous landscape for land cover classification. Even with spatial resolution enhancement of the Landsat 7 ETM+ reflectance bands from 30 m2

 to 15 m2,

features at the subpixel level and mixed pixelsy—e.g., confusion between urban,

barren and sands, or water and closed forest—provided a significant problem.

Another issue that influenced land cover accuracy results is the misregistration

between the imagery used to create land cover classification and the reference data

used in the accuracy assessment. This is especially predominant when land cover

accuracy assessments are completed by using aerial photography, videography, or,

as in this case, other satellite imagery. The PRGAP land cover used 18 Landsat 7

ETM+ images, which were mosaicked to create an island-wide cloud-free composite. The mosaic was then segmented into six geoclimatic zones to reduce spectral

variability and assist classification. With different preprocessing steps taken in

both the Landsat 7 ETM+ imagery set and the IKONOS imagery, juxtaposed with

variation in spatial and spectral characteristics of each data set, perfect registration

between the imagery was complex; therefore, some of the error in the accuracy

could be due to misregistration rather than misclassification.

Even though the six recoded PRGAP land cover units provided a significant aid

to image interpretation when using IKONOS imagery as a reference for the accuracy assessment, it is still possible that some error is due to the inability to confidently image-interpret a sample unit. This is particularly true in Puerto Rico, where

there are gradients between shrubland and forest that are difficult to distinguish.

Land Cover Classification Limitations

The land cover of Puerto Rico is a matrix of very stable and very dynamic elements in terms of species composition. The dynamic nature is due to both human

use and land cover change, the ability of woody vegetation to quickly replace

abandoned pastures, and dynamism related to natural disturbance, particularly

hurricanes and fire. Stable elements include those that were unsuitable for agriculture and adapted to the regime of natural disturbances. These include the coastal

mangroves, extremely wet cloud forest vegetation, and extremely dry vegetation on

rocky coastal areas. Vegetation description in Puerto Rico has a long history and

much detail for some parts of the island, particularly the Luquillo Mountains.

Other, more disturbed areas are less well described, and in general, description at

the plant community level, including all woody and herbaceous species present, is

limited. Little classification other than Dansereau (1966) attempted detailed plant

community description for the herbaceous communities on the island. The current
30

GENERAL TECHNICAL REPORT IITF-GTR-39

land cover and classification represents the land cover at the time of image acquisition (2003) and is an attempt to link these units with available published descriptions to the finest scale at which they are available.

Predicted Vertebrate Distributions and Species

Richness

All species range maps are predictions about the occurrence of those species within

a particular area (Csuti 1994). Traditionally, the predicted occurrences of most species begin with samples from collections made at individual point locations. Most

species range maps are small scale (e.g., >1:10,000,000) and derived primarily

from point data to construct field guides. These are suitable, at best, for approximating distribution at the regional level or counties. The purpose of the GAP

vertebrate species maps is to provide more precise information about the current

predicted distribution of individual native species according to actual habitat

characteristics within their general ranges and to allow calculation of predicted

area of distributions and associations to specific habitat characteristics.

The GAP maps are produced at a nominal scale of 1:100,000 or better and are

intended for applications at the landscape or “gamma” scale (heterogeneous areas

generally covering 1000 to 1 million hectares and made up of more than one kind

of natural community). Applications of these data to site- or stand-level analyses

(site—a microhabitat, generally 10 to 100 m2

; stand—a single habitat type, generally 0.1 to 1000 ha) (Whittaker 1977, see also Stoms and Estes 1993) will likely

reveal the limitations of using this process to incorporate differences in habitat

quality (e.g., understory condition) or necessary microhabitat features such as

standing dead trees.

Gap analysis uses the predicted distributions of animal species to evaluate their

conservation status relative to existing land management (Scott et al. 1993). However, the maps of species distributions may be used to answer a wide variety of

management, planning, and research questions relating to individual species or

groups of species. In addition to the maps, great utility may be found in the consolidated specimen collection records and literature that are assembled into databases used to produce the maps. Perhaps most importantly, as a first effort in

developing such detailed distributions, they should be viewed as testable hypotheses

to be confirmed or refuted in the field. We encourage biologists and naturalists to

conduct such tests and report their findings in the appropriate literature and to the

GAP such that new data may improve future iterations.
31

The Puerto Rico Gap Analysis Project

Before this effort, there were no maps available, digital or otherwise, showing

the likely present-day distribution of species by habitat type across their ranges.

Because of this, ordinary species (i.e., those not threatened with extinction nor

managed as game animals) are generally not given sufficient consideration in land

use decisions in the context of large geographic regions or in relation to their actual

habitats. Their decline, because of incremental habitat loss, results in one threatened or endangered species “surprise” after another. Frequently, the occurrence

records that exist for species with a wide range are artificially truncated by state or

other politically-based boundaries. As a result, creating a consistent spatial framework for storing, retrieving, manipulating, analyzing, and updating our knowledge

about the status of a given animal species is one of the most necessary and basic

elements for preventing further erosion of our biological resources.

Although there is a wealth of ecological and natural history information for

Puerto Rico, it is not available in a centralized location of consistent quality or

format. A great deal of the PRGAP effort was in compiling this information to

develop species occurrence, range, and predicted distribution databases and maps.

We created a relational database for the PRGAP to manage data on species occurrences, species habitat information, and literature citations (see volume 2). Each

animal species represented in the PRGAP relational database is associated with a

unique identifier (PRGAPSppID) that is used to link (relate) the species to records

on its occurrence, habitat preferences, and conservation status, which are stored in

multiple tables contained in the database. The objective is to identify, through documented sources or expert opinion, those physiographic and biological elements

with which a species is associated. Predicting species geographic distributions

requires building a database that identifies all the major physiographic and biological elements considered to influence the occurrence of a species across its range.

Published books, scientific papers, and reports about the ecology and life history of

individual species or groups of species are the primary sources for information

about the habitats in which a species can be expected to occur. Official Internet

sources, unpublished documents, and articles contained in popular media were also

used, but are considered less credible. Accuracy assessment was conducted for each

species by incorporating an expert opinion and review process.

Vertebrate Mapping Methods

PRGAP species list—

Over 470 vertebrate species (excluding fish) have been recorded in Puerto Rico and

its adjacent islands including those that occur in terrestrial, terrestrial aquatic, and
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GENERAL TECHNICAL REPORT IITF-GTR-39

marine environments (app. 2). Of these, about 436 are terrestrial vertebrate species.

Many of these are identified as migratory, wintering, accidental, or vagrant species

that do not breed regularly or at all in Puerto Rico. The PRGAP only considered

species that are known to breed in the project area, and that are regularly occurring

nonaccidentals (Csuti and Crist 1998). Csuti and Crist (1998) suggested, as a

general definition, that “regular breeders” are those species breeding in the state

during at least 5 of the past 10 years. However, this is often difficult to document.

The species list for the PRGAP project final analysis was compiled in cooperation with the DNER, the USFWS, and several nongovernmental sources. This list

does not include most of the nonnative species that are known to occur in Puerto

Rico. However, because several nonnative species are known to influence the

distribution or densities of native species or are valued as game species, we elected

to include a group of those that are considered regularly occurring in Puerto Rico.

Our final list of 177 species included 18 amphibians, 98 birds, 14 mammals, and 47

reptiles (fig. 10).

Mapping Standards and Data Sources—

The PRGAP mapped predicted species distributions in accordance with the standards of the GAP handbook as of 13 January, 2000. All GIS modeling of species

distributions was conducted on Dell workstations running ERDAS Imagine and

ArcINFO 9.0.

Figure 10—Terrestrial vertebrate species by taxonomic group included in the Puerto Rico Gap

analysis.
33

The Puerto Rico Gap Analysis Project

Minimum Mapping Unit (MMU)—

The PRGAP recognizes two mapping unit scales that are related to (1) defining a

species geographic range determined by its probability of occurrence or (2) a

predicted species distribution based on associated range, habitat, and life history

variables. The PRGAP has adopted a hexagon grid network for use in mapping

Puerto Rico’s biological diversity. This hexagon grid (PRGAP-HEX) provides a

uniform unit of area that can be used to represent the range and occurrence of

vertebrate species across a very heterogeneous landscape. Each hexagon has an area

of 24 km2

, which is the MMU for species geographic range distributions (fig. 11).

The PRGAP-HEX grid consists of 483 individual hexagons with 305 occurring only

over land, 161 over coastal areas, and 17 over open marine areas with small reefs

and cays. The hexagon shape is based on the U.S. Environmental Protection

Agency’s (USEPA) Environmental Monitoring and Assessment Program (EMAP)

typically used in gap analysis. However, EMAP represents only the conterminous

United States, and in lieu of EMAP coverage in the Caribbean, the PRGAP-HEX

grid (fig. 1) was developed by the USFS by tessellating the larger hexagonal grid

used in Caribbean Forest Inventory and Analysis (FIA).

Figure 11—A 24 km2 hexagon was used as the minimum

mapping unit for species geographic range distributions.
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GENERAL TECHNICAL REPORT IITF-GTR-39

The second mapping unit scale is derived from the 15-m2-pixel resolution

reflected in the PRGAP land cover layer (fig. 12), and that is used in mapping

species habitat and predicted distribution. The PRGAP uses a 15-m2-pixel resolution as the MMU for mapping predicted species distributions.

Mapping species geographic distribution—

To map a species geographic distribution, each hexagon was attributed with one of

eight categories for probability of occurrence of that species (table 7). Species

probability of occurrence information is derived from published literature, unpublished data sets, museum records, and expert opinion. A species record of occurrence may be confirmed when associated with a credible observation, including the

location, observation date, and observer’s name. Records may be probable based on

published range maps, location descriptions, or expert opinion, or predicted based

on the occurrence of habitat and expert opinion that the species is likely to occur.

Most of the records are for birds, followed by reptiles, amphibians, and mammals

(fig. 13).

Collecting species data—

We used a great variety of sources to develop species geographic ranges and their

predicted distributions based on habitat availability as inferred from PRGAP land

cover. Vertebrate occurrence data were gathered from DNER and include records

collected for the Critical Wildlife Areas Report, element occurrence records on

threatened and endangered species managed by the DNER’s Natural Heritage

Program, and from their long-term pigeon and dove survey. In addition, we included occurrence records obtained from:

• USGS Patuxent Wildlife Research Center—10 years of the USGS Breeding

Bird Survey Program.

• National Audubon Society Christmas Bird Count.

• USGS Cooperative Unit karst study with North Carolina State University.

• Luquillo Experimental Forest long-term ecological research studies.

• Puerto Rico Ornithological Society Puerto Rico Breeding Bird Atlas

project.

• Smithsonian Museum of Natural History.

• University of Puerto Rico, Mayagüez and Humacao campuses.

• IITF staff and scientists.

• University and independent researchers.

• Peer-reviewed publications, research reports, books, and published maps.
35

The Puerto Rico Gap Analysis Project

The Antillian crested hummingbird Orthorhyncus cristatus is known as the Zumbador Crestado

locally.

Iván Vicens
36

GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 12—Land cover of Puerto Rico. Developed at 15-m2 resolution using Landsat 7 ETM+ imagery acquired in 1999-2003 and ancillary data (legend follows).
37

The Puerto Rico Gap Analysis Project
38

GENERAL TECHNICAL REPORT IITF-GTR-39

Table 7—Confidence level assigned to hexagon records for each species

Confidence level Taxon Description

Confirmed All Confidently assumed or known to occur in the

hexagon. Sources include species locality records

and expert opinion.

Predicted All Predicted to occur based on a combination of

presence of suitable habitat and historical record

or presence in adjacent hexagon. Sources include

expert opinion only.

Probable All Probable occurrence based on a strong likelihood.

Sources include expert opinion, published range

maps, or range descriptions.

Historical included All Confidently assumed or known to have occurred

in the hexagon prior to 1970 and considered as

valid for recent distribution. Sources include

species locality records and expert opinion.

Historical excluded All Confidently assumed or known to have occurred

in the hexagon prior to 1970 but considered

invalid for recent distribution. Sources include

species locality records and expert opinion.

Questionable All Occurrence within hexagon was still in question

after expert review. Hexagons coded as questionable are not included in species current range

distribution. Sources include expert opinion only.

Excluded All Documented occurrence was excluded by expert

review after having been coded as confirmed,

predicted, or probable. Sources include expert

opinion only.

Absent All Considered absent from hexagon based on no

documented records of occurrence and expert

opinion. Sources include locality records and

expert opinion.

Figure 13—Occurrence

records for each taxonomic

group in the Puerto Rico

Gap Analysis Project

database.
39

The Puerto Rico Gap Analysis Project

These last sources were particularly useful when mapping reptiles, amphibians,

and bats. All of this information is maintained in the PRGAP relational database.

Approximating species geographic range extent—

For each species included in our analysis, we mapped its geographic distribution by

intersecting the species occurrence records with our PRGAP-HEX base map; the

hexagon cell represents the minimum mapping unit for interpreting species geographic range extent. The hexagon was preferred over other mapping units for three

reasons. First, the uniform shape and size is important when comparing relative

measures of density associated with species diversity. Second, the hexagon’s geometric properties allow it to be easily aggregated or tessellated to conform to the

convex shape of the Earth’s surface. Third, and equally important, the hexagon

border is determined arbitrarily and overcomes many problems associated with

delineating species range by county (Boone 1996) or other politically-based boundaries. An additional reason for using the PRGAP-HEX base map to delineate species range extents is that each hexagon is spatially related to data collected by the

Caribbean FIA Program. This provides for an opportunity to conduct future analyses of vertebrate biological diversity, forest inventory and use, tree species diversity,

and forest health.

Each vertebrate occurrence record with latitude and longitude coordinates was

attributed to a single hexagon that, in turn, was classified as “confirmed” or “historical included” depending on the date of record (table 7). Museum or other

records with locality information that could be assigned to a single hexagon were

also designated “confirmed” or “historical included.” Published distribution maps

and descriptions, and locality records that intersect more than one hexagon were

also used to attribute hexagons. However, this process was less straightforward

requiring different methods or steps depending on the record’s unique case. In these

cases, the hexagons intersecting these locality records were attributed or classified

as “probable” (table 7).

Methods specific to birds—

Breeding bird surveys represented a very important source of bird distribution data.

Observations from BBS routes were attributed to all the hexagons that were intersected by the routes. These hexagons were classified as “confirmed.” The following

references also represented valuable tools for mapping bird distributions: Biaggi

(1977), Oberle (2000), Raffaele (1983), and Raffaele et al. (1998).

We selected all the hexagons that intersected the distribution localities or areas

described in these references, and attributed them an occurrence classification of

“probable” (table 7).
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GENERAL TECHNICAL REPORT IITF-GTR-39

Methods specific to reptiles and amphibians—

Valuable amphibian distribution information came from the following references:

Joglar (1998), Rivero (1998), and Schwartz and Henderson (1991). Point distribution maps from some of these references were electronically scanned, points of

occurrence digitized, and then buffered (buffer distance differed depending upon

the inherent error associated with each respective published map) to reflect processing error. The respective buffered points were then intersected with a complete

species hexagon distribution data set (which includes hexagons still categorized as

“absent”) to augment the current distribution status. All hexagons selected using this

process were attributed as “probable.”

Methods specific to mammals—

A valuable source of bat distribution information was provided by Dr. M. Gannon

of Penn State University, and subsequently from Gannon et al. (2005). We digitized

the point locations from the distribution maps in this book. We assigned a buffer to

each of the digitized points depending on the observation or record description that

Gannon et al. (2005) provided. The buffers were intersected with the complete

species hexagon data sets to complement the existing distribution records, and

subsequently attributed with an occurrence category of “probable.”

Wildlife habitat relationships—

Species habitat models were based on species life-history information obtained

from the literature and from expert review. The information used to develop species habitat models has been documented in the PRGAP-VERT database, where we

compiled all available information on species taxonomic classification, conservation

status, worldwide distribution, distribution in Puerto Rico, associations with vegetation or floristics, geologic substrates, soils, climate or rainfall patterns, elevation,

topography, habitat structure, or other terrain features. We also documented information on species migratory, reproductive, and demographic patterns, dietary

habits, habitat use, and activity patterns. This information, as well as a specific

wildlife habitat model, is reported in volume 2 for each species.

Modeling of vertebrate species distributions for PRGAP—

Species habitat models were linked to specific mapped land cover units or other

information for which we have reliable spatial information. The predicted distributions include all habitats within hexagons with confirmed, probable, or predicted

occurrences, and in adjacent hexagons when there is contiguous habitat.
41

The Puerto Rico Gap Analysis Project

The barred anole Anolis straulus is know locally as the Lagartijo Manchado.

Javier Mercado
42

GENERAL TECHNICAL REPORT IITF-GTR-39

Vertebrate Distribution Results

Hexagon occurrence maps (see volume 3) provide a comprehensive set of species

occurrences for Puerto Rico. They indicate that although there are a number of

widespread species occurring nearly everywhere, there are also significant patterns

of coastal or montane species, species restricted to dry or wet environments, and

species restricted to particular islands. They also indicate the need for island-wide

systematic surveys and more comprehensive sampling of several species.

Predicted distribution maps (see volume 4) also provide a new body of information that is available for future analysis. They highlight the importance of the

diversity of habitats on the island, from wetlands to dry lands, from grassland to

forests, and the matrix of habitats that include mangrove and dry forests, forest

edges, and tracts of dense forest. Predicted distribution maps include information

on species taxonomy, conservation status, habitat model, natural history, major

literature sources, and, for many species, a photograph.

Amphibians—

We mapped the geographic distribution (see hexagon occurrence maps) of 18

amphibian species based on associated occurrence records. The majority of Puerto

Rico’s amphibians are in the genus Eleutherodactylus, the coquís of Puerto Rico. As

an example, Eleutherodactylus portoricensis is the Puerto Rican coquí, with a range

above 200 m in the central mountains (fig. 14) and habitat that includes forest,

woodland, and shrubland within this range (fig. 15).

Birds—

We mapped the distribution of 98 bird species. As an example, the yellowshouldered blackbird, Agelaius xanthomus, is an uncommon and endangered

species that is mostly restricted to the drier coastal areas (fig. 16) and inhabits

forests, woodlands, shrublands and open areas adjacent to forest (fig. 17).

Mammals—

We mapped the distribution of 14 mammal species, including the 13 bats found on

the islands. As an example, the Jamaican fruit-eating bat, Artibeus jamaicensis, is

common throughout the island (fig. 18) and inhabits forests, woodlands, shrublands,

and adjacent grasslands (fig. 19).

Reptiles—

We mapped the distribution of 47 reptile species. As an example, the Puerto Rican

boa, Epicrates inornatus, is widespread in its distribution, but uncommon (fig. 20).

It inhabits moist and wet forested areas or dense dry forest near water (fig. 21).
43

The Puerto Rico Gap Analysis Project Figure 14—Eleutherodactylus portoricensis hexagon occurrence map.
44

GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 15—Eleutherodactylus portoricensis predicted distribution map.
45

The Puerto Rico Gap Analysis Project Figure 16–Agelaius xanthomus hexagon occurrence map.
46

GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 17—Agelaius xanthomus predicted distribution map.
47

The Puerto Rico Gap Analysis Project Figure 18—Artibeus jamaicensis hexagon occurrence map.
48

GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 19—Artibeus jamaicensis predicted distribution map.
49

The Puerto Rico Gap Analysis Project Figure 20—Epicrates inornatus hexagon occurrence map.
50

GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 21—Epicrates inornatus predicted distribution map.
51

The Puerto Rico Gap Analysis Project

The Coquis are an important national symbol of the ecological richness of Puerto Rico.

Javier Mercado
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GENERAL TECHNICAL REPORT IITF-GTR-39

Vertebrate Richness

The GAP has often been associated with the mapping of species-rich areas or “hotspots.” Richness maps identify numbers of elements that co-occur by geographic

location. These are color coded or shaded in intensity from the highest numbers of

co-occurrence (richness) to the lowest. Areas with the greatest species richness may

or may not indicate best conservation opportunities. They may occur in already

protected areas or may represent mostly species that are currently protected or not

at risk. Still, they are often a useful starting point to examine conservation opportunities in combination with other analyses described in this report’s “Introduction”

and “Analysis” sections. They may be useful for other applications such as identifying places of interest for wildlife observation and study.

Species richness is depicted for all mapped taxa at a 15-m2

-pixel resolution

where each pixel is associated with a habitat type and by a color that represents the

number of vertebrate species potentially using that habitat, a measure of alpha

diversity (Whittaker 1977) (fig. 22). Warmer colors (reds) indicate higher vertebrate species richness (higher alpha diversity), cooler colors (blues) lower vertebrate species richness (lower alpha diversity). This is a good indicator of potential

species richness of a habitat type and possibly the relative habitat quality. Vertebrate

species richness is also mapped at the hexagon resolution of 24 km2

 (fig. 23) and

colors reflect the number of habitats for each hexagon and the relative beta diversity, or difference in vertebrate species composition between habitats (Whittaker

1997). Richness per pixel (alpha diversity) and per hexagon (beta diversity) may

show very different patterns. Mountain forested habitats tend to be more species

rich (higher alpha diversity), but coastal regions, with greater heterogeneity of

habitats, may have higher beta diversity.

Amphibian species richness patterns show higher diversity in the forested

mountain regions (figs. 24 and 25). Bird species richness patterns (figs. 26 and 27)

show higher diversity in coastal areas with a mix of open and closed forest habitats.

Mammal species richness (primarily bats) patterns indicate greater richness in the

northern limestone region where karst topography and caves occur (figs. 28 and

29). Reptile species richness (alpha diversity) is higher in the forested areas; beta

diversity for reptiles is highest in areas with a mix of either wet or dry forests or a

matrix of forest and grassland (figs. 30 and 31).
53

The Puerto Rico Gap Analysis Project

Figure 22—Predicted resident, endemic, and endangered terrestrial vertebrate species richness per 15-m2 pixel.

Figure 23—Predicted terrestrial vertebrate resident, endemic, and endangered species richness per 24-km2 hexagon.
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GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 24—Predicted amphibian species richness per 15-m2 pixel.

Figure 25—Predicted amphibian species richness per 24-km2 hexagon.
55

The Puerto Rico Gap Analysis Project

Figure 26—Predicted bird species richness per 15-m2 pixel.

Figure 27—Predicted bird species richness per 24-km2 hexagon.
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GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 28—Predicted mammal species richness per 15-m2 pixel.

Figure 29—Predicted mammal species richness per 24-km2

 hexagon.
57

The Puerto Rico Gap Analysis Project

Vertebrate Distribution Accuracy Assessment

Assessing the accuracy of the predicted vertebrate distributions is subject to many

of the same problems as assessing land cover maps, as well as a host of more

serious challenges related to both the behavioral aspects of species and the logistics

of detecting them. These are described further in the “Background” section of the

GAP handbook on the national GAP home page. It is, however, necessary to

provide some measure of confidence in the results of the gap analysis for all species

collectively, if not individually or by taxonomic group (comparison to stewardship

Figure 30—Predicted reptile species richness per 15-m2 pixel.

Figure 31—Predicted reptile species richness per 24-km2 hexagon.
58

GENERAL TECHNICAL REPORT IITF-GTR-39

Tomás A. Carlo

The endemic Puerto Rican Tody Todus mexicanus can be seen in the island’s Central Mountains.
59

The Puerto Rico Gap Analysis Project

and management status), and to allow users to judge the suitability of the distribution maps for their own uses. We, therefore, feel it is important to provide users

with a statement about the accuracy of predicted vertebrate distributions within

the limitations of available resources and practicalities of such an endeavor. We

acknowledge that distribution maps are never finished products but are continually

updated as new information is gathered. This reflects not only an improvement

over the modeling process, but also the opportunity to map true changes in species

distributions over time. However, we feel that assessing the accuracy of the current

maps provides useful information about their reliability to potential users.

Our goal was to produce maps that predict distribution of terrestrial vertebrates

and, from that, total species richness and species content with an accuracy of 80

percent or higher. Failure to achieve this accuracy indicates the need to refine the

data sets and models used for predicting distribution.

Vertebrate Data Limitations

We compiled all available information on species occurrences and obtained expert

review on the majority of species distributions. We did not set aside a data set for

accuracy assessment, but as new information and site species lists become available

it would be advisable to conduct such an accuracy assessment.

Land Stewardship

To fulfill the analytical mission of GAP, we compare the mapped distribution of

elements of biodiversity with different categories of land ownership and management. These comparisons are a start in assessing the likelihood of future threat to

a biotic element through habitat conversion—the primary cause of biodiversity

decline. We use the term “stewardship” in place of “ownership” in recognition that

legal ownership does not necessarily equate to the entity charged with management

of the resource, and that the mix of ownership and managing entities is a complex

and rapidly changing condition not suitably mapped by GAP. At the same time, it

is necessary to distinguish between stewardship and management status in that a

single category of land stewardship, such as a national forest, may contain several

degrees of management for biodiversity.

The purpose of comparing biotic distribution with stewardship is to provide a

method by which land stewards can assess their relative amount of responsibility

for the management of a species or plant community and identify other stewards
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GENERAL TECHNICAL REPORT IITF-GTR-39

sharing that responsibility. This information can reveal opportunities for

cooperative management of that resource, which directly supports the primary

mission of GAP to provide objective, scientific information to decisionmakers and

managers to make informed decisions regarding biodiversity. It is likely that a

steward that has previously borne the major responsibility for managing a species

may, through such analyses, identify a more equitable distribution of that responsibility. We emphasize, however, that GAP only identifies private land as a homogeneous category and does not differentiate individual tracts or owners, unless the

information was provided voluntarily to recognize a long-term commitment to

biodiversity maintenance.

After comparison to stewardship, it is also necessary to compare biotic occurrence to categories of management status. The purpose of this comparison is to

identify the need for change in management status of individual elements or areas

containing high degrees of diversity. Such changes can be accomplished in many

ways that do not affect the stewardship status. Although it will eventually be desirable to identify specific management practices for each tract and whether they are

beneficial or harmful to each element, GAP currently uses a scale of 1 to 4 to

denote relative degree of maintenance of biodiversity for each tract. A status of “1”

denotes the highest, most permanent level of maintenance, and “4” represents the

lowest level of biodiversity management, or unknown status. This is a highly

subjective area, and we recognize several limitations in our approach, although we

maintain certain principles in assigning the status level. Our first principle is that

land ownership is not the primary determinant in assigning status. The second

principle is that although data are imperfect and all land is subject to changes in

ownership and management, we can use the intent of a land steward as evidenced

by legal and institutional factors to assign status. In other words, if a land steward

institutes a program backed by legal and institutional arrangements that are intended for permanent biodiversity maintenance, we use that as the guide for

assigning status.

The characteristics used to determine status are as follows:

• Permanence of protection from conversion of natural land cover to

unnatural (human-induced barren, exotic-dominated, arrested succession).

• Relative amount of the tract managed for natural cover.

• Inclusiveness of the management, i.e., single feature or species versus all

biota.

• Type of management and degree that it is mandated through legal and

institutional arrangements.
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The Puerto Rico Gap Analysis Project

The four status categories can generally be defined as follows (after Crist et al.

1995, Edwards et al. 1995, Scott et al. 1993,):

Status 1: An area having permanent protection from conversion of natural land

cover and a mandated management plan in operation to maintain a natural state

within which disturbance events (of natural type, frequency, and intensity) are

allowed to proceed without interference or are mimicked through management.

Status 2: An area having permanent protection from conversion of natural

land cover and a mandated management plan in operation to maintain a primarily

natural state, but which may receive use or management practices that degrade the

quality of existing natural communities.

Status 3: An area having permanent protection from conversion of natural land

cover for the majority of the area, but subject to extractive uses of either a broad,

low-intensity type or localized intense type. It also confers protection to federally

listed endangered and threatened species throughout the area.

Status 4: Lack of irrevocable easement or mandate to prevent conversion of

natural habitat types to anthropogenic habitat types. Allows for intensive use

throughout the tract. Also includes those tracts for which the existence of such

restrictions or sufficient information to establish a higher status is unknown.

Mapping Standards

The PRGAP land stewardship map is a 1:100,000-scale version of land ownership

and management of Puerto Rico’s protected areas, distinguishing local, state, and

federal jurisdictions from private lands and delineating areas managed for the

long-term maintenance of natural ecological processes and biodiversity. This

layer therefore contains attributes for ownership, management, and for the level of

biodiversity protection. We used the Lambert Conformal Conic projection, the

North American 1983 Grid Coordinate System, and the North American Datum

(NAD) 1983 State Plane Puerto Rico Virgin Islands Federal Information Processing

Standard (FIPS) 5200 datum. The purpose of the PRGAP land stewardship is to

provide a complete digital map of protected areas of Puerto Rico describing land

ownership and land management. These data are intended to aid in state-level

assessment of natural resources and are not intended for use at a scale greater than

1:100,000.
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GENERAL TECHNICAL REPORT IITF-GTR-39

Stewardship Mapping Methods

Stewardship mapping—

Protected area data sets were obtained from various GIS coordinators in government

and nongovernmental agencies in Puerto Rico; the DNER, USFS, USFWS, the

Conservation Trust of Puerto Rico, and the Puerto Rico Planning Board. These data

sets were provided via disks, e-mail, downloaded from agency electronic bulletin

boards, or as hard copy from management plans. Data sets were received in different formats: ArcView1

 shapefiles, ArcInfo coverages, and Arc export files. The data

were brought into the land stewardship ArcGIS geodatabase of the PRGAP. After

the data were converted to common projections, edge matching and editing were

performed. Final mapping was done using ArcGIS 9.0.

Quality control was in the form of site interviews with all land managers, at

which time a review of stewardship boundaries was conducted. Suggestions of

boundary changes were incorporated when documentation was available from

agencies to confirm the corrections.

Management status categorization—

Initially, we used information from the protected area’s management plans in order

to assign a GAP status code to each land unit. However, only 7 of the 77 protected

areas have current management plans. The Puerto Rico Forest Bureau of the DNER

is in the process of writing management plans for the state forests. Five additional

plans were in development, and three were expired at the time of our interviews.

Old management plans do not correctly describe current management practices.

The Master Plan for the Commonwealth Forests of Puerto Rico (DRNA 1976)

was developed for the state forests when they were being managed as multiple-use

forests for timber production, resource management, and recreation. Today, the

forests are not producing timber or wood products as in the past, although a small

fraction of what was being produced in the 1970s and 1980s is still being used for

craft products and fence posts for the state forests themselves. The forests are also

seed sources for many of the DNER tree nurseries, which provide trees for reforestation projects on private lands and the forests. The forests are also being managed

for biodiversity protection. As there were few active management plans to refer to

for information, we conducted one-on-one interviews with land managers. The site

visits made it possible to gather information on the land unit’s boundaries, species

1

 The use of trade or firm names in this publication is for reader information and does not

imply endorsement by the U.S. Department of Agriculture of any product or service.
63

The Puerto Rico Gap Analysis Project

The Sierra palm Prestoea montana is abundant on the steep slopes of the Luquillo Mountains in northeastern Puerto

Rico.

Mariano Solórzano
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GENERAL TECHNICAL REPORT IITF-GTR-39

occurrence records, research areas, areas open to the public, threats (urban encroachment, invasive species, trespassing destruction of land, pollution, exotic

species), and needs of the protected area managers to better reach their objectives.

Stewardship Mapping Results

We identified 90 stewardship areas for Puerto Rico. Seventy-seven stewardship

areas have some management for conservation (table 8). Land ownership of these

areas is shared among 20 organizations (table 9, fig. 32) with the DNER being

the primary landowner. Management of land stewardship areas is primarily shared

among 13 organizations (figs. 33 and 34, table 10) with the DNER, the USFS, and

the USFWS being the primary governmental land managers and the Conservation

Trust of Puerto Rico being the primary nongovernmental land manager.

Altogether, 7.6 percent of Puerto Rico has some management for conservation

(GAP status 1, 2, or 3) and 7.4 percent of Puerto Rico has good management of

conservation (GAP status 1 or 2) (fig. 35). Most of this management is under GAP

status 2, management for natural resources, and this includes most of the commonwealth forests and reserves. Areas with a priority on biodiversity conservation

(GAP status 1) include two management areas in the Luquillo Experimental Forest

(Caribbean National Forest): The Baño del Oro and the El Toro Wilderness area

(figs. 35 and 36). Fifty-nine percent of the stewardship areas are managed by

commonwealth agencies, 30 percent by federal agencies, and 11 percent by nongovernmental or private agencies (fig. 37).

Stewardship Map Limitations

These data and maps are a compilation of ownership maps provided by a variety

of sources that are individually responsible for their accuracy. They were created

solely for the purpose of conducting the analyses described in this report and are

not suitable for locating boundaries on the ground or determining precise area

measurements of individual tracts. Land ownership, management, and land use are

very dynamic. At the writing of this report, several areas were under consideration

for either development, conservation, or some combination of these activities. This

includes the former naval bases at Roosevelt Roads and Sabana Seca, public and

private lands in the Northeast Ecological Corridor, the Piñones area, a potential

ecological corridor linking the central mountain reserves with the northern karst

and coastal plain, and the ecologically valuable northern karst region.
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The Puerto Rico Gap Analysis Project

Table 8—Stewardship areas in Puerto Rico, area and percentage of total by management class

Gap status

Area name Class 1 2 3 4 Total

Ha % Ha % Ha % Ha % Ha %

Management for biodiversity conservation Area Natural Protegida Cañón de San Cristóbal Nongovernmental 497 0.9 497 < 0.1 Area Natural Protegida Jorge Sotomayor del Toro Nongovernmental 16 < 0.1 16 < 0.1 Area Natural Protegida Rio Encantado Nongovernmental 284 0.5 284 < 0.1 Bosque de Pterocarpus de Dorado Nongovernmental 12 0.1 12 < 0.1 Bosque del Pueblo de Adjuntas Nongovernmental 418 0.8 418 < 0.1 Bosque Estatal de Aguirre Commonwealth 432 0.8 432 < 0.1 Bosque Estatal de Boquerón Commonwealth 623 5.7 623 < 0.1 Bosque Estatal de Cambalache Commonwealth 649 1.2 649 < 0.1 Bosque Estatal de Carite Commonwealth 2700 4.9 2700 0.3 Bosque Estatal de Ceiba Commonwealth 119 0.2 118 5.5 237 < 0.1 Bosque Estatal de Cerrillos Commonwealth 220 0.4 220 < 0.1 Bosque Estatal de Guajataca Commonwealth 955 1.7 955 0.1 Bosque Estatal de Guánica Commonwealth 3832 6.9 3832 0.4 Bosque Estatal de Maricao Commonwealth 4168 7.6 4168 0.5 Bosque Estatal de Monte Guilarte Commonwealth 1705 3.1 1705 0.2 Bosque Estatal de Piñones Commonwealth 732 1.3 732 < 0.1 Bosque Estatal de Río Abajo Commonwealth 2284 4.1 2284 0.3 Bosque Estatal de Susúa Commonwealth 1298 2.4 1298 0.1 Bosque Estatal de Toro Negro Commonwealth 2763 5.0 2763 0.3 Bosque Estatal de Tres Picachos Commonwealth 2097 3.8 2097 0.2 Bosque Estatal de Vega Commonwealth 482 0.9 482 < 0.1 Bosque La Olimpia Nongovernmental 185 0.3 185 < 0.1 Bosque Urbano de San Patricio Commonwealth 40 0.1 40 < 0.1 Bosque Urbano del Nuevo Milenio Commonwealth 176 0.3 176 < 0.1 Bosque Urbano Doña Inez Mendoza Nongovernmental 99 4.6 99 < 0.1 Cabo Rojo National Wildlife Refuge Federal 758 35.1 758 < 0.1 Camp Santiago U.S. Military Reservation Federal

/nongovernmental 2916 5.3 2213 0.3 5129 0.6

 Camuy Caves Park Commonwealth 104 4.8 104 < 0.1 Corredor Ecologico de San Juan Commonwealth 24 1.1 24 < 0.1 Culebra National Wildlife Refuge Federal 603 5.5 603 < 0.1 Desecheo National Wildlife Refuge Federal 131 0.2 131 < 0.1 El Yunque National Forest Federal 7162 65.2 4267 7.7 11 429 1.3
66

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 8—Stewardship areas in Puerto Rico, area and percentage of total by management class (continued) Gap status Area name Class 1 2 3 4 Total Ha % Ha % Ha % Ha % Ha % Estación Experimental Agrícola Commonwealth 22 0.2 22 < 0.1 Finca Guillermeti Nongovernmental 195 0.4 195 < 0.1 Jardín Botánico de la Universidad de Puerto Rico Commonwealth 125 5.8 125 < 0.1 Laguna Cartagena National Wildlife Refuge Federal 109 0.2 314 14.6 423 < 0.1 Parque Nacional Sun Bay Commonwealth 23 1.1 23 < 0.1 Refugio de Vida Silvestre de Boquerón Commonwealth 235 0.4 235 < 0.1 Refugio de Vida Silvestre Lago Guajataca Commonwealth 272 12.6 272 < 0.1 Refugio de Vida Silvestre Lago La Plata Commonwealth 188 8.7 188 < 0.1

 Refugio de Vida Silvestre Lago Luchetti Commonwealth 94 4.4 94 < 0.1 Reserva Nacional de Investigación Estuarina

de Bahía de Jobos Commonwealth 414 3.8 35 0.1 449 < 0.1

 Reserva Natural Aguas Buenas Commonwealth 709 6.5 709 < 0.1 Reserva Natural Arrecifes de la Cordillera Commonwealth 21 0.2 63 0.1 4 0.2 88 < 0.1 Reserva Natural Bahía Ballena Nongovernmental 97 0.2 97 < 0.1 Reserva Natural Bosque de Pterocarpus de Humacao Nongovernmental 285 2.6 285 < 0.1 Reserva Natural Caja de Muertos Commonwealth 188 0.3 188 < 0.1 Reserva Natural Caño La Boquilla Commonwealth 70 0.6 70 < 0.1 Reserva Natural Caño Martin Peña Commonwealth 107 0.2 107 < 0.1 Reserva Natural Caño Tiburones de Arecibo Commonwealth 1933 3.5 1933 0.2

 Reserva Natural Cerro El Buey Nongovernmental 323 0.6 323 < 0.1 Reserva Natural Cueva del Indio Commonwealth 6 0.0 6 < 0.1 Reserva Natural de Humacao Commonwealth 898 1.6 898 0.1 Reserva Natural de Isla de Mona Commonwealth 15 0.1 5671 10.3 5686 0.6 Reserva Natural de la Bahía Bioluminiscente

de Vieques Commonwealth 414 0.7 414 < 0.1

 Reserva Natural de la Finca Seven Seas Commonwealth 83 0.8 83 < 0.1 Reserva Natural de La Parguera Nongovernmental 408 3.7 408 < 0.1 Reserva Natural de Laguna de Joyuda Commonwealth 178 0.3 178 < 0.1 Reserva Natural de Punta Guaniquilla Commonwealth 15 0.0 15 < 0.1 Reserva Natural del Río Espíritu Santo Commonwealth 871 1.6 871 < 0.1 Reserva Natural Finca Belvedere Commonwealth 162 0.3 162 < 0.1 Reserva Natural Hacienda Buena Vista Nongovernmental 35 0.1 35 < 0.1 Reserva Natural Hacienda la Esperanza Nongovernmental 924 1.7 924 0.1
67

The Puerto Rico Gap Analysis Project

 Table 8—Stewardship areas in Puerto Rico, area and percentage of total by management class (continued)

Gap status

Area name Class 1 2 3 4 Total

Ha % Ha % Ha % Ha % Ha %

Reserva Natural Inés María Mendoza (Punta Yeguas) Nongovernmental 130 1.2 130 < 0.1

 Reserva Natural La Cienaga Las Cucharillas Commonwealth 396 0.7 396 < 0.1 Reserva Natural La Parguera Commonwealth 23 < 0.1 23 < 0.1 Reserva Natural Laguna Tortuguero Commonwealth 273 2.5 992 1.8 1265 0.1

 Reserva Natural Las Cabezas de San Juan Nongovernmental 225 0.4 225 < 0.1 Reserva Natural Las Piedras del Collado Commonwealth 8 0.1 8 < 0.1 Reserva Natural Mata de Platano Field Station Nongovernmental 144 1.3 144 < 0.1 Reserva Natural Pantano de Cibuco Commonwealth 267 0.5 267 < 0.1 Reserva Natural Punta Guaniquilla Nongovernmental 166 0.3 166 < 0.1 Reserva Natural Punta Petrona Commonwealth 175 0.3 175 < 0.1 Reserva Natural Punta Tuna Commonwealth 52 0.1 52 < 0.1 San Juan National Historic Site Federal 32 1.5 32 < 0.1 Servidumbre Escenica Río Portugues Nongovernmental 14 < 0.1 14 < 0.1 Vieques National Wildlife Refuge Federal 7070 12.8 7070 0.8

No management for biodiversity conservation

 Fort Allen Naval Radio Station Federal 366 < 0.1 366 < 0.1 Fort Buchanan Military Reservation Federal 276 < 0.1 276 < 0.1 Isla Grande Airport Federal 102 < 0.1 102 < 0.1 Parque Nacional Punta Guilarte Commonwealth 101 < 0.1 101 < 0.1 Ramey Air Force Base Federal 1102 0.1 1102 0.1 Reserva Natural Torrecilla Alta (proposed) Commonwealth 1599 0.2 1599 0.2 Roosevelt Roads Naval Reservation Federal 3483 0.4 3483 0.4 Sabana Seca U.S. Naval Radio Station Federal 920 0.1 920 0.1 Sierra Bermeja Private 899 0.1 899 0.1 U.S. Military Reservation Aguadilla Federal 459 < 0.1 459 < 0.1 U.S. Naval Radio Transmitter Facility Aguada Federal 164 < 0.1 164 < 0.1 U.S. Naval Radio Transmitter Facility Aguadilla Federal 156 < 0.1 156 < 0.1 Non-Stewardship Areas Private/other 814 730 98.6 814 730 91.0

Total 10 983 100 55 205 100 2156 100 826 571 100 894 913 100

Note: Seventy-seven of the 90 stewardship areas have some management for biodiversity conservation (GAP status 1 through 3). Thirteen areas have no management for biodiversityconservation and are either proposed for future conservation management or have other civilian and military uses.
68

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 9—Landowners of stewardship areas, area and percentage of total by management class Gap status Owner Class 1 2 3 4 Total Ha % Ha % Ha % Ha % Ha % ACT Commonwealth 175 0.3 175 < 0.1 Citizens of the Karst Foundation Nongovernmental 144 1.3 195 0.4 341 < 0.1 Commonwealth and other public or private entities Commonwealth 861 1.6 23 886 < 0.1 Conservation Trust of Puerto Rico Nongovernmental 835 7.6 2483 4.5 3330 0.4 Department of Natural and Environmental Resources Commonwealth 1932 17.6 29 665 53.7 23 1.1 31 694 3.5 Department of Natural and Environmental

Resources and the Conservation Trust of Commonwealth/Puerto Rico nongovernmental 97 0.2 97 < 0.1

Electrical Energy Authority Commonwealth 173 0.3 555 26.0 754 < 0.1

Industrial Development Company Commonwealth 63 0.1 4 0.2 67 < 0.1Land Administration Commonwealth 273 2.5 1993 3.6 340 15.9 2629 0.3Land Authority Commonwealth 22 0.2 4436 8.0 80 3.7 1700 0.2 6250 0.7Luis Muñóz Marín Foundation 99 4.7 104 < 0.1National Parks Company of Puerto Rico Commonwealth 414 0.7 415 < 0.1U.S. Army Corps of Engineers Federal 220 0.4 220 < 0.1U.S. Department of Defense Federal 2916 5.3 118 5.5 5758 0.7 8802 1.0U.S. Fish and Wildlife Service Federal 575 5.2 7202 13.0 758 35.5 8588 1.0

U.S. Forest Service Federal 7162 65.2 4267 7.7 11 502 1.3

U.S. National Park Service Federal 32 1.5 34 < 0.1U.S. Navy Federal 45 < 0.1 3483 0.4 3528 0.4University of Puerto Rico Commonwealth 11 0.1 125 5.8 141 < 0.1Private/other Private 28 0.3 815 329 98.7 815 329 91.1

Total 10 983 100 55 204 100 2134 100 826 294 100 894 913 100
69

The Puerto Rico Gap Analysis Project Figure 32—Landowners of stewardship areas in Puerto Rico (Gould et al. 2008e).
70

GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 33—Primary land managers and the number of hectares managed in Puerto Rico under GAP

management status 1, 2, 3, or 4. Entities with clear bars have no management for conservation (GAP

status 4). Entities with dark bars are in part or all managed for conservation (GAP status 1 through

3). Note the scale is logarithmic. DNER = Puerto Rico Department of Natural and Environmental

Resources, ELAPR = Estado Libre Associado de Puerto Rico (the commonwealth government),

NOAA = the National Oceanic and Atmospheric Administration.
71

The Puerto Rico Gap Analysis Project Figure 34—Land management of stewardship areas in Puerto Rico (Gould et al. 2008f).
72

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 10—Land managers of stewardship areas, area and percentage of total Gap status Manager 1 2 3 4 Total

Ha % Ha % Ha % Ha % Ha %

Citizens of the Karst Foundation 144 1.31 195 0.36 341 < 0.1Industrial Development Company 101 < 0.1 101 < 0.1Conservation Trust of Puerto Rico 705 6.42 2580 4.73 3296 0.37Conservation Trust of Puerto Rico and Department

of Natural and Environmental Resources 130 1.18 131 < 0.1

Department of Natural and Environmental

Resources 1802 16.41 36 847 67.48 559 25.91 39 318 4.40

Department of Natural and Environmental

Resources and NOAA 414 3.77 35 < 0.1 453 < 0.1

Commonwealth of Puerto Rico 3483 0.42 3483 0.39

Land Administration 24 1.12 25 < 0.1Land Authority 1599 0.19 1599 0.18Luis Muñóz Marín Foundation 99 4.61 104 < 0.1National Parks Company of Puerto Rico 406 0.74 128 5.92 541 < 0.1Other nongovernmental entities 2916 5.34 2921 0.33Other private entities 899 0.11 899 0.10University of Puerto Rico 22 0.20 125 5.77 153 < 0.1U.S. Air Force 1102 0.13 1102 0.12U.S. Army 2673 0.32 2673 0.30U.S. Fish and Wildlife Service 603 5.49 7311 13.39 1072 49.72 9054 1.01

U.S. Forest Service 7162 65.21 4267 7.81 11 502 1.29U.S. National Park Service 32 1.49 34 < 0.1U.S. Navy 45 < 0.1 118 5.45 1983 0.24 2151 0.24

None or unknown 815 033 98.57 815 033 91.06

Total 10 983 100 54 601 100 2156 100 826 874 100 894 913 100
73

The Puerto Rico Gap Analysis Project Figure 35—Priority given to conservation (GAP status levels 1 through 4) in stewardship areas in Puerto Rico (Gould et al. 2008g).
74

GENERAL TECHNICAL REPORT IITF-GTR-39

Figure 36—Number of hectares and managing agencies in GAP status 1, 2, 3, and 4

for Puerto Rico. Note scale is logarithmic. NGO = nongovernmental organization.

Figure 37—Number of hectares of areas with some management for biodiversity

conservation (GAP status 1 through 3), by managing agency.
75

The Puerto Rico Gap Analysis Project

Analysis Based on Stewardship and

Management Status

This section describes the method and results of the gap analysis as used by the Gap

Analysis Program. The primary objective of GAP is to provide information on the

distribution and status of elements of biological diversity. Although GAP seeks to

identify habitat types and species not adequately represented in the current network

of biodiversity management areas, a standard definition of “adequate representation” for land cover types or individual species is difficult (Noss et al. 1995). A

practical solution to this problem is to report both percentages and absolute area of

each element in protected areas and allow the user to determine which classes are

adequately represented. There are many other factors that should be considered in

such determinations:

• Loss or gain in historical distribution.

• Nature of the spatial distribution.

• Immediate versus long-term risk.

• Degree of local adaptation among populations of the biotic elements that

are worthy of individual conservation consideration.

Such analyses are beyond the scope of this project, but we encourage their

application coupled with field confirmation of the mapped distributions.

Currently, land cover types and terrestrial vertebrates are the primary focus of

GAP’s mapping efforts; however, other components of biodiversity, such as aquatic

organisms or selected groups of invertebrates may be incorporated into GAP distributional data sets. Where appropriate, GAP data may also be analyzed to identify

the location of a set of areas in which most or all land cover types or species are

predicted to be represented. The use of “complementarity” analysis, that is, an

approach that additively identifies a selection of locations that together may represent biodiversity rather than “hot spots of species richness” may prove most effective for guiding biodiversity maintenance efforts. Several approaches have been

developed that facilitate this process (Csuti et al. 1997, Jennings 2000, Peterson and

Klusa 2003, Pressey et al. 1993, Sowa et al. 2007, Williams et al. 1996, Vierling et

al. 2007). These areas become candidates for field validation and may be incorporated into a system of areas managed for the long-term maintenance of biological

diversity.

The network of Conservation Data Centers (CDCs) and Natural Heritage

Programs established cooperatively by The Nature Conservancy and various state

agencies maintain detailed databases on the locations of rare elements of
76

GENERAL TECHNICAL REPORT IITF-GTR-39

biodiversity. The GAP cooperatively uses these data to develop predicted distributions of potentially suitable habitat for these elements, which may be valuable for

identifying research needs and preliminary considerations for restoration or reintroduction. Conservation of such elements, however, is best accomplished through the

fine-filter approach of the above organizations as described in the introduction. It is

not the role of GAP to duplicate or disseminate Heritage Program or CDC element

occurrence records. Users interested in more specific information about the location, status, and ecology of populations of such species are directed to their state

Heritage Program or CDC.

Analysis Methods

The gap analysis is accomplished by first producing maps of land cover (fig. 12),

predicted distributions for selected animal species (fig. 15 and see volume 4), and

land stewardship and management status (fig. 35). Intersecting the land stewardship

and management map with the distribution of the elements results in tables that

summarize the area and percentage of total mapped distribution of each element in

different land stewardship and management categories. We created a raster image of

the stewardship GAP status categories 1 through 4 using ERDAS Imagine 9.1 and

intersected this with our land cover classes and with our individual species predicted habitat distributions. The resulting output includes the number of hectares of

each species or land cover unit in each GAP status category.

Analysis Results

The data are provided in a format that allows users to carry out inquiries about

the representation of each element in different land stewardship and management categories as appropriate to their own management objectives. This forms the

basis of GAP’s mission to provide landowners and managers with the information

necessary to conduct informed policy development, planning, and management for

biodiversity maintenance.

Land cover analysis—

Table 11 provides the area in hectares of each mapped cover type by status and the

percentage of the total area in that cover type. For example, a typical entry indicates

that freshwater Pterocarpus swamp covers 261 ha and that 172 ha is ranked status 1

or 2, which represents 66.1 percent of the 261 ha in that type.
77

The Puerto Rico Gap Analysis Project

Mangrove forests play an important role in harboring fish, crabs, and birds along Puerto Rico’s coastlines.

Mariano Solórzano
78

GENERAL TECHNICAL REPORT IITF-GTR-39

As a coarse indicator of the status of the elements, we provide a breakdown

along five levels of protection (0 to <1 percent, 1 to <10 percent, 10 to <20 percent, 20 to <50 percent, ≥50 percent). The <1 percent level indicates those elements with essentially none of their distribution in a protected status; while levels

of 10, 20, and 50 percent have been recommended in the literature as necessary

amounts of conservation (McNeely and Miller 1994, Noss and Cooperrider 1994,

Odum and Odum 1972, Ride 1975). Summaries of the analysis according to the

thresholds described above are shown in table 11.

Land cover with <1 percent in GAP status 1 or 2—

These eight land cover classes range from the smallest to largest in extent. They

make up 43 percent of the island and are primarily subject to intensive human use

such as agriculture, housing, and development. The class of moist grasslands and

pastures covers nearly one-fourth of the island and is primarily abandoned agricultural land. Given the resilience of the natural vegetation in Puerto Rico, this land

cover type has potential for management for reforestation or as natural grasslands.

As a group, 0.7 percent is protected under GAP status 1 or 2.

Land cover with 1 to <10 percent in GAP status 1 or 2—

Twenty-seven land cover units fall in this category and they account for 44 percent

of the island. They range from an extent of less than 1 to over 6 percent of the

island. They contain a number of young secondary forest and woodland land cover

classes, as well as artificial and natural barrens, active and abandoned shade coffee

plantations, dry grasslands and pastures, riparian forests, and four mature secondary

forest classes. As a group, 3.8 percent of these units are protected under GAP status

1 or 2.

Land cover with l0 to <20 percent in GAP status 1 or 2—

Four land unit classes fall in this category and they account for 1.7 percent of the

island. They include two woodland or shrubland classes–which usually occur on

abandoned agricultural land, a dryland riparian forest, and palm plantations. For the

group, 16.2 percent is protected under GAP status 1 or 2.

Land cover with 20 to <50 percent in GAP status 1 or 2—

Fourteen land cover classes fall in this group, and they account for 6.1 percent of

the island. They include a number of ecologically important areas including beaches

and shorelines, mature forests, wetlands, mangrove complexes, and Sierra palm

forest. For the group, 36.5 percent is protected under GAP status 1 or 2.
79

The Puerto Rico Gap Analysis Project

The Mona Island rock iquana Cyclura stejnegeri is endemic to Mona Island

Nestor Pérez
80

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 11—Area in hectares, percentage of total, and protection category for each land cover type Total Protection Landcover type habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category Ha % Ha % Ha % Ha % Ha % Ha % Puerto Rico 894 912 100 66 275 7.4 11 240 1.3 55 035 6.1 2253 0.3 826 385 92 4 Dry forest, woodland, and shrubland Mature secondary lowland dry alluvial semideciduous forest 1548 0.2 732 47.3 32 2.0 700 45.3 1 0.1 814 52.6 2 Young secondary lowland dry alluvial semideciduous forest 2881 0.3 62 2.2 5 0.2 57 2.0 14 0.5 2805 97.4 4 Lowland dry alluvial shrubland and woodland 4472 0.5 299 6.7 33 0.7 266 6.0 166 3.7 4007 89.6 4 Lowland dry riparian forest 806 0.1 120 14.9 6 0.8 114 14.1 0 0 686 85.1 3 Lowland dry riparian shrubland and woodland 423 0.5 15 3.5 1 0.3 14 3.2 0 0 408 96.5 4

Mature secondary lowland dry limestone

evergreen forest 982 0.1 207 21.1 0 0 207 21.1 0 0 776 78.9 2

Mature secondary lowland dry limestone

semideciduous forest 10 669 1.2 3871 36.3 33 0.3 3838 36.0 3 0 6795 63.7 2

Young secondary lowland dry limestone

semideciduous forest 3922 0.4 163 4.2 23 0.6 140 3.6 13 0.3 3745 95.5 4

Lowland dry limestone woodland and

shrubland 8181 0.9 3535 43.2 40 0.5 3495 42.7 0 0 4645 56.8 2

Lowland dry limestone shrubland 4694 0.5 1336 28.5 66 1.4 1270 27.1 64 1.4 3294 70.2 2Lowland dry cactus shrubland 47 0 47 100 0 0 47 100 0 0 0 0.0 1Coastal dwarf woodland and shrubland 103 0 103 100 0 0 103 100 0 0 0 0.0 1Lowland dry limestone cliffside semideciduous

forest 12 0 12 100 0 0 12 100 0 0 0 0.0 1

Lowland dry limestone cliffside shrubland

and woodland 49 0 49 100 0 0 49 100 0 0 0 0.0 1

Abandoned dry forest plantation 81 0 81 100 0 0 81 100 0 0 0 0.0 1Mature secondary lowland dry noncalcareous

semideciduous forest 7062 0.8 3818 54.1 319 4.5 3499 49.5 0 0 3244 45.9 1

Young secondary lowland dry noncalcareous

semideciduous forest 3644 0.4 285 7.8 4 0.1 281 7.7 1 0 3358 92.2 4

Lowland dry noncalcareous shrubland and

woodland 13 307 1.5 2214 16.6 203 1.5 2011 15.1 6 0.1 11 086 83.3 3

Mature secondary dry and moist serpentine

semideciduous forest 1839 0.2 379 20.6 0 0 379 20.6 0 0 1460 79.4 2

Young secondary dry and moist serpentine

semideciduous forest 1960 0.2 1081 55.1 0 0 1081 55.1 0 0 879 44.9 1

Dry and moist serpentine woodland and shrubland 841 0.1 86 10.2 0 0 86 10.2 0 0 755 89.8 3
81

The Puerto Rico Gap Analysis Project

Table 11—Area in hectares, percentage of total, and protection category for each land cover type (continued)

Total Protection

Landcover type habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category

Ha % Ha % Ha % Ha % Ha % Ha %

Moist forest, woodland, and shrublandMature secondary lowland moist alluvial evergreen

forest 2256 0.3 165 7.3 8 0.4 157 7.0 1 0 2091 92.6 4

Young secondary lowland moist alluvial evergreen

forest 6675 0.8 316 4.7 13 0.2 303 4.5 10 0.1 6350 95.1 4

Lowland moist alluvial shrubland and woodland 5141 0.6 128 2.5 7 0.1 121 2.4 6 0.1 5007 97.4 4Lowland moist riparian forest 916 0.1 48 5.3 3 0.3 46 5.0 6 0.7 862 94.0 4Lowland moist riparian shrubland and woodland 508 0.1 30 6.0 0 0 30 6.0 0 0 478 94.0 4Mature secondary moist limestone evergreen and

semideciduous forest 46 423 5.2 4370 9.4 121 0.3 4249 9.4 4 0 42 049 90.6 4

Young secondary moist limestone evergreen and

semideciduous forest 15 494 1.7 169 1.1 5 0 164 1.1 94 0.6 15 231 98.3 4

Moist limestone shrubland and woodland 10 338 1.2 298 2.9 10 0.1 288 2.8 6 0.1 10 034 97.1 4Mature secondary lowland moist noncalcareous

evergreen forest 40 446 4.5 1305 3.2 255 0.6 1050 2.6 0 0 39 140 96.8 4

Young secondary lowland moist noncalcareous

evergreen forest 52 980 5.9 438 0.8 161 0.3 277 0.5 110 0.2 52 432 99.0 5

Lowland moist noncalcareous shrubland and

woodland 44 523 5.0 461 1.0 100 0.2 361 0.8 28 0.1 44 034 98.9 4

Lowland moist abandoned and active coffee

plantations 11 068 1.2 121 1.1 23 0.2 98 0.9 0 0 10 947 98.9 4

Wet forest, woodland, and shrublandMature secondary montane wet alluvial evergreen

forest 613 0.1 139 22.7 10 1.6 130 21.1 0 0 474 77.3 2

Young secondary montane wet alluvial evergreen

forest 988 0.1 17 1.7 1 0.1 16 1.6 0 0 971 98.3 4

Montane wet alluvial shrubland and woodland 900 0.1 6 0.7 0 0 6 0.7 0 0 894 99.3 5Mature secondary montane wet noncalcareous

evergreen forest 25 207 2.8 642 4.3 0 0 642 4.3 0 0 14 326 95.7 4

Montane wet evergreen abandoned and active

coffee plantation 54 859 6.1 839 1.5 0 0 839 1.5 0 0 54 021 98.5 4

Mature primary and secondary montane wet

noncalcareous evergreen tabonuco forest 8715 1.0 5114 57.8 2028 22.9 3086 34.9 0 0 3742 42.3 1

Mature primary and secondary montane wet

noncalcareous evergreen palo colorado cloudforest 3712 0.4 7870 50.5 3401 21.8 4470 28.7 0 0 7713 49.5 1
82

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 11—Area in hectares, percentage of total, and protection category for each land cover type (continued) Total Protection Landcover type habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category Ha % Ha % Ha % Ha % Ha % Ha % Mature primary and secondary montane wet noncalcareous evergreen sierra palm forest 11 953 1.3 3323 37.8 1388 15.8 1935 22.0 0 0 5458 62.2 2

Mature primary and secondary montane wet

noncalcareous evergreen elfin woodland cloudforest 1537 0.2 1639 55.8 215 7.3 1424 48.5 0 0 1297 44.2 1

Young secondary montane wet noncalcareous

evergreen forest 21 651 2.4 381 1.8 64 0.3 317 1.5 0 0 21 271 98.2 4

Montane wet evergreen noncalcareous shrubland

and woodland 24 090 2.7 469 2.0 45 0.2 424 1.8 0 0 23 621 98.1 4

Mature secondary montane wet serpentine

evergreen forest 3379 0.4 2889 85.5 0 0 2889 85.5 0 0 490 14.5 1

Young secondary montane wet serpentine

evergreen forest 651 0.1 422 64.8 0 0 422 64.8 0 0 230 35.2 1

Wet serpentine shrubland and woodland 358 0 220 61.5 0 0 220 61.5 0 0 138 38.5 1Flooded forest, woodland, and shrublandMangrove forest and shrubland 8700 1.0 3999 46.0 890 10.2 3109 35.7 17 0.2 4683 53.8 2Freshwater Pterocarpus swamp 261 0 172 66.1 75 28.9 97 37.2 88 33.9 0 0.0 1

GrasslandsDry grasslands and pastures 42 176 4.7 2851 6.8 209 0.5 2642 6.3 462 1.1 38 863 92.1 4Dry cactus grassland and shrubland 213 0 202 94.9 0 0 202 94.9 0 0 11 5.1 1Moist grasslands and pastures 218 945 24.5 1984 0.9 304 0.1 1679 0.8 81 0 216 880 99.1 5Herbaceous wetlandsSeasonally flooded herbaceous nonsaline

wetlands 18 225 2.0 894 4.9 106 0.6 788 4.3 163 0.9 17 167 94.2 4

Seasonally flooded herbaceous saline wetlands 4865 0.5 1290 26.5 86 1.8 1204 24.8 1 0 3574 73.5 2Emergent herbaceous nonsaline wetlands 1102 0.1 107 9.7 0 0 107 9.7 39 3.5 956 86.8 4Emergent herbaceous saline wetlands 882 0.1 332 37.6 31 3.6 300 34.1 0 0 550 62.4 2AgricultureHay and row crops 25 764 3.0 179 0.7 25 0.1 154 0.6 9 0 25 576 99.3 5Woody agriculture and plantations 492 0.1 76 15.4 1 0.2 75 15.2 0 0 417 84.6 3
83

The Puerto Rico Gap Analysis Project

Table 11—Area in hectares, percentage of total, and protection category for each land cover type (continued)

Total Protection

Landcover type habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category

Ha % Ha % Ha % Ha % Ha % Ha %

Natural barrensRocky cliffs and shelves 397 0 215 54.1 59 14.7 157 39.4 2 0.5 180 45.4 1Gravel beaches and stony shoreline 82 0 31 37.7 9 10.4 22 27.4 0 0 51 62.3 2Fine to coarse sandy beaches, mixed sand and gravel

beaches 1155 0.1 260 22.5 33 2.9 227 19.7 21 1.8 874 75.7 2

Riparian and other natural barrens 449 0.1 6 1.3 0 0 6 1.32 2 0.4 441 98.2 4Salt and mudflats 1493 0.2 439 29.4 144 9.6 296 19.8 260 17.4 793 53.1 2Developed areasArtificial barrens 8674 1.0 108 1.2 0 0 108 1.2 0 0 8566 98.8 4Salt production 20 0 0 0 0 0 0 0 20 100 0 0.0 5High-density urban development 52 335 5.9 58 0.1 0 0 58 0.1 23 0.04 52 254 99.8 5Low-density urban development 37 237 4.2 225 0.6 30 0.1 194 0.5 33 0.1 36 979 99.3 5WaterFreshwater 4433 0.5 242 5.5 1 0 241 5.4 520 11.7 3671 82.8 4Saline water 4005 0.5 2291 57.2 616 15.4 1675 41.8 64 1.6 1650 41.2 1Aquaculture 101 0 0 0 0 0 0 0 0 0 101 100.0 5Note: Protection category 1 = ≥50 percent protected, 2 = 20 to < 50 percent protected, 3 = 10 to < 20 percent protected, 4 = 1 to < 10 percent protected, 5 = < 1 percent protected.
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GENERAL TECHNICAL REPORT IITF-GTR-39

Land cover with at least 50 percent in GAP status 1 or 2—

Seventeen land cover units are over 50 percent protected under GAP status 1 or 2

and they account for 5.1 percent of the island. They include important primary and

mature secondary forest types in the Luquillo Mountains, freshwater Pterocarpus

swamps, forests on serpentine substrates, and a number of dryland habitats unique to

Mona Island and the Guánica Biosphere Reserve.

Land cover analysis limitations and discussion—

Assessing the conservation status of natural land cover is limited by several factors:

• GAP has typically found the accuracy of the mapped distributions of

natural communities at the floristic (e.g., alliance) level to be substantially

lower and more variable than that of animal distributions.

• Any aggregation of biotic units (e.g., above species) is a surrogate for

species or lower levels of biotic organization and will underrepresent

conservation needs (Pressey and Logan 1995).

• We do not distinguish the degree of natural condition or value of the

mapped units because of management manipulation, exotic invasion,

or spatial configuration. Considering an aggregation of species such as

we have mapped to be sufficiently represented in existing conservation

areas cannot be determined solely by the percentage of the community

represented because the aggregation has unmapped variation in species

composition that we could not measure. Until individual plant species

distributions can be mapped, it is not possible to assure that the full range

of plant biodiversity is represented, and surrogates must be used.

Predicted animal species distribution analysis—

Table 12 provides the area in hectares of each mapped species by status and the

percentage of the total habitat area of that species. For example, a typical entry

indicates the habitat area of the yellow-shouldered blackbird, Agelaius xanthomus,

is 61,073 ha (6.8 percent of the island) and of that area 14,363 ha is ranked GAP

status 1 or 2, which represents 23.5 percent of the habitat used by that species.

As a coarse indicator of the status of the elements, we provide a breakdown

along five levels of protection (<1 percent, 1 to <10 percent, 10 to <20 percent, 20

to <50 percent, ≥50 percent). The <1-percent level indicates those elements with

essentially none of their distribution in a protected status; levels of 10, 20, and 50

percent have been recommended in the literature as necessary amounts of conservation (McNeely and Miller 1994, Noss and Cooperrider 1994, Odum and Odum
85

The Puerto Rico Gap Analysis Project

1972, Ride 1975). Summaries of the analysis according to the thresholds described

above are shown in table 12. Table 13 lists the locally or federally listed endangered, endemic, and vulnerable species.

Species with <1 percent in GAP status 1 or 2—

Four species have less than 1 percent of their habitat protected. Two species of

gecko are common in urban areas and not in need of protection. One bird, Carduelis

cucullata, is nonnative and has not been seen recently. Eleutherodactylus cooki,

the guajón or rock coqui, has limited habitat and none of it is protected. This species

and its habitat are strongly in need of protection. A fifth species, Eleutherodactylus

juanariveroii, the coqui llanero, is a recently discovered species of coqui that has yet

to be incorporated into the complete PRGAP database. It has a very limited known

range and the area of sighting is unprotected. Plans are underway to protect the area

but they conflict with ongoing development plans, so the outcome and survival of

this species is not certain (fig. 38) (Ríos-López and Thomas 2007).

Species with 1 to <10 percent in GAP status 1 or 2—

Seventy-seven species have 1 to less than 10 percent of their habitat protected.

Many are widespread and occur in disturbed habitats. A few, such as the blind

snake Typhlops platycephalus, have limited habitat (15 percent of the island) and

the majority of this is unprotected (98 percent of its habitat).

Species with l0 to <20 percent in GAP status 1 or 2—

Thirty-two species have 10 to less than 20 percent of their habitat protected. This

includes the broad-winged hawk, Buteo platypterus brunnescens, an important

competitor of the red-tailed hawk, Buteo jamaicensis, both of which prey on the

endangered Puerto Rican parrot, Amazona vittata. These 32 species are an even mix

of those with widespread and limited habitat extent.

Species with 20 to <50 percent in GAP status 1 or 2—

Forty-three species have 20 to less than 50 percent of their habitat protected. A

number of these species are limited in the extent of their habitat. All but one, the

wrinkled frog, Eleutherodactylus wightmanae, with a habitat extent covering 11

percent, are found on less than 10 percent of the island. A number of endangered

species in this group would benefit from greater protection as well as habitat improvement and expansion. Many are limited to less extensive habitats such as saline

and freshwater ponds and wetlands, or high mountain areas.
86

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 12—Predicted distribution area and percentage of the total island for all resident, endemic, and endangered terrestrial vertebrate species in Puerto Rico Common name Common name Protected Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category Ha % Ha % Ha % Ha % Ha % Ha % Puerto Rico 894 912 100 66 275 7.4 11 240 1.3 55 035 6.1 2253 0.3 826 385 92 4 Accipiter striatus Puerto Rican sharp- Halcón de sierra venator shinned hawk 84 589 9.5 20 647 24.4 6752 8.0 13 895 16.4 0 0 63 941 75.6 2 Agelaius Yellow-shouldered Mariquita de xanthomus blackbird Puerto Rico 61 073 6.8 14 363 23.5 1263 2.1 13 100 21.4 578 0.9 46 133 75.5 2 Alsophis portoricensis Puerto Rican racer Culebra corredora 826 049 92.4 60 162 7.3 9875 1.2 50 287 6.1 1581 0.2 764 306 92.5 4 Amazona ventralis Hispaniolan parrot Cotorra de la Española 1570 0.2 203 12.9 9 0.6 194 12.3 134 8.5 1233 78.5 3 Amazona vittata Puerto Rican parrot Cotorra

Puertorriqueña 5648 0.6 5479 97.0 4788 84.8 691 12.2 0 0 169 3.0 1

Ameiva alboguttata Puerto Rican Siguan de la Mona 4576 0.5 4576 100 0 0 4576 100 0 0 0 0 1

ground lizard

Ameiva desechensis Puerto Rican Siguana de Desecheo 121 0 121 100 0 0 121 100 0 0 0 0 1

ground lizard

Ameiva exsul Puerto Rican Siguana común 370 939 41.5 9042 2.4 943 0.3 8099 2.2 891 0.2 361 005 97.3 4

ground lizard

Ameiva wetmorei Blue-tailed ground Siguana de rabo azul 23 550 2.6 5547 23.6 225 1.0 5322 22.6 220 0.9 17 783 75.5 2

lizard

Ammodramus Grasshopper sparrow Gorrión chicharra 110 886 12.4 5527 5.0 292 0.3 5235 4.7 559 0.5 104 799 94.5 4

savannarum

Amphisbaena bakeri Baker’s legless Culebrita ciega de 107 775 12.1 5725 5.3 3 0 5722 5.3 129 0.1 101 920 94.6 4

lizard Baker

Amphisbaena caeca Puerto Rican worm Culebrita ciega 283 610 31.7 29 875 10.5 4207 1.5 25 668 9.1 257 0.1 253 477 89.4 3

lizard común

Amphisbaena schmidti Schmidt’s worm Culebrita ciega de 71 972 8.0 2339 3.3 82 0.1 2257 3.1 45 0.1 69 587 96.7 4

lizard Schmidt

Amphisbaena xera North American Culebrita ciega 80 480 9.0 6441 8.0 226 0.3 6215 7.7 275 0.3 73 763 91.7 4

worm lizard del seco

Anas bahamensis White-cheeked Pato quijada colorada 4557 0.5 2770 60.8 563 12.3 2207 48.4 64 1.4 1722 37.8 1

pintail

Anolis cooki Cook’s anole Lagartijo del seco 25 291 2.8 5546 21.9 1031 4.1 4514 17.8 963 3.8 18 783 74.3 2

Anolis cristatellus Crested anole Lagartijo común 876 825 98.0 55 661 6.3 10 535 1.2 45 127 5.1 1668 0.2 819 495 93.5 4

Anolis cuvieri Puerto Rican Lagarto verde 270 691 30.3 29 342 10.8 7645 2.8 21 697 8.0 178 0.1 241 171 89.1 3

giant anole

Anolis desechensis Anole Lagartijo de Desecheo 121 0 121 100 0 0 121 100 0 0 0 0 1Anolis evermanni Emerald anole Lagartijo verde 286 364 32.0 29 070 10.2 7630 2.7 21 440 7.5 130 0 257 164 89.8 3

Anolis gundlachi Yellow-bearded Lagartijo barba 266 628 29.8 28 556 10.7 7167 2.7 21 389 8.0 104 0 237 968 89.3 3

anole amarilla
87

The Puerto Rico Gap Analysis Project

Table 12—Predicted distribution area and percentage of the total island for all resident, endemic, and endangered terrestrial vertebrate species inPuerto Rico (continued)

Common name Common name Protected

Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category

Ha % Ha % Ha % Ha % Ha % Ha %

Anolis krugi Upland grass anole Lagartijo jardinero 296 095 33.1 7358 2.5 846 0.3 6512 2.2 85 0 288 652 97.5 4

de montana

Anolis monensis Mona anole Lagartijo de la Mona 5571 0.6 5571 100 14 0.3 5557 99.7 0 0 0 0 1

Anolis occultus Puerto Rican Lagartijo pigmeo 297 730 33.3 30 376 10.2 7769 2.6 22 607 7.6 106 0 267 248 89.8 3

pygmy anole

Anolis poncensis Dryland grass Lagartijo jardinero 106 899 12.0 9040 8.5 396 0.4 8644 8.1 730 0.7 97 129 90.9 4

anole de Ponce

Anolis pulchellus Common grass Lagartijo jardinero 422 871 47.3 11 695 2.8 1150 0.3 10 545 2.5 994 0.2 410 182 97.0 4

anole

Anolis roosevelti Culebra Island Chipojo de Culebra 14 893 1.7 7464 50.1 492 3.3 6972 46.8 18 0.1 7411 49.8 1

giant anole

Anolis stratulus Barred anole Lagartijo manchado 403 298 45.1 40 885 10.1 8269 2.1 32 617 8.1 259 0.1 362 155 89.8 3

Anthracothorax Antillean mango Zumbador dorado 508 304 56.8 17 066 3.4 1586 0.3 15 480 3.0 808 0.2 490 430 96.5 4

dominicus

Anthracothorax viridis Green mango Zumbador verde 419 555 46.9 32 052 7.6 7868 1.9 24 184 5.8 270 0.1 387 233 92.3 4

Aratinga canicularis Orange-fronted Periquito 18 690 2.1 449 2.4 21 0.1 428 2.3 66 0.4 18 175 97.2 4

parakeet frentianaranjado

Aratinga chloroptera Hispaniolan parakeet Periquito de la Española 5437 0.6 5437 100 0 0 5437 100 0 0 0 0 1Aratinga pertinax Brown-throated Periquito de curazao 4733 0.5 881 18.6 275 5.8 606 12.8 1 0 3851 81.4 3

parakeet

Ardea alba Great egret Garzón blanco 84 803 9.5 8050 9.5 1519 1.8 6530 7.7 553 0.7 76 200 89.9 4

Arrhyton exiguum Culebra garden Culebra de jardín 510 750 57.1 49 657 9.7 9795 1.9 39 863 7.8 722 0.1 460 370 90.1 4

snake

Artibeus jamaicensis Jamaican fruit- Muciérlago frutero 636 621 71.2 55 972 8.8 9832 1.5 46 140 7.2 638 0.1 580 011 91.1 4

eating bat

Asio flammeus Short-eared owl Múcaro real 177 239 19.8 21 374 12.1 2771 1.6 18 603 10.5 510 0.3 155 356 87.7 3

Brachyphylla Antillean fruit- Murciélago 376 550 42.1 38 110 10.1 8547 2.3 29 563 7.9 498 0.1 337 942 89.7 3

cavernarum eating bat cavernícola

Bubulcus ibis Cattle egret Garza del ganado 339 102 37.9 13 935 4.1 2197 0.6 11 738 3.5 1187 0.3 323 980 95.5 4

Bufo lemur Puerto Rican Sapo concho 241 0 75 31.2 0 0 75 31.1 0 0 166 68.8 2

crested toad

Bufo marinus Giant toad Sapo común 525 164 58.7 17 367 3.3 2338 0.4 15 028 2.9 1194 0.2 506 604 96.5 4

Buteo jamaicensis Red-tailed hawk Guaraguao colirrojo 874 087 97.7 62 670 7.2 10 367 1.2 52 304 6.0 1363 0.2 810 054 92.7 4

Buteo platypterus Puerto Rican broad- Guaraguao de bosque 188 451 21.1 25 871 13.7 7071 3.8 18 800 10 95 0.1 162 485 86.2 3

brunnescens winged hawk

Butorides virescens Green heron Martinete verde 41 920 4.7 8646 20.6 1686 4.0 6960 16.6 649 1.5 32 625 77.8 2
88

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 12—Predicted distribution area and percentage of the total island for all resident, endemic, and endangered terrestrial vertebrate species in Puerto Rico (continued) Common name Common name Protected Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category Ha % Ha % Ha % Ha % Ha % Ha % Caprimulgus Puerto Rican Guabairo de Puerto 15 411 1.7 3788 24.6 56 0.4 3733 24.2 24 0.2 11 599 75.3 2 noctitherus nightjar Rico Carduelis cucullata Red siskin Cardenalito 0 0 0 0 0 0 0 0 0 0 0 0 5 Cathartes aura Turkey vulture Aura tiñosa 282 785 31.6 18 440 6.5 1721 0.6 16 719 5.9 1382 0.5 262 964 93.0 4 Catoptrophorus Willet Playero aliblanco 3924 0.4 1468 37.4 292 7.4 1177 30 315 8.0 2140 54.5 2 semipalmatus Charadrius vociferus Killdeer Chorlo sabanero 100 307 11.2 4303 4.3 748 0.7 3556 3.5 901 0.9 95 102 94.8 4 Charadrius wilsonia Wilson’s plover Chorlo marítimo 12 303 1.4 5686 46.2 1276 10.4 4409 35.8 355 2.9 6262 50.9 2

Chlorostilbon Puerto Rican Zumbadorcito de 327 825 36.7 34 673 10.6 8570 2.6 26 103 8.0 133 0 293 019 89.4 3

maugaeus emerald Puerto Rico

Chordeiles gundlachii Antillean nighthawk Querequeqúe 172 738 19.3 16 805 9.7 664 0.4 16141 9.3 1280 0.7 154 654 89.5 4

Antillano

Coccyzus americanus Yellow-billed cuckoo Pájaro bobo 29 348 3.3 11 758 40.1 918 3.1 10841 36.9 181 0.6 17 410 59.3 2

pechiblanco

Coccyzus minor Mangrove cuckoo Pájaro bobo menor 247 501 27.7 28 646 11.6 1652 0.7 26 994 10.9 545 0.2 218 310 88.2 3Coereba flaveola Bananaquit Reinita común 534 523 59.8 48 711 9.1 9357 1.8 39 353 7.4 605 0.1 485 208 90.8 4

Columbina passerina Common ground- Rolita 581 754 65.1 22 035 3.8 2213 0.4 19 822 3.4 1373 0.2 558 346 96.0 4

dove

Contopus latirostris Lesser Antillean Bobito Antíllano 169 101 18.9 13 598 8.0 384 0.2 13 214 7.8 106 0.1 155 396 91.9 4

pewee Menor

Crotophaga ani Smooth-billed ani Garrapatero 308 413 34.5 16 138 5.2 1553 0.5 14 586 4.7 998 0.3 291 277 94.4 4

Cyclura stejnegeri Mona Island rock Iguana de la Mona 5483 0.6 5483 100 0 0 5483 100 0 0 0 0 1

iguana

Cypseloides niger Black swift Vencejo negro 64 818 7.2 16 944 26.1 7151 11.0 9793 15.1 0 0 47 874 73.9 2

Dendrocygna arborea West Indian Chiriría Caribeña 11 184 1.3 5421 48.5 1037 9.3 4384 39.2 97 0.9 5666 50.7 2

whistling-duck

Dendrocygna bicolor Fulvous whistling- Chiriría bicolor 29 607 3.3 11 032 37.3 595 2.0 10 436 35.2 393 1.3 18 182 61.4 2

duck

Dendroica adelaidae Adelaide’s warbler Reinita mariposera 35 604 4.0 3335 9.4 162 0.5 3173 8.9 206 0.6 32 063 90.1 4

Dendroica angelae Elfin-woods warbler Reinita de bosque enaño 14 223 1.6 7177 50.5 2470 17.4 4707 33.1 0 0 7046 49.5 1

Dendroica petechia Yellow warbler Reinita amarilla 12 296 1.4 5471 44.5 1096 8.9 4376 35.6 41 0.3 6783 55.2 2

Diploglossus pleei Puerto Rican Celestus 375 606 42.0 19 769 5.3 4591 1.2 15 178 4.0 575 0.2 355 263 94.6 4

galliwasp

Egretta caerulea Little blue heron Garza azul 34 002 3.8 8654 25.5 1645 4.8 7009 20.6 409 1.2 24 939 73.3 2Egretta thula Snowy egret Garza blanca 32 069 3.6 8150 25.4 1645 5.1 6504 20.3 668 2.1 23 252 72.5 2
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The Puerto Rico Gap Analysis Project

Table 12—Predicted distribution area and percentage of the total island for all resident, endemic, and endangered terrestrial vertebrate species inPuerto Rico (continued)

Common name Common name Protected

Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category

Ha % Ha % Ha % Ha % Ha % Ha %

Egretta tricolor Tricolored heron Garza pechiblanca 8237 0.9 3982 48.3 713 8.7 3269 39.7 0 0 4255 51.7 2

Elaenia martinica Caribbean elaenia Juí blanco 29 912 3.3 11 047 36.9 247 0.8 10 800 36.1 106 0.4 18 758 62.7 2

Eleutherodactylus Antillean frog Churí 593 293 66.3 16 943 2.9 2102 0.4 14 840 2.5 696 0.1 575 654 97.0 4

antillensis

Eleutherodactylus Grass coqui Coquí de las hierbas 294 705 33.0 6430 2.2 652 0.2 5778 2.0 131 0 288 145 97.8 4

brittoni

Eleutherodactylus Whistling frog Coquí pitito 413 883 46.3 26 150 6.3 1996 0.5 24 154 5.8 369 0.1 387 363 93.6 4

cochranae

Eleutherodactylus Rock coqui Coquí guajón 3864 0.4 0 0 0 0 0 0 0 0 3864 100 5

cooki

Eleutherodactylus Common coqui Coquí de las yerbas 1398 0.2 428 30.6 161 11.5 267 19.1 4 0.3 966 69.1 2

coqui

Eleutherodactylus Eneida’s coqui Coquí de eneida 134 307 15.0 22 936 17.1 7544 5.6 15 392 11.5 0 0 111 371 82.9 3

eneidae

Eleutherodactylus Cricket coqui Coquí grillo 44 987 5.0 3469 7.7 365 0.8 3105 6.9 2 0 41 517 92.3 4

gryllus

Eleutherodactylus Tree-hole frog Coquí de Hedrick 10 764 1.2 8287 77.0 4993 46.4 3293 30.6 0 0 2477 23.0 1

hedrickiEleutherodactylus Golden coqui Coquí dorado 3332 0.4 1445 43.4 2 0 1444 43.3 0 0 1887 56.6 2

jasperi

Eleutherodactylus Warty coqui Coquí martillito 387 0 290 74.9 170 43.9 120 31.0 0 0 97 25.1 1

locustus

Eleutherodactylus Mona coqui Coquí de la Mona 5557 0.6 5557 100 0 0 5557 100 0 0 0 0 1

monensis

Eleutherodactylus Puerto Rican Coquí de la montaña 73 158 8.2 17 912 24.5 5991 8.2 11 921 16.3 0 0 55 245 75.5 2

portoricensis coqui

Eleutherodactylus Ground coqui Coquí caoba 179 182 20 27 435 15.3 7221 4.0 20 214 11.3 11 0 151 737 84.7 3

richmondi

Eleutherodactylus Burrowing coqui Coquí duende 303 0 303 100 255 84.0 48 16.0 0 0 0 0 1

unicolor

Eleutherodactylus Wrinkled frog Coquí melodioso 98 708 11.0 24 311 24.6 7106 7.2 17 205 17.4 1 0 74 396 75.4 2

wightmanae

Epicrates inornatus Puerto Rican boa Culebrón 414 379 46.3 37 178 9.0 8827 2.1 28 351 6.8 286 0.1 376 916 91.0 4

Epicrates monensis Virgin Islands tree Culebrón de la 2795 0.3 469 16.8 280 10 189 6.8 1 0 2325 83.2 3

granti boa Isla Virgin

Epicrates monensis Mona boa Culebrón de la Mona 5436 0.6 5436 100 0 0 5436 100 0 0 0 0 1

monensis

Eptesicus fuscus Big brown bat Murciélago Marrón 481 270 53.8 30 873 6.4 8252 1.7 22 621 4.7 668 0.1 449 730 93.4 4

mayor
90

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 12—Predicted distribution area and percentage of the total island for all resident, endemic, and endangered terrestrial vertebrate species in Puerto Rico (continued) Common name Common name Protected Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category Ha % Ha % Ha % Ha % Ha % Ha % Erophylla sezekorni Buffy flower bat Murciélago las flores 582 629 65.2 39 759 6.8 8611 1.5 31 148 5.3 816 0.1 542 054 93.0 4 Estrilda melpoda Orange-cheeked Veterano 381 550 42.7 8714 2.3 789 0.2 7925 2.1 1028 0.3 371 807 97.4 4 waxbill Estrilda troglodytes Black-rumped Veterano orejicolorado 147 112 16.5 4820 3.3 487 0.3 4333 2.9 734 0.5 141 558 96.2 4 waxbill Eulampis holosericeus Green-throated carib Zumbador de pecho azul 71 073 7.9 6805 9.6 638 0.9 6167 8.7 495 0.7 63 773 89.7 4 Euphonia musica Antillean euphonia Jilguero 152 797 17.1 27 323 17.9 6948 4.5 20 376 13.3 0 0 125 473 82.1 3 Euplectes afer Yellow-crowned Napoleon tejedor 7232 0.8 754 10.4 152 2.1 602 8.3 31 0.4 6447 89.1 3 bishop

Euplectes franciscanus Orange bishop Obispo colorado 98 783 11.0 3479 3.5 312 0.3 3167 3.2 712 0.7 94 591 95.8 4

Falco sparverius American kestrel Halcón común 159 196 17.8 11 658 7.3 690 0.4 10 968 6.9 995 0.6 146 543 92.1 4Fulica caribaea Caribbean coot Gallinazo Caribeño 1130 0.1 233 20.6 0 0 233 20.6 265 23.4 632 55.9 2Gallinula chloropus Common moorhen Gallareta común 6821 0.8 2599 38.1 591 8.7 2008 29.4 278 4.1 3944 57.8 2

Geotrygon chrysia Key West quail-dove Paloma perdíz aurea 214 244 24.0 20 868 9.7 393 0.2 20 475 9.6 117 0.1 193 259 90.2 4Geotrygon montana Ruddy quail-dove Paloma perdíz rojiza 133 503 14.9 25 348 19.0 6736 5.0 18 612 13.9 0 0 108 155 81.0 3Geotrygon mystacea Bridled quail-dove Paloma perdíz de 41 483 4.6 9293 22.4 311 0.8 8981 21.7 3 0 32 188 77.6 2

Martinica

Hemidactylus brookii Brook’s house gecko Salamanquesa 36 345 4.1 148 0.4 10 0 138 0.4 44 0.1 36 154 99.5 5

Hemidactylus mabouia Cosmopolitan house Salamanquesa 45 264 5.1 105 0.2 4 0 101 0.2 7 0 45 151 99.8 5

gecko

Herpestes javanicus Small Indian Mangosta 775 094 86.7 55 684 7.2 9645 1.2 46 039 5.9 1302 0.2 718 108 92.6 4

mongoose

Himantopus mexicanus Black-necked stilt Viuda Mexicana 33 746 3.8 8380 24.8 1596 4.7 6784 20.1 365 1.1 25 001 74.1 2

Hirundo rustica Barn swallow Golondrina horquilla 51 415 5.7 4274 8.3 429 0.8 3845 7.5 323 0.6 46 818 91.1 4

Icterus dominicensis Greater Antillean Calandria 391 912 43.8 35 398 9.0 8006 2.0 27 392 7.0 543 0.1 355 970 90.8 4

oriole

Ixobrychus exilis Least bittern Martinetito 2830 0.3 871 30.8 113 4.0 758 26.8 50 1.8 1909 67.5 2Lasiurus borealis Eastern red bat Murciélago rojo 74 097 8.3 14 842 20.0 6763 9.1 8079 10.9 207 0.3 59 048 79.7 2Leptodactylus White-lipped frog Ranita de blanco 492 760 55.1 32 821 6.7 8004 1.6 24 817 5.0 499 0.1 459 440 93.2 4

albilabris

Lonchura cucullata Bronze mannikin Diablito 196 080 21.9 5694 2.9 307 0.2 5387 2.7 682 0.3 189 704 96.7 4

Lonchura malabarica Warbling silverbill Gorrión picoplata 56 303 6.3 3859 6.9 156 0.3 3703 6.6 669 1.2 51 776 92.0 4

Lonchura malacca Chestnut mannikin Monjita tricolor 389 502 43.6 8582 2.2 710 0.2 7871 2.0 1054 0.3 379 866 97.5 4

Lonchura punctulata Nutmeg mannikin Gorrión canela 146 489 16.4 3830 2.6 453 0.3 3377 2.3 686 0.5 141 972 96.9 4

Loxigilla portoricensis Puerto Rican Gallito 210 806 23.6 29 121 13.8 7298 3.5 21 823 10.4 5 0 181 680 86.2 3

bullfinch
91

The Puerto Rico Gap Analysis Project

Table 12—Predicted distribution area and percentage of the total island for all resident, endemic, and endangered terrestrial vertebrate species inPuerto Rico (continued)

Common name Common name Protected

Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category

Ha % Ha % Ha % Ha % Ha % Ha %

Mabuya mabouya Slippery-backed Lucía 106 920 12.0 19 860 18.6 776 0.7 19 084 17.8 315 0.3 86 744 81.1 3

sloanei mabuya

Margarops fuscatus Pearly-eyed thrasher Zorzal pardo 638 729 71.4 51 274 8.0 9696 1.5 41 578 6.5 873 0.1 586 582 91.8 4

Megascops nudipes Puerto Rican Múcaro común 455 750 51.0 41 999 9.2 9504 2.1 32 495 7.1 549 0.1 413 202 90.7 4

screech-owl

Melanerpes Puerto Rican Carpintero 463 506 51.8 47 677 10.3 9080 2.0 38 597 8.3 552 0.1 415 278 89.6 3

portoricensis woodpecker

Mimus polyglottos Northern Ruisenor 546 630 61.1 9657 1.8 1352 0.2 8305 1.5 989 0.2 535 985 98.1 4

mockingbird

Molossus molossus Velvety free-tailed Murciélago casero 707 108 79.1 56 607 8.0 9966 1.4 46 640 6.6 915 0.1 649 586 91.9 4

bat

Molothrus bonariensis Shiny cowbird Tordo lustrosso 395 788 44.3 9120 2.3 1131 0.3 7989 2.0 835 0.2 385 833 97.5 4

Monophyllus redmani Puerto Rican long- Murciélago 696 229 77.9 49 194 7.1 9259 1.3 39 935 5.7 1563 0.2 645 473 92.7 4

portoricensis tongued bat

Mormoops blainvillii Antillean ghost- Murciélago barbicacho 350 556 39.2 23 391 6.7 619 0.2 22 772 6.5 298 0.1 326 867 93.2 4

faced bat

Myiarchus antillarum Puerto Rican Juí de Puerto Rico 409 729 45.8 44 959 11.0 9294 2.3 35 665 8.7 494 0.1 364 276 88.9 3

flycatcher

Nesospingus Puerto Rican tanager Llorosa de Puerto Rico 161 567 18.1 24 184 15.0 7106 4.4 17 078 10.6 2 0 137 381 85.0 3

speculiferus

Noctilio leporinus Greater bulldog bat Murciélago pescador 12 125 1.4 3177 26.2 659 5.4 2518 20.8 629 5.2 8319 68.6 2

Nyctanassa violacea Yellow-crowned Yaboa comun 23 119 2.6 7631 33.0 1764 7.6 5867 25.4 914 4.0 14 573 63.0 2

night-heron

Nycticorax nycticorax Black-crowned Yaboa real 14 627 1.6 4857 33.2 1091 7.5 3766 25.7 310 2.1 9461 64.7 2

night-heron

Orthorhyncus Antillean crested Zumbador crestado 94 472 10.6 16 441 17.4 1474 1.6 14 968 15.8 182 0.2 77 849 82.4 3

cristatus hummingbird

Oxyura jamaicensis Ruddy duck Pato chorizo 6841 0.8 3255 47.6 828 12.1 2427 35.5 367 5.4 3219 47.1 2

Patagioenas inornata Puerto Rican plain Paloma sabanera 306 755 34.3 4093 1.3 1170 0.4 2923 1.0 278 0.1 302 384 98.6 4

pigeon

Patagioenas White-crowned Paloma cabeciblanca 14 489 1.6 6649 45.9 1305 9.0 5344 36.9 157 1.1 7683 53.0 2

leucocephala pigeon

Patagioenas squamosa Scaly-naped pigeon Paloma turca 408 863 45.7 43 938 10.7 8594 2.1 35 343 8.6 256 0.1 364 670 89.2 3

Pelecanus occidentalis Brown pelican Pelícano pardo 32 200 3.6 9502 29.5 2160 6.7 7342 22.8 671 2.1 22 027 68.4 2Petrochelidon fulva Cave swallow Golondrina de cuevas 553 717 61.9 23 443 4.2 2660 0.5 20 783 3.8 1932 0.3 528 343 95.4 4

Phyllodactylus Greater Antillean Salamanquesa bandeada 11 663 1.3 3968 34.0 295 2.5 3673 31.5 0 0 7695 66.0 2

wirshingi leaf-toed gecko
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GENERAL TECHNICAL REPORT IITF-GTR-39 Table 12—Predicted distribution area and percentage of the total island for all resident, endemic, and endangered terrestrial vertebrate species in Puerto Rico (continued) Common name Common name Protected Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category Ha % Ha % Ha % Ha % Ha % Ha % Podilymbus podiceps Pied-billed grebe Zaramago 2696 0.3 1258 46.7 131 4.9 1127 41.8 0 0 1437 53.3 2 Porphyrio martinica Purple gallinule Gallareta azul 161 0 137 85.4 0 0 137 85.4 0 0 23 14.6 1 Porzana flaviventer Yellow-breasted Gallito amarillo 827 0.1 471 56.9 95 11.5 376 45.4 46 5.6 310 37.5 1 crake Progne dominicensis Caribbean martin Golondrina de iglesias 109 909 12.3 2736 2.5 277 0.3 2458 2.2 86 0.1 107 087 97.4 4 Pteronotus parnellii Parnell’s mustached Murciélago bigotudo 451 275 50.5 32 586 7.2 9077 2.0 23 510 5.2 2100 0.5 416 589 92.3 4 portoricensis bat mayor Pteronotus quadridens Sooty mustached Murciélago bigotudo 291 484 32.6 29 257 10 7799 2.7 21 458 7.4 268 0.1 261 958 89.9 3 bat menor

Quiscalus niger Greater Antillean Mozambique 443 537 49.6 19 175 4.3 2418 0.5 16 757 3.8 1994 0.4 422 368 95.2 4

grackle

Rallus longirostris Clapper rail Pollo de mangle 11 117 1.2 5068 45.6 1313 11.8 3754 33.8 133 1.2 5916 53.2 2

Saurothera vieilloti Puerto Rican Pájaro bobo mayor 349 779 39.1 31 973 9.1 7334 2.1 24 638 7.0 115 0 317 691 90.8 4

lizard-cuckoo

Sphaerodactylus Gaige’s dwarf Salamanquita de 59 087 6.6 10 385 17.6 4489 7.6 5897 10 2 0 48 700 82.4 3

gaigeae gecko Pandura

Sphaerodactylus Klauber’s dwarf Salamanquita negra 248 909 27.8 28 977 11.6 7292 2.9 21 685 8.7 104 0 219 828 88.3 3

klauberi gecko

Sphaerodactylus Desecheo gecko Salamanquita de 15 0 15 100 0 0 15 100 0 0 0 0 1

levinsi Desecheo

Sphaerodactylus Common dwarf Salamanquita común 503 967 56.4 49 185 9.8 9609 1.9 39 576 7.9 585 0.1 454 197 90.1 4

macrolepis gekco

Sphaerodactylus Monito gecko Salamanquita de Monito 14 0 14 100 14 100 0 0 0 0 0 0 1

micropithecus

Sphaerodactylus Mona dwarf Salamanquita de la 5531 0.6 5531 100 0 0 5531 100 0 0 0 0 1

monensis gekco Mona

Sphaerodactylus Nichol’s dwarf Salamanquita pigmea 7566 0.8 2059 27.2 80 1.1 1979 26.2 31 0.4 5476 72.4 2

nicholsi gecko

Sphaerodactylus Roosevelt’s dwarf Salamanquita de 22 353 2.5 5584 25.0 187 0.8 5397 24.1 223 1.0 16 545 74.0 2

roosevelti gecko Roosevelt

Sphaerodactylus Townsend’s dwarf Salamanquita del 20 898 2.3 5787 27.7 68 0.3 5719 27.4 2 0 15 108 72.3 2

townsendi gecko sureste

Spindalis Puerto Rican Reina mora 433 764 48.5 37 416 8.6 8781 2.0 28 635 6.6 275 0.1 396 072 91.3 4

portoricensis spindalis

Stenoderma rufum Desmarest’s fig- Murciélago rojo 444 969 49.8 42 943 9.7 8132 1.8 34 811 7.8 379 0.1 401 648 90.3 4

eating bat frutero

Tachybaptus dominicus Least grebe Tigua 341 0 46 13.5 0 0 46 13.5 45 13.1 250 73.4 3
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Table 12—Predicted distribution area and percentage of the total island for all resident, endemic, and endangered terrestrial vertebrate species inPuerto Rico (continued)

Common name Common name Protected

Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category

Ha % Ha % Ha % Ha % Ha % Ha %

Tadarida brasiliensis Brazilian free-tailed Murciélago de 410 380 45.9 27 999 6.8 8331 2.0 19 668 4.8 1568 0.4 380 813 92.8 4

bat cola libre

Tiaris bicolor Black-faced Gorrión negro 311 963 34.9 12 478 4.0 1483 0.5 10 995 3.5 1253 0.4 298 232 95.6 4

grassquit

Tiaris olivacea Yellow-faced Gorrion barba 433 023 48.4 17 034 3.9 2184 0.5 14 850 3.4 1334 0.3 414 654 95.8 4

grassquit amarilla

Todus mexicanus Puerto Rican tody San Pedrito 379 296 42.4 37 504 9.9 8638 2.3 28 866 7.6 158 0 341 634 90.1 4

Turdus plumbeus Red-legged thrush Zorzal de patas

coloradas 380 984 42.6 38 371 10.1 8282 2.2 30 090 7.9 251 0.1 342 362 89.9 3

Typhlops granti Grant’s blind snake Vibora de Grant 11 831 1.3 3891 32.9 206 1.7 3685 31.2 0 0 7939 67.1 2

Typhlops hypomethes Blind snake Vibora universitaria 5721 0.6 2747 48.0 204 3.6 2543 44.4 0 0 2974 52.0 2

Typhlops monensis Mona blind snake Vibora de la Mona 106 0 106 100 0 0 106 100 0 0 0 0 1Typhlops platycephalus Blind snake Vibora de cabeza 9096 1.0 186 2.0 9 0.1 177 1.9 2 0 8909 97.9 4

aplastada

Typhlops richardii Richard’s blind Vibora común 2925 0.3 650 22.2 518 17.7 133 4.5 3 0.1 2272 77.7 2

snake

Typhlops rostellatus Puerto Rican wet- Vibora de pico 247 592 27.7 20 629 8.3 7314 3.0 13 315 5.4 364 0.1 226 599 91.5 4

land blind snake

Tyrannus Loggerhead Clerigo 343 451 38.4 14 218 4.1 1103 0.3 13 116 3.8 292 0.1 328 942 95.8 4

caudifasciatus kingbird

Tyrannus dominicensis Gray kingbird Pitirre gris 725 432 81.1 27 482 3.8 3202 0.4 24 280 3.3 1515 0.2 696 435 96.0 4

Vireo altiloquus Black-whiskered Julian chivi 541 744 60.6 40 513 7.5 9037 1.7 31 476 5.8 795 0.1 500 437 92.4 4

vireo

Vireo latimeri Puerto Rican vireo Bienteveo de 151 020 16.9 15 704 10.4 588 0.4 15 116 10 106 0.1 135 210 89.5 3

Puerto Rico

Zenaida asiatica White-winged dove Tórtola aliblanca 487 655 54.5 44 150 9.1 6408 1.3 37 742 7.7 575 0.1 442 931 90.8 4

Zenaida aurita Zenaida dove Tórtola cardosantera 572 274 64.0 32 579 5.7 2798 0.5 29 781 5.2 1411 0.2 538 284 94.1 4Zenaida macroura Mourning dove Tórtola rabilarga 267 912 30 25 008 9.3 1477 0.6 23 530 8.8 1208 0.5 241 696 90.2 4

Note: Protection category 1 = ≥50 percent protected, 2 = 20 to < 50 percent protected, 3 = 10 to < 20 percent protected, 4 = 1 to < 10 percent protected, 5 = < 1 percent protected.
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GENERAL TECHNICAL REPORT IITF-GTR-39 Table 13—Federally listed endangered (LE), threatened (LT) and partial status (PS) species and those considered critically endangered (CE), endangered (EN), vulnerable (VU), or data deficient (DD) by the Puerto Rico Department of Natural and Environmental Resources indicating their area protected Common name Common name Protected Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category Ha % Ha % Ha % Ha % Ha % Ha % Puerto Rico 894 912 100 66 275 7.4 11 240 1.3 55 035 6.1 2253 0.3 826 385 92 4 Accipiter striatus Puerto Rican sharp- Halcón de sierra 84 589 9.5 20 647 24.4 6752 8.0 13 895 16.4 0 0.0 63 941 75.6 2 venator (LE, CR) shinned hawk Agelaius xanthomus Yellow-shouldered Mariquita de Puerto 61 073 6.8 14 363 23.5 1263 2.1 13 100 21.4 578 0.9 46 133 75.5 2 (LE, EN) blackbird Rico Amazona vittata Puerto Rican parrot Cotorra Puertorriqueña 5648 0.6 5479 97.0 4788 84.8 691 12.2 0 0.0 169 3.0 1 (LE, CR) Ammodramus Grasshopper sparrow Gorrión chicharra 110 886 12.4 5527 5.0 292 0.3 5235 4.7 559 0.5 104 799 94.5 4 savannarum

(PS, DD)

Anas bahamensis White-cheeked pintail Pato quijada colorada 4557 0.5 2770 60.8 563 12.3 2207 48.4 64 1.4 1722 37.8 1

(VU)

Anolis cooki (EN) Cook’s anole Lagartijo del Seco 25 291 2.8 5546 21.9 1031 4.1 4514 17.8 963 3.8 18 783 74.3 2

Anolis poncensis (VU) Dryland grass anole Lagartijo jardinero 106 899 12.0 9040 8.5 396 0.4 8644 8.1 730 0.7 97 129 90.9 4

de Ponce

Anolis roosevelti Culebra Island Chipojo de Culebra 14 893 1.7 7464 50.1 492 3.3 6972 46.8 18 0.1 7411 49.8 1

(LE, CR) giant anole

Brachyphylla Antillean fruit- Murciélago cavernícola 376 550 42.1 38 110 10.1 8547 2.3 29 563 7.9 498 0.1 337 942 89.7 3

cavernarum (VU) eating bat

Bufo lemur (LT, EN) Puerto Rican Sapo concho 241 0.0 75 31.2 0 0.0 75 31.1 0 0.0 166 68.8 2

crested toad

Buteo platypterus Puerto Rican Guaraguao de bosque 188 451 21.1 25 871 13.7 7071 3.8 18 800 10.0 95 0.1 162 485 86.2 3

brunnescens broad-winged(LE, CR) hawk

Caprimulgus Puerto Rican Guabairo de Puerto 15 411 1.7 3788 24.6 56 0.4 3733 24.2 24 0.2 11 599 75.3 2

noctitherus nightjar Rico(LE, EN)

Carduelis cucullata Red siskin Cardenalito 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 5

(DD)

Coccyzus americanus Yellow-billed Pájaro bobo 29 348 3.3 11 758 40.1 918 3.1 10 841 36.9 181 0.6 17 410 59.3 2

(PS) cuckoo pechiblanco

Cyclura stejnegeri Mona Island Iguana de la Mona 5483 0.6 5483 100.0 0 0.0 5483 100.0 0 0.0 0 0.0 1

(LT, EN) Rock Iguana

Dendrocygna arborea West Indian Chiriría Caribeña 11 184 1.3 5421 48.5 1037 9.3 4384 39.2 97 0.9 5666 50.7 2

(LE, CR) whistling duck

Dendroica angelae Elfin-woods warbler Reinita de bosque enaño 14 223 1.6 7177 50.5 2470 17.4 4707 33.1 0 0.0 7046 49.5 1

(VU)

Eleutherodactylus cooki Rock coqui Coquí guajón 3864 0.4 0 0.0 0 0.0 0 0.0 0 0.0 3864 100.0 5

(LT, VU)
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Table 13—Federally listed endangered (LE), threatened (LT) and partial status (PS) species and those considered critically endangered (CE), endangered(EN), vulnerable (VU), or data deficient (DD) by the Puerto Rico Department of Natural and Environmental Resources indicating their area protected(continued)

Common name Common name Protected

Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category

Ha % Ha % Ha % Ha % Ha % Ha %

Eleutherodactylus Eneida’s coqui Coquí de Eneida 134 307 15.0 22 936 17.1 7544 5.6 15 392 11.5 0 0.0 111 371 82.9 3

eneidae (CR)

Eleutherodactylus Tree-hole frog Coquí de Hedrick 10 764 1.2 8287 77.0 4993 46.4 3293 30.6 0 0.0 2477 23.0 1

hedricki (DD)

Eleutherodactylus Golden coqui Coquí dorado 3332 0.4 1445 43.4 2 0.0 1444 43.3 0 0.0 1887 56.6 2

jasperi (LT, CR)

Eleutherodactylus Warty coqui Coquí martillito 387 0.0 290 74.9 170 43.9 120 31.0 0 0.0 97 25.1 1

locustus (VU)

Eleutherodactylus Puerto Rican coqui Coquí de la 73 158 8.2 17 912 24.5 5991 8.2 11 921 16.3 0 0.0 55 245 75.5 2

portoricensis (VU) montaña

Eleutherodactylus Ground coqui Coquí caoba 179 182 20.0 27 435 15.3 7221 4.0 20 214 11.3 11 0.0 151 737 84.7 3

richmondi (VU)

Epicrates inornatus Puerto Rican boa Culebrón 414 379 46.3 37 178 9.0 8827 2.1 28 351 6.8 286 0.1 376 916 91.0 4

(LE, VU)

Epicrates monensis Virgin Islands Culebrón de la 2795 0.3 469 16.8 280 10.0 189 6.8 1 0.0 2325 83.2 3

granti (LE, CR) tree boa Isla Virgin

Epicrates monensis Mona boa Culebrón de la 5436 0.6 5436 100.0 0 0.0 5436 100.0 0 0.0 0 0.0 1

monensis (LE, VU) Mona

Erophylla sezekorni Buffy flower bat Murciélago las 582 629 65.2 39 759 6.8 8611 1.5 31 148 5.3 816 0.1 542 054 93.0 4

(VU) flores

Fulica caribaea Caribbean coot Gallinazo Caribeño 1130 0.1 233 20.6 0 0.0 233 20.6 265 23.4 632 55.9 2

(VU)

Gallinula chloropus Common moorhen Gallareta común 6821 0.8 2599 38.1 591 8.7 2008 29.4 278 4.1 3944 57.8 2

(PS)

Geotrygon chrysia Key West quail-dove Paloma perdíz aurea 214 244 24.0 20 868 9.7 393 0.2 20 475 9.6 117 0.1 193 259 90.2 4

(DD)

Geotrygon mystacea Bridled quail-dove Paloma perdíz de 41 483 4.6 9293 22.4 311 0.8 8981 21.7 3 0.0 32 188 77.6 2

(DD) Martinica

Himantopus Black-necked stilt Viuda Mexicana 33 746 3.8 8380 24.8 1596 4.7 6784 20.1 365 1.1 25 001 74.1 2

mexicanus (PS)

Icterus dominicensis Greater Antillean Calandria 391 912 43.8 35 398 9.0 8006 2.0 27 392 7.0 543 0.1 355 970 90.8 4

(DD) oriole

Mabuya mabouya Slippery-backed Lucía 106 920 12.0 19 860 18.6 776 0.7 19 084 17.8 315 0.3 86 744 81.1 3

sloanei (VU) mabuya

Monophyllus Puerto Rican long- Murciélago 696 229 77.9 49 194 7.1 9259 1.3 39 935 5.7 1563 0.2 645 473 92.7 4

redmani tongued batportoricensis (VU)
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GENERAL TECHNICAL REPORT IITF-GTR-39 Table 13—Federally listed endangered (LE), threatened (LT) and partial status (PS) species and those considered critically endangered (CE), endangered (EN), vulnerable (VU), or data deficient (DD) by the Puerto Rico Department of Natural and Environmental Resources indicating their area protected (continued) Common name Common name Protected Species (English) (Spanish) Total habitat Status 1 or 2 Status 1 Status 2 Status 3 Status 4 category Ha % Ha % Ha % Ha % Ha % Ha % Oxyura jamaicensis Ruddy duck Pato chorizo 6841 0.8 3255 47.6 828 12.1 2427 35.5 367 5.4 3219 47.1 2 (VU) Patagioenas inornata Puerto Rican Paloma sabanera 306 755 34.3 4093 1.3 1170 0.4 2923 1.0 278 0.1 302 384 98.6 4 (LE, EN) plain pigeon Pelecanus occidentalis Brown pelican Pelícano pardo 32 200 3.6 9502 29.5 2160 6.7 7342 22.8 671 2.1 22 027 68.4 2 (LE, EN) Porzana flaviventer Yellow-breasted Gallito amarillo 827 0.1 471 56.9 95 11.5 376 45.4 46 5.6 310 37.5 2 (DD) crake

Rallus longirostris Clapper rail Pollo de mangle 11 117 1.2 5068 45.6 1313 11.8 3754 33.8 133 1.2 5916 53.2 2

(PS)

Sphaerodactylus Gaige’s dwarf gecko Salamanquita 59 087 6.6 10 385 17.6 4489 7.6 5897 10.0 2 0.0 48 700 82.4 3

gaigeae (DD) de Pandura

Sphaerodactylus Desecheo gecko Salamanquita de 15 0.0 15 100.0 0 0.0 15 100.0 0 0.0 0 0.0 1

levinsi (DD) Desecheo

Sphaerodactylus Monito gecko Salamanquita de 14 0.0 14 100.0 14 100.0 0 0.0 0 0.0 0 0.0 1

micropithecus Monito(LE, CR)

Tachybaptus Least grebe Tigua 341 0.0 46 13.5 0 0.0 46 13.5 45 13.1 250 73.4 3

dominicus (DD)

Typhlops granti Grant’s blind snake Vibora de Grant 11 831 1.3 3891 32.9 206 1.7 3685 31.2 0 0.0 7939 67.1 2

(DD)

Typhlops monensis Mona blind snake Vibora de la Mona 106 0.0 106 100.0 0 0.0 106 100.0 0 0.0 0 0.0 1

(DD)

Note: Protection category 1 = ≥50 percent protected, 2 = 20 to < 50 percent protected, 3 = 10 to < 20 percent protected, 4 = 1 to < 10 percent protected, 5 = < 1 percent protected.
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The Puerto Rico Gap Analysis Project Figure 38—Eleutherodactylus portoricensis predicted distribution map (based on Río-López and Thomas 2007 and personal communication).
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GENERAL TECHNICAL REPORT IITF-GTR-39

Species with at least 50 percent in GAP status 1 or 2—

Twenty-one species have at least 50 percent of their predicted habitat protected.

These include a number of species found only on forest reserves or particular

protected satellite islands (Mona and Desecheo). All of these species have very

limited habitat and none exceed 2 percent of the island. Many of these species could

benefit from management to extend their range to other suitable habitat, as is being

done with the Puerto Rican parrot, Amazona vittata. Improving habitat inside and

outside the current reserves, and increasing the number and extent of reserves would

also benefit this group.

Endangered species representation in GAP status 1 and 2—

Forty-seven species are listed as either federally threatened or endangered or given

partial status, or are locally listed by the DNER as vulnerable, endangered, critically

endangered, or data deficient (table 13). The extent of habitat for 70 percent of

these species is typically less than 5 percent of the island. Eighty-three percent of

the species have a habitat extent less than 20 percent of the island. Eleutherodactylus

cooki, the guajón or rock coqui, is the least protected, with no protected habitat.

Ten species have less than 10 percent of their habitat protected and 18 species have

less than 20 percent of their habitat protected. Five species are found only in

reserves with 100 percent of their habitat protected.

Analysis Limitations

When applying the results of our analyses, it is critical that the following limitations are considered: (1) the limitations described for each of the component parts

(land cover mapping, animal species mapping, stewardship mapping) of the analyses, (2) the spatial and thematic map accuracy of the components, and (3) the

suitability of the results for the intended application.

Conclusions and Management Implications

Puerto Rico faces a number of problems that are common to much of the world.

Population is increasing while land area is not, and there is an ongoing reassessment of land use policy and practice to accommodate growing populations, shifting

economies, and changing public value systems. Puerto Rico shares similarities with

the Eastern United States with its history of agricultural abandonment, relatively

high population density, and abundance of “forested” suburban areas in the wildland-urban interface. Puerto Rico has affinities with other tropical regions with
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The Puerto Rico Gap Analysis Project

Melanerpes portoricensis, the Puerto Rican woodpecker, is endemic to the island.

Javier Mercado
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GENERAL TECHNICAL REPORT IITF-GTR-39

high biodiversity and an abundance of rare species. Puerto Rico shares similarities

with other islands in that land limitations are evident and abrupt at the terrestrialmarine interface and species diversity is controlled by the isolation of the island as

well as climatic, evolutionary, and historical factors.

Puerto Rico is unique in its assemblage of species and characteristics that combine all of the above, as well as the character of its people, government, policies,

and attitudes regarding conservation. There are a number of new initiatives within

governmental and nongovernmental agencies and private citizens groups to develop

better integrated, island-wide land use planning and policy. This is a very positive

step. There are also a number of initiatives to consider regional planning for conservation efforts including Caribbean-wide assessments of biodiversity and the

development of bird conservation efforts encompassing the Atlantic flyway. The

PRGAP is a valuable resource for these efforts both in the information we are

making available and as a model of data management and information development

and transfer.

The PRGAP is a comprehensive assemblage of information on Puerto Rico’s

land cover, vertebrate occurrences, natural history information, and land stewardship. The results of this project are:

• An important body of information on current land cover attributes,

terrestrial vertebrate species distributions, literature availability, and

conservation priorities. This information can serve as an educational tool, a

reference data set, as input for modeling, as a data set for analyses, and as a

source of new hypotheses for research.

• A set of integrated geospatial information on land cover, land use,

population dynamics, habitat use and dynamics, terrain characteristics,

physiography, geology, climate, hydrology, land ownership and

management, and species occurrences. This information is well documented

as to its sources and carefully co-registered for integrated scientific

analyses. It is designed to be used for analyses of PRGAP data, as well as

to produce map products and datasets that will be useful for education and

research.

• A set of databases that relate information on vertebrate occurrences,

taxonomy, conservation status, habitat characteristics, scientific literature,

life history traits, habitat distributions, and land management priorities.

These data sets have complete information on the terrestrial resident,

endemic, and endangered species in Puerto Rico, but also a framework and
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initial taxonomic information on all of Puerto Rico and the U.S. Virgin

Island’s 490 terrestrial, freshwater, and marine species. The databases are

designed to be maintained, updated, and expanded as long as institutional

support is available.

• A network of public and private individuals and organizations that have

provided input or will be future users of PRGAP products and information.

This report is a preliminary analysis of this comprehensive data set, and there is

still much information to be derived from the PRGAP. Some of these initial

findings indicate that the various physiographic regions of the island have distinct

roles in the maintenance of biodiversity of different taxonomic groups. The bird

species show highest diversity in the coastal plain and hills, with their mix of open

forest, grasslands, wetlands, saline and freshwater ponds, and streams. This part of

the island is under the greatest development pressure. The PRGAP information also

indicates that a number of mature forest habitats are key to the survival of endangered and endemic species. The northern karst region is an important center of both

mammal and reptile diversity. Reptile diversity and the presence of a number of

endemic species is also high in the dryland habitats. The moist forested mountain

peaks serve as reservoirs of amphibian diversity.

Protection of this landscape diversity is not uniform in Puerto Rico. The least

protected areas are the former agricultural lands in the coastal plain, low hills, and

the karst region, and these should be conservation priorities. Protection of diversity

could be improved by (1) total protected area of at least 15 percent; (2) expanded

reserves in the limestone hills and coastal plain, particularly the matrix of wetland

and upland vegetation as found on the former Roosevelt Roads and Sabana Seca

Naval Bases, the Northeast Ecological Corridor, Piñones, and other areas; (3) better

regulation of development in the periphery of existing reserves to maintain the

integrity of hydrologic systems in wetlands and to maintain viable corridors and

buffer zones in the upland reserves; (4) development of ecological corridors,

including riparian corridors to connect existing reserves with green space; (5)

development of small and intermediate-sized parks and open space within urban

areas that serve as habitat as well as recreational and educational resources for

communities; (6) the protection of unique habitats such as mountain valleys that

shelter Eleutherodactylus cooki (the guajón) and the freshwater nonforested wetlands that shelter Eleutherodactylus juanariveroii (the coqui llanero); and (7) the

restoration of formerly extensive habitats such as the freshwater swamps or riparian

forests of Pterocarpus officinalis Jacq. and the moist lowland ausubo (Manilkara

bidentata (A. DC.) Chev.) forests.
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GENERAL TECHNICAL REPORT IITF-GTR-39

Concluciones e Implicaciones de Manejo

Puerto Rico enfrenta una serie de problemas que son comunes al resto del mundo.

La población aumenta, mientras no así el área, y hay una continua reevaluación de

las políticas y las prácticas de uso del terreno para acomodar ese aumento. Además

Puerto Rico experimenta el vaivén económico, y el cambio en los valores del

sistema público. Puerto Rico comparte con el área este de los Estados Unidos una

historia de abandono agrícola, densidad poblacional relativamente alta y en

aumento, y abundancia de áreas suburbanas forestadas en la interfase urbanosilvestre. Puerto Rico tiene afinidades con otras regiones tropicales con gran

biodiversidad y abundancia de especies raras. También comparte similitudes con

otras islas en términos de las limitaciones de espacio, evidentes y abruptas en la

franja marítimo-terrestre, y en que la diversidad de las especies está controlada por

su aislamiento así como por factores climáticos, evolutivos e históricos.

Puerto Rico es único en su ensamblaje de especies y características que

combinan todo lo anteriormente mencionado, así como el carácter de su gente, su

gobierno, y sus actitudes con relación a la conservación. Existen en la isla un

número de nuevas iniciativas dentro de agencias tanto gubernamentales como nogubernamentales, y agrupaciones privadas con el fin de desarrollar políticas y

planes de uso de terreno a nivel isla mejor integrados, lo cual es un paso muy

positivo. También existen una serie de iniciativas para considerar planes regionales

en los esfuerzos de conservación incluyendo avalúos de conservación a lo largo del

Caribe y el desarrollo de esfuerzos de conservación de aves a lo largo de lo que se

conoce como la ruta migratoria del Atlántico (conocida en inglés como Atlantic

Flyway). El Proyecto Análisis Gap de Puerto Rico (PRGAP) es un recurso valioso

para tales esfuerzos tanto en la información que se está haciendo disponible, y

como modelo de manejo de datos y transferencia y desarrollo de información.

El PRGAP es una colección abarcadora de información de cobertura del

terreno en la isla, las distribuciones e historia natural de especies vertebradas, así

como el dominio de áreas de manejo con propósitos de conservación. Los

resultados de este proyecto se podrían resumir como:

• Una colección comprensiva de información de atributos de cobertura

terrestre actualizados, distribución de especies vertebradas terrestres,

disponibilidad de literatura científica, y prioridades de conservación. Esta

información representa una herramienta educacional, un conjunto de datos

de referencia, una herramienta de datos para modelación, un conjunto de

datos para análisis, y una fuente de hipótesis para la investigación.
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The Puerto Rico Gap Analysis Project

• Un conjunto de información geo-espacial sobre la cobertura del terreno, el

uso de suelos, dinámica de poblaciones, uso y dinámica del hábitat,

características del terreno, fisiografía, geología, clima, hidrología, tenencia

y manejo de tierras, y distribución geográfica de las especies de

vertebrados. Las fuentes de información están bien documentadas, y

cuidadosamente co-registradas para análisis científicos integrados. Está

diseñada para ser usada en el análisis de la información de los datos de

PRGAP, así como para la preparación de productos cartográficos y

cubiertas digitales de utilidad educacional y en lo que a la investigación

científica se refiere.

• Un conjunto de bases de datos que entrelazan datos de avistamientos de

vertebrados, taxonomía, estatus de conservación, características de hábitat,

literatura científica, rasgos de su historia natural, distribución de hábitat, y

prioridades de conservación. Este conjunto de datos incluyen información

completa sobre especies terrestres residentes, endémicas o amenazadas en

Puerto Rico, pero también una estructura inicial de información

taxonómica de todas las especies de vertebrados marinos, acuáticos de agua

dulce, y terrestres de Puerto Rico y las Islas Vírgenes de Estados Unidos.

Estas bases de datos están diseñadas para ser mantenidas, actualizadas, y

expandidas mientras dure el apoyo institucional.

• Una red de agencias públicas, individuos privados y organizaciones que han

proveído insumo y son o serán futuros usuarios de los productos y la

información de PRGAP.

Este reporte representa un análisis preliminar de este abarcador conjunto de

datos, y todavía existe el potencial de derivar nueva información del PRGAP.

Algunos de los hallazgos iniciales indican que las distintas regiones fisiográficas de

la isla tienen diferentes roles en mantener la biodiversidad de los diferentes grupos

taxonómicos estudiados. Las aves muestran mayor diversidad en los llanos y lomas

costeras, con su composición de bosques abiertos, pastizales, humedales, charcas y

quebradas de agua dulce y salada. Esta porción de la isla está bajo la mayor presión

de desarrollo urbano. La información del PRGAP también indica que un número

de ambientes de bosque maduro son clave para la supervivencia de especies

amenazadas y/o endémicas. La región del Carso es importante para la supervivencia

de la diversidad de mamíferos y reptiles. La zona de vida seca muestra también una

gran diversidad de reptiles y especies endémicas. Por otro lado, los bosques

húmedos de los picos montañosos representan un santuario para la diversidad de

anfibios.
104

GENERAL TECHNICAL REPORT IITF-GTR-39

La protección de esta diversidad de paisajes no es uniforme en Puerto Rico. Las

áreas menos protegidas son aquellos terrenos de las llanuras costeras que otrora

fuesen utilizados para la agricultura, las colinas bajas y la región del Carso, por lo

que estas regiones deberían representar prioridades de conservación. La protección

de la biodiversidad puede mejorarse, por ejemplo, (1) aumentando a 15% la

superficie total de áreas protegidas; (2) expandiendo las reservas en las regiones del

Carso y las planicies costeras, particularmente la matriz de humedales y vegetación

de tierras elevadas como aquellas localizadas en las antigua bases navales de

Roosevelt Roads y Sabana Seca, el Corredor Ecológico del Noreste, Piñones, y

otras áreas; (3) mejorando la regulación del desarrollo urbano en la periferia de las

reservas existentes para así mantener la integridad de los sistemas hidrológicos en

humedales y mantener corredores viables y zonas de amortiguamiento en las

reservas ubicadas a mayor elevación; (4) desarrollando corredores ecológicos, tales

como corredores ribereños, para conectar reservas existentes con espacios verdes;

(5) desarrollando parques medianos y pequeños y espacios abiertos entre áreas

urbanas que sirvan como hábitat así como recursos educativos y recreacionales para

las comunidades; (6) Protegiendo los hábitat únicos tales como los valles

montañosos que sirven de santuario al Coquí Guajón (Eleutherodactylus cooki), y

los humedales, ya sea los no boscosos de agua dulce, que cobijan a la recién

descubierta especie de anfibio Coquí Llanero, provisionalmente Eleutherodactylus

juanariveroii; y (7) restaurando hábitat antiguamente extensos, tales como los

pantanos de agua dulce o bosques ribereños de Pterocarpus officinalis Jacq. y los

bosques de ausubo (Manilkara bidentata (A. DC.) Chev.) de las planicies húmedas.
105

The Puerto Rico Gap Analysis Project

Product Use and Availability

How to Obtain Products

It is the goal of the Gap Analysis Program and the USGS Biological Resources

Division (BRD) to make the data and associated information as widely available as

possible. Use of the data requires specialized GIS software and substantial computing power. Additional information on how to use the data or obtain GIS services is

provided below and on the GAP home page (URL below). While a CD-ROM or

DVD of the data will be the most convenient way to obtain the data, it may also

be downloaded via the Internet from the national GAP home page at http://

www.gap.uidaho.edu/.

The home page will also provide, over the long term, the status of our state’s

project, future updates, data availability, and contacts. Within a few months of this

project’s completion, CD-ROMs or DVDs of the final report and data should be

available at a nominal cost—the above home page will provide ordering information. To find information on this state GAP project’s status and data, follow the

links to “Current Projects” and then to the particular state of interest.

Disclaimer

Following is the official BRD disclaimer as of 29 January 1996, followed by

additional disclaimers from GAP. Prior to using the data, you should consult the

GAP home page for the current disclaimer.

Although these data have been processed successfully on a computer system at

the BRD, no warranty expressed or implied is made regarding the accuracy or

utility of the data on any other system or for general or scientific purposes, nor

shall the act of distribution constitute any such warranty. This disclaimer applies

both to individual use of the data and aggregate use with other data. It is strongly

recommended that these data are directly acquired from a BRD server [see above

for approved data providers] and not indirectly through other sources which may

have changed the data in some way. It is also strongly recommended that careful

attention be paid to the content of the metadata file associated with these data. The

BRD shall not be held liable for improper or incorrect use of the data described or

contained herein.

These data were compiled with regard to the following standards. Please be

aware of the limitations of the data. These data are meant to be used at a scale of

1:100,000 or smaller (such as 1:250,000 or 1:500,000) for the purpose of assessing

the conservation status of animals and vegetation types over large geographic
106

GENERAL TECHNICAL REPORT IITF-GTR-39

regions. The data may or may not have been assessed for statistical accuracy. Data

evaluation and improvement may be ongoing. The BRD makes no claim as to the

data’s suitability for other purposes. This is writable data, which may have been

altered from the original product if not obtained from a designated data distributor

identified above.

Metadata

Proper documentation of information sources and processes used to assemble GAP

data layers is central to the successful application of GAP data.

Metadata is a description of the content, quality, lineage, contact, condition,

and other characteristics of data. It is a valuable tool that preserves the usefulness

of data over time by detailing methods for data collection and data set creation. It

greatly minimizes duplication of effort in the collection of expensive digital data

and fosters sharing of digital data resources. Metadata supports local data asset

management such as local inventory and data catalogs, and external user communities such as clearinghouses and Web sites. It provides adequate guidance for end-use

application of data such as detailed lineage and context. Metadata makes it possible

for data users to search, retrieve, and evaluate data set information by providing

standardized terms for geospatial and biological data.

The Federal Geographic Data Committee approved the Content Standard for

Digital Geospatial Metadata (FGDC-STD-001-1998) in June 1998, and the National Biological Information Infrastructure (NBII) (http://www.nbii.gov) developed the Biological Data Profile (approved in 1999) that adds fields for biological

information such as taxonomy, analytical tools, and methodology to the FGDC

standard core set of elements (http://www.nbii.gov/datainfo/metadata/standards/).

Executive Order 12906 requires that any spatial data sets generated with federal

dollars will have FGDC-compliant metadata.

Each PRGAP spatial data layer is accompanied by its metadata (*.xml or .sgml

file) in the same directory. We also include an additional directory (called

“meta_master’), which will include each metadata file in four forms (*.txt, *.xml,

*.html, and *.sgml).

Appropriate Use of These Data

The PRGAP is an important source of island-wide biological GIS information. The

data were created with the expectation that they would be used for other applications. Therefore, we list below both appropriate and inappropriate uses. This list is
107

The Puerto Rico Gap Analysis Project

in no way exhaustive but should serve as a guide to assess whether a proposed use

can or cannot be supported by GAP data. For most uses, it is unlikely that GAP

will provide the only data needed, and for uses with a regulatory outcome, field

surveys should verify the result. In the end, it will be the responsibility of each data

user to determine if GAP data can answer the question being asked, and if they are

the best tool to answer that question.

Scale—

The data were produced with an intended application at the regional level, that is,

geographic areas from tens of thousands to millions of hectares in size. The data

provide a coarse-filter approach to analysis, meaning that not every occurrence

of every plant community or animal species habitat is mapped–only larger, more

generalized distributions. The data are based on the USGS 1:100,000 scale of

mapping in both detail and precision. The PRGAP data can be used as a map for

a particular geographic area, or to provide regional context for a particular area.

The distinction can be made with the following example: You could use PRGAP

land cover to determine the approximate amount of mangrove forest occurring in a

municipality, or you could map mangrove with high-resolution aerial photography

to determine a more precise amount. You could use PRGAP data to determine the

approximate percentage of all mangrove forest in the region that occurs in a municipality, and thus gain a sense of how important the municipality’s distribution is to

maintaining that vegetation type.

Appropriate uses—

The above example illustrates two appropriate uses of the data: as a coarse map for

a large area such as a municipality, and to provide context for fine-level maps. The

following is a general list of applications:

• Island-wide biodiversity planning.

• Regional (Councils of Government) planning.

• Regional habitat conservation planning.

• Municipality or watershed comprehensive planning.

• Large-area resource management planning.

• Modeling future biodiversity or species distribution scenarios.

• Assessing habitat change.

• Coarse-filter evaluation of potential impacts or benefits of major projects or

plan initiatives on biodiversity, such as utility or transportation corridors,

wilderness proposals, or regional open space and recreation proposals.
108

GENERAL TECHNICAL REPORT IITF-GTR-39

• Determining relative amounts of management responsibility for specific

biological resources among land stewards to facilitate cooperative

management and planning.

• Basic research on regional distributions of plants and animals and to help

target both specific species and geographic areas for needed research.

• Environmental impact assessment for large projects or military activities.

• Estimation of potential economic impacts from loss of biological resourcebased activities.

• Education at all levels and for both students and citizens.

Inappropriate uses—

It is far easier to identify appropriate uses than inappropriate ones; however, there is

a “fuzzy line” that is eventually crossed when the differences in resolution of the

data, size of geographic area being analyzed, and precision of the answer required

for the question are no longer compatible. Examples include:

• Using the data to map small areas (less than thousands of hectares),

typically requiring mapping resolution at 1:24,000 scale and using aerial

photographs or ground surveys.

• Combining GAP data with other data finer than 1:100,000 scale to produce

new hybrid maps or answer queries.

• Generating specific overly precise areal measurements from the data

(minimum mapping unit size and accuracy affect this precision).

• Establishing exact boundaries for regulation or acquisition.

• Establishing definite occurrence or nonoccurrence of any feature for an

exact geographic area (for land cover, the percentage of accuracy will

provide a measure of probability).

• Determining abundance, health, or condition of any feature.

• Establishing a measure of accuracy of any other data by comparison with

GAP data.

• Altering the data in any way and redistributing them as a PRGAP data

product.

• Using the data without acquiring and reviewing the metadata and this

report.
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The Puerto Rico Gap Analysis Project

Acknowledgments

This work has been accomplished with the assistance and support of a great number

of people. Their support and enthusiasm for conservation has in many cases been

inspirational. First of all we thank Jaime Collazo and Eileen Helmer for bringing

this project to Puerto Rico and the U.S. Forest Service International Institute of

Tropical Forestry (IITF). The institutional support provided IITF and the Puerto

Rico Department of Natural and Environmental Resources (DNER) was very

important. Thanks to Ariel Lugo, José Berrios, and José Chabert. Kevin Gergley,

program manager of the USGS Biological Resources Division, National Gap

Program has supported this work. The staff at the National Gap office, Jocelyn

Aycrigg, Jill Maxwell, Anne Davidson, and Nicole Coffey, have been a great help.

Steve Williams and Alexa McKerrow, working with Jaime Collazo and the North

Carolina and Southeast Gap projects have offered advice and experience since the

project was initiated. Tom Brandeis assisted in developing the PRGAP hexagon

layer.

This report draws on the GAP handbook template for final reports.

We have been assisted by a number of volunteers at IITF, and we appreciate the

support of Elizabeth Hernandez, coordinator of the volunteer program, in recruiting and assisting local, national, and international volunteers. Those volunteers

include Milagros González, Jessica Acosta, Isabel Pares, and Angelica Vega.

A number of scientists at IITF have been a source of feedback, expertise and

encouragement including Wayne Arendt, Grizelle González, Eileen Helmer, Jean

Lodge, Ariel Lugo, Frank Wadsworth, Pete Weaver, and Joe Wunderle. International Institute of Tropical Forestry field biologists were a source of expertise about

the flora and fauna of the island, including Iván Vicens, Javier Mercado, Matthew

Anderson, Marcos Rodríguez, and Juan Rivera. The project was completed in the

IITF Geographic Information Systems and Remote Sensing Laboratory, and we

would like to thank all the people in the lab for their support and enthusiasm,

including Olga Ramos-González, Brook Edwards, and ecologist Carlos Rodríguez.

A number of biologists and program managers at the DNER have offered time

and expertise including Eduardo Ventosa, María Camacho, José Sustache, Eduardo

Cintrón, Edgardo González, Ernesto Diaz, Nora Álvarez, and Danny Dávila.

We also thank Jorge Paniagua, Sandra Coria, and Jerry Bauer for help in

producing a brochure about the project.
110

GENERAL TECHNICAL REPORT IITF-GTR-39

The scientific content and credibility in this report is due in great part to the

participation of a number of field biologists. Thanks to José Salguero, from the

Puerto Rican Ornithological Society, who was involved in many aspects of the

PRGAP Analysis Project and contributed with bird occurrence records, also special

thanks to Luis J. Villanueva (University of Puerto Rico, Río Piedras Campus),

Fernando Bird Picó and Juan A. Rivero (University of Puerto Rico, Mayagüez

Campus), who contributed with amphibian and reptile occurrence records. Michael

Gannon provided bat distribution data. Frank Rivera Milán provided comments and

data on pigeons. Jaime Collazo provided a number of important bird occurrence

records for the karst area. Special thanks to the many reserve and forest managers

who welcomed us and often took time from their responsibilities to escort us

during ground visits. They provided us with information about protected lands in

Puerto Rico. We appreciate your time and support.

We are grateful to the reviewers who carried out priceless efforts reviewing the

distribution and predicted habitat maps and the species accounts. Special thanks to

expert reviewers Javier Mercado, Matthew Anderson, Alberto Estrada, Alberto

Puente, Carlos Díaz, Francisco J. Vilella, Gilbert Bonilla Martinez, Jorge Saliva,

Joseph Wunderle, Katsi Ramos Alvarez, Luis J. Villanueva, Luis Jorge Rivera

Herrera, Ricardo López Ortiz, Roseanne Medina Miranda, Sondra Vega, Verónica

Anadón, Gail Susana Ross, and many others who supported and contributed to the

PRGAP.

Research at the International Institute of Tropical Forestry is done in collaboration with the University of Puerto Rico.
111

The Puerto Rico Gap Analysis Project

Agradecimientos

Este trabajo fue hecho con la ayuda y el apoyo de un gran número de personas. Su

apoyo y entusiasmo por la conservación han sido, en muchos casos, inspirador.

Primero quiero darles las gracias a Jaime Collazo y a Eileen Helmer por traer este

proyecto a Puerto Rico y al Instituto Internacional de Dasonomía Tropical (IITF

por sus siglas en inglés). El apoyo institucional del IITF y del Departamento de

Recursos Naturales y Ambientales de Puerto Rico (DRNA) fue de suma

importancia, gracias a Ariel Lugo, José Berríos, y José Chabert. Kevin Gergley,

administrador de la División de Recursos Biológicos del Servicio Geológico de los

Estados Unidos (USGS) del Programa Nacional de GAP, ha apoyado este trabajo.

El personal en la oficina Nacional de GAP, Jocelyn Aycrigg, Jill Maxwell, Anne

Davidson, y Nicole Coffey, han sido de gran ayuda. Steve Williams y Alexia

McKerrow, trabajando con Jaime Collazo y los proyectos de GAP de Carolina del

Norte y el Sureste de Estados Unidos han ofrecido consejos y experiencia desde que

el inicio del proyecto. Tom Brandeis participó en el desarrollo de la capa de

hexágonos del Proyecto Gap Analysis de Puerto Rico.

Este reporte hace uso del texto del Manual GAP para los reportes finales.

Hemos recibido la generosa ayuda de varios voluntarios en el IITF y nos

gustaría reconocer el apoyo de Elizabeth Hernández, coordinadora del programa de

voluntarios, por enlistar y ayudar a voluntarios locales, e internacionales. Esos

voluntarios incluyen Milagros González, Jessica Acosta, Isabel Pares, y Angélica

Vega.

Varios científicos en IITF han sido una fuente de conocimiento, consejos y

apoyo incluyendo a: Wayne Arendt, Grizelle González, Jean Lodge, Ariel E. Lugo,

Frank Wadsworth, Peter Weaver, y Joseph Wunderle. Los biólogos de campo del

IITF fueron una fuente de conocimiento experto sobre la flora y fauna de la Isla, e

incluye a: Iván Vicens, Javier Mercado, Matthew Anderson, Marcos Rodríguez, y

Juan Rivera. El proyecto fue completado en el Laboratorio de GIS y Percepción

Remota del IITF y le doy gracias a todas las personas en el laboratorio por su

apoyo y entusiasmo, incluyendo a: Olga Ramos-González, Maya Quiñones, Brook

Edwards, y Carlos Rodríguez.

Varios biólogos y directores en el DRNA han ofrecido de su tiempo y

experiencia, incluyendo a: Eduardo Ventosa, María Camacho, José Sustache,

Eduardo Cintrón, Edgardo González, Ernesto Díaz, Nora Álvarez, y Danny Dávila.

Queremos darles las gracias a Jorge Paniagua, Sandra Coria, y Jerry Bauer por

su ayuda en la producción del folleto informativo del PRGAP.
112

GENERAL TECHNICAL REPORT IITF-GTR-39

El contenido científico y credibilidad en este informe son debidos en gran parte

a la participación de varios biólogos de campo. Nuestros agradecimientos a José

Salguero, de la Sociedad Ornitológica Puertorriqueña, quién estuvo envuelto en

muchos aspectos del Proyecto PRGAP y contribuyó con registros de avistamientos

de aves. Gracias también a Luis J. Villanueva (Universidad de Puerto Rico, Recinto

de Río Piedras), Fernando Bird Picó y Juan A. Rivero (Universidad de Puerto Rico,

Recinto de Mayagüez), que contribuyeron con registros de ocurrencia de anfibios y

reptiles. Michael Gannon proveyó datos de distribución de murciélagos. Frank

Rivera Milán proveyó comentarios y datos sobre columbidos. Jaime Collazo de la

Universidad Estatal de Carolina del Norte proveyó varios registros importantes de

observaciones de aves para el área de karso. Gracias especiales a los muchos

directores de reservas y bosques que nos dieron la bienvenida y a menudo tomaron

tiempo de sus responsabilidades para acompañarnos durante nuestras visitas de

campo. Ellos nos proveyeron con información sobre tierras protegidas en Puerto

Rico. Apreciamos su tiempo y apoyo.

Gracias a las personas que participaron de la revisión de los datos por sus

esfuerzos imprescindibles sobre los mapas de distribución y predicción de hábitat,

así como los avistamientos de especies. Un agradecimiento especial a todos los

expertos: Javier Mercado, Matthew Anderson, Alberto Estrada, Alberto Puente,

Carlos Díaz, Francisco J. Vilella, Gilbert Bonilla Martínez, Jorge Saliva, Joseph

Wunderle, Katsi Roseanne Medina Miranda, Sondra Vega, Verónica Anadón, Gail

Susana Ross y muchos otros que nos brindaron apoyo y contribuyeron al PRGAP.

¡Muchas gracias!

Las investigaciones en el Instituto Internacional de Dasonomía Tropical se

hacen en colaboración de la Universidad de Puerto Rico.

English Equivalents

When you know: Multiply by: To find:

Millimeters (mm) 0.0394 Inches

Meters (m) 3.28 Feet

Square meters (m2

) 10.76 Square feet

Kilometers (km) 0.621 Miles

Square kilometers (km2

) 0.386 Square miles

Degrees Celsius (°C) 1.8 °C + 32 Degrees Fahrenheit
113

The Puerto Rico Gap Analysis Project

Literature Cited

Ahern, F.; Belward, A.; Churchill, P.; Davis, R.A.; Janetos, A.; Justice, C.O.;

Loveland, T.; Malingreau, J.-P.; Maiden, M.; Skole, D.; Taylor, V.;

Yasuoka, Y.; Zhu, Z. 1999. Strategy for global observation of forest cover.

Report 2. Ottawa, Canada: Canadian Center for Remote Sensing. 58 p.

Aide, T.M.; Grau, H.R. 2004. Enhanced: globalization, migration, and Latin

America ecosystems. Science. 305(5692): 1915–1916.

Alvárez, M. 1982. A comparison of the structure and ecology of Pterocarpus

officinalis (Jacq.) forested wetlands in Puerto Rico. Río Piedras, PR: University

of Puerto Rico. M.S. thesis.

Areces-Mallea, A.E.; Weakley, A.S.; Li, X. ; Sayre, R.G. ; Parrish, J.D.;

Tipton, C.V.; Boucher, T. 1999. A guide to Caribbean vegetation types:

preliminary classification system and descriptions. Arlington, VA: The Nature

Conservancy. 59 p.

Austin, M.P. 1991. Vegetation: data collection and analysis. In: Margules, C.R.;

Austin, M.P., eds. Nature conservation: cost effective biological surveys and

data analysis. East Melbourne, Australia: Commonwealth Scientific and Research Organization: 37–41.

Bawiec, W.J. 2001. Geology, geochemistry, geophysics, mineral occurrences and

mineral resource assessment for the Commonwealth of Puerto Rico. Open File

report 98–38. Reston, VA: U.S. Geological Survey. [CD-ROM].

Biaggi, V. 1997. Las aves de Puerto Rico. 4th ed. San Juan, PR: Editorial de la

Universidad de Puerto Rico. 389 p.

Birdsey, R.A.; Weaver, P.L. 1982. The forest resources of Puerto Rico. Resour.

Bull. SO-85. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. 60 p.

Birdsey, R.A.; Weaver, P.L. 1987. Forest area trends in Puerto Rico. Res. Note.

SO-331. New Orleans, LA: U.S. Department of Agriculture, Forest Service,

Southern Forest Experiment Station. 5 p.

Boone, R.B. 1996. An analysis of terrestrial vertebrate diversity in Maine. Orono,

ME: University of Maine. Ph.D. thesis.
114

GENERAL TECHNICAL REPORT IITF-GTR-39

Borhidi, A. 1996. Phytogeography and vegetation ecology of Cuba. Budapest,

Hungary: Akadémiai Kaidó. 859 p.

Bourgeron, P.S.; Humphries, H.C.; DeVelice, R.L.; Jensen, M.E. 1994.

Ecological theory in relation to landscape and ecosystem characterization.

In: Jensen, M.E.; Bourgeron, P.S., eds. Ecosystem management: principles

and applications. Gen. Tech. Rep. PNW-GTR-318. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station:

58–72. Vol. II.

Britton, N.L.; Wilson, P. 1924. Scientific survey of Porto Rico and the Virgin

Islands. Botany of Porto Rico and the Virgin Islands. New York, NY: New York

Academy of Sciences. 626 p. Vol. 5.

Chinea, J.D. 2002. Tropical forest succession on abandoned farms in the Humacao

Municipality of eastern Puerto Rico. Forest Ecology and Management. 167:

195–207.

Cintrón, G. 1991. Introduction to Mona Island. Acta Cientifica. 5: 6–9.

Crist, P.; Thompson, B.; Prior-Magee, J. 1995. A dichotomous key of land

management categorization. Unpublished manuscript. On file with: New

Mexico Cooperative Fish and Wildlife Research Unit, University of New

Mexico, Las Cruces, NM 88003.

Csuti, B. 1994. Methods for developing terrestrial vertebrate distribution maps for

gap analysis (version 1). In: Scott, J.M.; Jennings, M.D., eds. A handbook for

gap analysis. Moscow, ID: Idaho Cooperative Fish and Wildlife Research Unit,

University of Idaho: 2.1–2.5.2.

Csuti, B.; Crist, P. 1998. Methods for assessing accuracy of animal distribution

maps (version 2.01). In: A handbook for conducting gap analysis. Moscow, ID:

U.S. Geological Survey Gap Analysis Program. http://www.gap.uidaho.edu/

handbook. (December 2006).

Csuti, B.; Polasky, S.; Williams, P.H.; Pressey, R.L.; Camm, J.D.; Kershaw,

M.; Kiester, A.R.; Downs, B.; Hamilton, R.; Huso, M.; Sahr, K. 1997. A

comparison of reserve selection algorithms using data on terrestrial vertebrates

in Oregon. Biological Conservation. 80: 83–97
115

The Puerto Rico Gap Analysis Project

Daly, C.; Helmer, E.H.; Quiñones, M. 2003. Mapping the climate of Puerto Rico,

Vieques and Culebra. International Journal of Climatology. 23(11):

1359–1381.

Dansereau, P. 1966. Studies on the vegetation of Puerto Rico. 1. Description and

integration of the plant communities. Spec. Publ. 1. Mayagüez, PR: Institute of

Caribbean Science. 45 p.

Davis, F.W.; Stine, P.A.; Stoms, D.M.; Borchert, M.I.; Hollander, A.D. 1995.

Gap analysis of the actual vegetation of California. 1. The southwestern region.

Madroño. 42: 40–78.

Delannoy, C.A. 2005. Aves. In: Joglar, R.L., ed. Biodiversidad de Puerto Rico—

vertebrados terrestres y ecosistemas. San Juan, PR: Instituto de Cultura

Puertorriqueña: 241–278.

Diamond, J. 1986. The design of a nature reserve system for Indonesian New

Guinea. In: Soulé, M.E., ed. Conservation biology: the science of scarcity and

diversity. Sunderland, MA: Sinauer Associates: 485–503.

Driscoll, R.S.; Merkel, D.L.; Hagihara, J.S.; Radloff, D.L. 1983. A component

land classification for the United States: status report. Tech. Note 360. Denver,

CO: U.S. Department of the Interior, Bureau of Land Management. 30 p.

Driscoll, R.S.; Merkel, D.L.; Radloff, D.L.; Snyder, D.E.; Hagihara, J.S. 1984.

An ecological land classification framework for the United States. Misc. Publ.

1439. Washington, DC: U.S. Department of Agriculture, Forest Service. 56 p.

Edwards, T.C., Jr.; Homer, C.H.; Bassett, S.D.; Falconer, A.; Ramsey, R.D.;

Wight, D.W. 1995. Utah gap analysis: an environmental information system.

Tech. Rep. 95-1. Logan, UT: Utah Cooperative Fish and Wildlife Research

Unit, Utah State University. 1189 p. + 2 CD-ROMs.

Eusse, A.M.; Aide, T.M. 1999. Patterns of litter production across a salinity

gradient in a Pterocarpus officinalis tropical wetland. Plant Ecology. 145:

307–315.

Ewel, J.J.; Whitmore, J.L. 1973. The ecological life zones of Puerto Rico and the

U.S. Virgin Islands. Res. Pap. ITF-18. Río Piedras, PR: U.S. Department of

Agriculture, Forest Service, Institute of Tropical Forestry. 72 p.
116

GENERAL TECHNICAL REPORT IITF-GTR-39

Federal Geographic Data Committee [FGDC]. 1997. Vegetation classification

and information standards. Reston, VA: Vegetation subcommittee, FGDC

Secretariat. 35 p.

Figueroa Colón, J.C.; Woodbury, R.O. 1996. Rare and endangered plant species

of Puerto Rico and the Virgin Islands: an annotated checklist. Annals of New

York Academy of Sciences. 776: 65–71.

Fitzpatrick-Lins, K. 1981. Comparison of sampling procedures and data analysis

for a land-use and land-cover map. Photogrammetric Engineering and Remote

Sensing 47: 343–351.

Forman, R.T.T.; Godron, M. 1986. Landscape ecology. New York: John Wiley

and Sons. 640 p.

Franco, P.A.; Weaver, P.L.; Eggen-McIntosh, S. 1997. Forest resources of Puerto

Rico, 1990. Ashville, NC: U.S. Department of Agriculture, Forest Service,

Southern Research Station. 45 p.

Franklin, J.F. 1993. Preserving biodiversity: species, ecosystems, or landscapes?

Ecological Applications. 3(2): 202–205.

Gannon, M.R.; Kurta A.; Rodríguez-Durán, A.; Willig, M.R. 2005. Bats of

Puerto Rico—an island focus and Caribbean perspective. Lubbock, TX: Texas

Tech University Press. 239 p.

U.S. Geological Survey [USGS]. 2000. A handbook for conducting gap analysis.

Moscow, ID: Gap Analysis Program. http://www.gap.uidaho.edu/gap/handbook.

(September 10, 2007).

Gould, W.A.; Edwards, B.; Jiménez, M.E; Martinuzzi, S. 2008a. Physiography

of the Puerto Rican Archipelago. Scale 1: 260,000. IITF-RMAP-07. Río

Piedras, PR: U.S. Department of Agriculture, Forest Service, International

Institute of Tropical Forestry.

Gould, W.A.; Fevold, B.; Jiménez, M.E. 2008b. PRGAP hex map. Scale 1:

260,000. IITF-RMAP-08. Río Piedras, PR: U.S. Department of Agriculture,

Forest Service, International Institute of Tropical Forestry.

Gould, W.A.; González, G.; Carrero Rivera, G. 2006. Structure and composition

of vegetation along an elevational gradient in Puerto Rico. Journal of Vegetation

Science. 17: 653–664.
117

The Puerto Rico Gap Analysis Project

Gould, W.A.; Jiménez, M.E.; Martinuzzi, S. 2008c. Geoclimate and topography:

defining landscape units in Puerto Rico. Scale 1: 260,000. IITF-RMAP-06. Río

Piedras, PR: U.S. Department of Agriculture, Forest Service, International

Institute of Tropical Forestry.

Gould, W.A.; Martinuzzi, S.; Jiménez, M.E.; Edwards, B.; Ramos-González,

O.M. 2008d. Simplified landforms of Puerto Rico. Scale 1: 260,000. IITFRMAP-04. Río Piedras, PR: U.S. Department of Agriculture, Forest Service,

International Institute of Tropical Forestry.

Gould, W.A.; Solórzano, M.; Jiménez, M.E.; Alarcón, C. 2008e. Land stewardship in Puerto Rico: federal, commonwealth, and non-governmental ownership

of conservation areas. Scale 1: 260,000. IITF-RMAP-13. Río Piedras, PR: U.S.

Department of Agriculture, Forest Service, International Institute of Tropical

Forestry.

Gould, W.A.; Solórzano, M.; Jiménez, M.E.; Alarcón, C. 2008f. Land stewardship in Puerto Rico: federal, commonwealth, and non-governmental conservation management areas. Scale 1: 260,000. IITF-RMAP-14. Río Piedras, PR:

U.S. Department of Agriculture, Forest Service, International Institute of

Tropical Forestry.

Gould, W.A.; Solórzano, M.; Jiménez, M.E.; Alarcón, C. 2008g. Land stewardship in Puerto Rico: management priorities for conservation. Scale 1: 260,000.

IITF-RMAP-15. Río Piedras, PR: U.S. Department of Agriculture, Forest

Service, International Institute of Tropical Forestry.

Goward, S.N.; Tucker, C.J.; Dye, D.G. 1985. North American vegetation patterns

observed with NOAA-7 advanced very high resolution radiometer. Vegetatio.

64: 3–14.

Grau, H.R.; Aide, T.M.; Zimmerman, J.K.; Thomlinson, J.R.; Helmer, E.;

Zou, X. 2003. The ecological consequences of socioeconomic and land-use

changes in postagriculture Puerto Rico. Bioscience. 53: 1159–1168.

Grossman, D.; Goodin, K.L.; Li, X.; Wisnewski, C.; Faber-Langendoen, D.;

Anderson, M.; Sneddon, L.; Allard, D.; Gallyoun, M.; Weakley, V. 1994.

Standardized national vegetation classification system. Report by The Nature

Conservancy and Environmental Systems Research Institute for the NBS/NPS

Vegetation Mapping Program. Denver, CO: U.S. Department of the Interior,

National Biological Service.
118

GENERAL TECHNICAL REPORT IITF-GTR-39

Helmer E.H. 2004. Forest conservation and land development in Puerto Rico.

Landscape Ecology. 19: 29–40.

Helmer, E.H.; Ramos-González, O.M.; López, T. del M.; Díaz, W. 2002.

Mapping the forest type and land cover of Puerto Rico, a component of

the Caribbean biodiversity hotspot. Caribbean Journal of Science. 38(3–4):

165–183.

Holdridge, L.R. 1967. Life zone ecology. San Jose, Costa Rica: Tropical Science

Center. 206 p.

Jenkins, R.E. 1985. Information methods: why the heritage programs work. The

Nature Conservancy News. 35(6): 21–23.

Jennings, M.D. 1993. Natural terrestrial cover classification: assumptions and

definitions. Gap Analysis Tech. Bull. 2. Moscow, ID: Idaho Cooperative Fish

and Wildlife Research Unit, University of Idaho. 28 p.

Jennings, M.D. 2000. Gap analysis: concepts, methods, and recent results. Landscape Ecology. 15: 5–20.

Joglar L.R. 1998. Los coquíes de Puerto Rico, su historia natural y conservación.

Río Piedras, PR: Editorial de la Universidad de Puerto Rico. 232 p.

Joglar, R.L., ed. 2005. Biodiversidad de Puerto Rico–vertebrados terrestres y

ecosistemas. San Juan, PR: Instituto de Cultura Puertorriqueña. 563 p.

Kiester, A.R.; Scott, J.M.; Csuti, B.; Noss, R.F.; Butterfield, B.; Sahr, K.;

Whit, D. 1996. Conservation prioritization using GAP data. Conservation

Biology. 10: 1332–1342.

Kirkpatrick, J.B. 1983. An iterative method for establishing priorities for the

selection of nature reserves: an example from Tasmania. Biological Conservation. 25: 127–134.

Küchler, A.W.; Zonneveld, I.S., eds. 1988. Vegetation mapping. Dordrecht, The

Netherlands: Lower Academic Publishers. 635 p.

Landis, J.R.; Koch, G.C. 1977. The measurement of observer agreement for

categorical data. Biometrics. 33: 159–174.

Levin, S.A. 1981. The problem of pattern and scale in ecology. Ecology.

73: 1942–1968.
119

The Puerto Rico Gap Analysis Project

Lin, T.; Laporte, N.T. 2003. Using IKONOS and aerial videography to validate

Landsat land cover maps of central African tropical rain forests. Abstract B41B0880. San Francisco, CA: American Geophysical Union.

Lins, K.S.; Kleckner, R.L. 1996. Land cover mapping: an overview and history

of the concepts. In: Scott, J.M.; Tear, T.H.; Davis, F., eds. Gap analysis: a

landscape approach to biodiversity planning. Bethesda, MD: American Society

for Photogrammetry and Remote Sensing: 57–65.

Lopez, T. del M.; Aide, T.M.; Thomlinson, J.R. 2001. Urban expansion and the

loss of prime agricultural lands in Puerto Rico. Ambio. 30: 49–54.

Lugo, A.E. 1988. Estimating reductions in the diversity of tropical forest species.

In: Wilson, E.O.; Peter, F.M., eds. Biodiversity. Washington, DC: National

Academy Press: 58–69

Lugo, A.E. 2005. Los Bosques. In: Joglar, R.L., ed. Biodiversidad de Puerto

Rico–vertebrados terrestres y ecosistemas. San Juan, PR: Instituto de Cultura

Puertorriqueña: 241–278.

Lugo, A.E.; Helmer, E. 2004. Emerging forest on abandoned land: Puerto Rico’s

new forest. Forest Ecology and Management. 190: 145–161.

Maingi, J.K.; Marsh, S.E.; Kepner, S.E.; Edmonds, W.G. 2002. An accuracy

assessment of 1992 Landsat-MSS derived land cover for the upper San Pedro

watershed (U.S./Mexico). EPA/600/R-02/040. Washington, DC: Environmental

Protection Agency. 28 p.

Margules, C.R.; Nicholls, A.O.; Pressey, R.L. 1988. Selecting networks of

reserves to maximize biological diversity. Biological Conservation. 43: 63–76.

Martinuzzi, S.; Gould, W.A.; Ramos-González, O.M. 2006. Creating cloud-free

Landsat ETM+ data sets in tropical landscapes: cloud and cloud-shadow removal. Gen. Tech. Rep. IITF-32. Río Piedras, PR: U.S. Department of Agriculture, Forest Service, International Institute of Tropical Forestry. 12 p.

Martinuzzi, S.; Gould, W.A.; Ramos-González, O.M. 2007a. Land development, land use, and urban sprawl in Puerto Rico: integrating remote sensing and

population census data. Landscape and Urban Planning. 79: 288–297.

Martinuzzi, S.; Gould, W.A.; Ramos-González, O.M.; Edwards, B. 2007b.

Development of a landform model for Puerto Rico and its application for land

cover change analysis. Caribbean Journal of Science. 43(2): 161–171.
120

GENERAL TECHNICAL REPORT IITF-GTR-39

McNeely, J.A.; Miller, K.R., eds. 1994. National parks conservation and development. Washington, DC: Smithsonian Institution. 825 p.

Mount H.R.; Lynn, W.C. 2004. Soil survey laboratory data and soil descriptions

for Puerto Rico and the Virgin Islands. Soil Survey Investigation Report 50.

Lincoln, NE: U.S. Department of Agriculture, Natural Resources and Conservation Service. 202 p.

Mueller-Dombois, D.; Ellenberg, H. 1974. Aims and methods of vegetation

ecology. New York: John Wiley & Sons. 547 p.

Myers, N.; Mittermeier, R.A.; Mittermeier, C.G.; da Fonseca, G.A.B.; Kent, J.

2000. Biodiversity hotspots for conservation priorities. Nature. 203: 853–858.

Nicholls, A.O.; Margules, C.R. 1993. An upgraded reserve selection algorithm.

Biological Conservation. 64: 165–169.

Noss, R.F. 1987. From plant communities to landscapes in conservation inventories:

a look at The Nature Conservancy (USA). Biological Conservation. 41: 11–37.

Noss, R.F. 1990. Indicators for monitoring biodiversity: a hierarchical approach.

Conservation Biology. 4: 355–364.

Noss, R.F.; Cooperrider, A.Y. 1994. Saving nature’s legacy. Washington, DC:

Island Press. 416 p.

Noss, R.F.; LaRoe, E.T., III; Scott, J.M. 1995. Endangered ecosystems of the

United States: a preliminary assessment of loss and degradation. Biol. Rep. 28,

Washington, DC: U.S. Department of the Interior, National Biological Service.

95 p.

Oberle, M.W. 2000. Puerto Rico’s birds in photographs. 2nd ed. San Juan, PR:

Editorial Humanitas. 129 p.

Odum, E.D.; Odum, H.T. 1972. Proceedings of the North American Wildlife and

Natural Resources conference. 39: 178.

Pares, I.; Gould, W.A.; Aide, T.M. [In press]. Suburban growth and forest

expansion following agricultural abandment in Puerto Rico (1991–2000).

Ecology and Society. Special issue.

Peterson, A.T.; Kluza, D.A. 2003. New distributional modelling approaches for

gap analysis. Animal Conservation. 6: 47–54.
121

The Puerto Rico Gap Analysis Project

Pressey, R.L.; Humphries, C.J.; Margules, C.R.; Vane-Wright, R.I.; Williams,

P.H. 1993. Beyond opportunism: key principles for systematic reserve selection. Trends in Ecology and Evolution. 8: 124–128.

Pressey, R.L.; Logan, V.S. 1995. Reserve coverage and requirements in relation to

partitioning and generalization of land classes: analyses for western New South

Wales. Conservation Biology. 9(6): 1506–1517.

Pressey, R.L.; Nicholls, A.O. 1989. Application of a numerical algorithm to the

selection of reserves in semi-arid New South Wales. Biological Conservation.

50: 263–278.

Raffaele, H.A. 1983. A guide to the birds of Puerto Rico and the Virgin Islands.

San Juan, PR: Fondo Educativo Interamericano. 255 p.

Raffaele, H.A.; Wiley, J.; Garrido, O.; Keith, A.; Raffaele, J. 1998. A guide to

the birds of the West Indies. Princeton, NJ: Princeton University Press. 512 p.

Ramos-González, O.M.; Lugo, A.E. 1994. Mapa de la vegetación de Puerto Rico.

Acta Científica. 8: 63–66.

Ramos-González, O.M. 2001. Assessing vegetation and land cover changes

in northeastern Puerto Rico: 1978-1995. Caribbean Journal of Science.

37: 95–106.

Ride, W.L.D. 1975. Towards an integrated system: a study of selection and acquisition of national parks and nature reserves in Western Australia. In: Fenner, F.,

ed. A national system of ecological reserves in Australia. Canbaberra, Australia:

Australian Academy of Sciences: 64–85.

Ríos-López, N.; Thomas, R. 2007. New palustrian Eleutherodactylus (Anura:

Leptodactylidae) from Puerto Rico. Zootaxa. 1512: 51–64.

Rivera, L.W.; Aide, T.M. 1998. Forest recovery in the karst region of Puerto Rico.

Forest Ecology and Management. 108: 63–75.

Rivero, J.A. 1998. Los anfibios y reptiles de Puerto Rico [The amphibians and

reptiles of Puerto Rico]. 2nd ed. San Juan, PR: Editorial de la Universidad de

Puerto Rico. 510 p.

Rodríguez-Durán, A. 2005. Murciélagos. In: Joglar, R.L., ed. Biodiversidad de

Puerto Rico—Vertebrados Terrestres y Ecosistemas. San Juan, PR: Instituto de

Cultura Puertorriqueña: 241–278.
122

GENERAL TECHNICAL REPORT IITF-GTR-39

Scott, J.M.; Davis, F.; Csuti, B.; Noss, R.; Butterfield, B.; Groves, C.; Anderson, H.; Caicco, S.; D’Erchia, F.; Edwards, T.C., Jr.; Ulliman, J.; Wright,

G. 1993. Gap analysis: a geographic approach to protection of biological

diversity. Onalaska, WI: Wildlife Monographs. 123 p.

Schwartz, A.; Henderson, W.R. 1991. Amphibians and reptiles of the West Indies:

descriptions, distributions, and natural history. Gainesville, FL: University of

Florida Press. 720 p.

Sowa, S.P.; Annis, G.; Morey, M.E.; Diamond, D.D. 2007. A gap analysis and

comprehensive conservation strategy for riverine ecosystems of Missouri.

Ecological Monographs. 77(3): 301–334.

Specht, R.L. 1975. The report and its recommendations. In: Fenner, F., ed.

A national system of ecological reserves in Australia. Report 19. Canberra,

Australia: Australian Academy of Sciences: 11–16.

Stoms, D.M. 1994. Actual vegetation layer. In: Scott, J.M.; Jennings, M.D., eds.

A handbook for gap analysis. Moscow, ID: Idaho Cooperative Fish and Wildlife

Research Unit: 1.1–1.6.

Stoms, D.; Estes, J. 1993. A remote sensing research agenda for mapping

and monitoring biodiversity. International Journal of Remote Sensing.

14: 1839–1860.

United Nations Educational, Scientific, and Cultural Organization

[UNESCO]. 1973. International classification and mapping of vegetation.

Paris. 102 p.

U.S. Fish and Wildlife Service [USFWS]. 2005. National wetlands inventory

digital database. http://wetlands.fws.gov/mapper_tool.htm. (December 2005).

U.S. Geological Survey [USGS]. 2000. A handbook for conducting gap analysis.

Moscow, ID: Gap Analysis Program. http://www.gap.uidaho.edu/gap/handbook.

(September 10, 2007).

van Genderen, J.L.; Lock, B.F. 1977. Testing land use map accuracy. Photogrammetric Engineering and Remote Sensing. 43: 1135–1137.

Wadsworth, F. 1951. Caribbean Forester. 12(3): 93–114.

Weaver, P.L. 1990. Succession in the Elfin Woodland of the Luquillo Mountains of

Puerto Rico. Biotropica. 22: 83–89.
123

The Puerto Rico Gap Analysis Project

Weaver, P.L. 1991. Environmental gradients affect forest composition in the

Luquillo Mountains of Puerto Rico. Interciencia. 16: 142–151.

Weaver, P.L. 1994. Baño de Oro Natural Area Luquillo Mountains, Puerto Rico.

Gen. Tech. Rep. SO-111. New Orleans, LA: U.S. Department of Agriculture,

Forest Service, Southern Forest Experimental Station.

Weaver, P.L.; Medina, E.; Pool, D.; Dugger, K.; Gonzales-Liboy, J.; Cuevas, E.

1986. Ecological observations in the dwarf cloud forest of the Luquillo Mountains in Puerto Rico. Biotropica. 18: 79–85.

Whittaker, R.H. 1960. Vegetation of the Siskiyou Mountains, Oregon and

California. Ecological Monographs. 30(3): 279–338.

Whittaker, R.H. 1965. Dominance and diversity in land plant communities.

Science. 147: 250–259.

Whittaker, R.H. 1977. Species diversity in land communities. Evolutionary

Biology. 10: 1–67.

Williams, P.H.; Gibbons, D.; Margules, C.; Rebelo, A.; Humphries, C.;

Pressey, R. 1996. A comparison of richness hotspots, rarity hotspots, and

complementary areas for conserving diversity of British birds. Conservation

Biology. 10: 155–174.

Zhu, Z.; Yang, L.; Stehman, S.V.; Czaplewski, R.L. 2000. Accuracy assessment of the U.S. Geological Survey regional land-cover mapping program:

New York and New Jersey region. Photogrammetric Engineering and Remote

Sensing. 66: 1425–1438.
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Appendix 1. Example GAP Applications

Businesses and Nongovernment Organizations

The following are some examples of applications of Gap Analysis Program (GAP)

data by the private sector:

• The Wyoming Natural Heritage Program (a private nongovernment

organization) transformed the endangered and sensitive species database

into a spatially referenced digital geographic information system using the

GAP digital base map and other GAP spatial data.

• Hughes Corp. is experimenting with the Utah and Nevada GAP digital base

maps, simulating images to aid the development of new space-based

remote-sensing devices.

• The Nature Conservancy used the Wyoming GAP data to develop a map of

ecoregions of Wyoming.

• Weyerhaeuser Corp. is using the Arkansas GAP data in managing their

lands in Arkansas.

• IBM Corp. is funding a project at the University of California-Santa

Barbara that, in part, uses GAP data in the development of visualization

software.

• The New Mexico GAP vegetation data is being used for an environmental

assessment of a proposed spaceport, a state/private venture.

• The Conservation Trust of Puerto Rico and NatureServe are using Puerto

Rico Gap data in their development of island-wide biodiversity

conservation plans.

County and City Planning

Examples of the use of GAP by local governments:

• California GAP biological data were combined with the Southern

California Association of Governments land ownership data to show which

ownerships and jurisdictions were needed for joint conservation planning

and management of a particular natural community or species, maximizing

efficiency and minimizing the potential for yet another conservation crisis.

• In California, county and city planners of several jurisdictions, wildlife

agencies, developers of the 4S Ranch property, and the state Natural

Communities Conservation Planning program used the GAP regional data,
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as well as more detailed information, to conserve 1,640 acres of habitat

within a 2,900-acre planned development.

• Day-to-day county planning operations in Piute, Grande, and Washington

counties, Utah.

• County planners in Piute County, Utah, used GAP data to optimize the

siting of a proposed sawmill for aspen with respect to the distribution of

aspen stands.

• Missoula County, Montana, used the GAP land cover map of the area as a

base map for its comprehensive long-range plan.

• Snohomish County, Washington, used the GAP land cover map in meeting

state requirements for a growth management plan.

• The City of Bainbridge Island, Washington, used GAP data to assist them

in development of a watershed planning project.

State Uses

The following are some examples of uses of GAP data by state agencies:

• The GAP database of species habitats was used by the Tennessee Wildlife

Resources Agency (TWRA) to update its book “Species in Need of

Management.”

• Images of land cover derived from GAP Landsat thematic mapper data are

used by TWRA for locating particular habitat types. Information on the

locations of these habitat types is provided by TWRA to the public for a

wide variety of public service functions, from education to cooperative

resource management.

• Early GAP data developed by TWRA were used to help identify an

extremely important area of the state with high biodiversity that was

subsequently purchased by the state for conservation.

• Preliminary findings from GAP were used by TWRA to develop three

resource management initiatives.

• The Tennessee GAP project, which is being carried out primarily by

TWRA, is the foundation of a multiagency, long-term biodiversity program

for Tennessee.

• GAP data have been used by the Tennessee Forestry Stewardship Program

to help develop a district program for nine conservation planning districts,

outlining Best Management Practices (BMPs) for biological conservation

on private lands.
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• GAP data are being used extensively by TWRA in the preparation of

project proposals to the North American Waterfowl Conservation Program.

These proposals require that biodiversity issues are addressed in specific

detail. The use of GAP data on occurrence of land cover types and

terrestrial vertebrates has made this possible.

• The Wyoming Department of Fish and Game used GAP data to assist them

in transforming the Wildlife Observation System database into a spatially

referenced geographic information system.

• The Utah Division of Wildlife Resources and the Bear River Water

Conservancy District used the Utah GAP land cover map in a resource

management assessment for mitigating conflicts between a proposed

groundwater withdrawal project and the maintenance of an elk calving area

in the Uinta Mountains.

• The Utah Division of Wildlife Resources, the Rocky Mountain Elk

Foundation, and Sheik Safari International used the Utah GAP land cover

map to identify critical elk habitat. The environmental profile of these

areas was then used to identify other similar areas for elk habitat

enhancement.

• The Utah Division of Wildlife Resources used the Utah GAP land cover

map for a rapid ecological assessment of the Echo Henefer Wildlife

Management Area.

• The Washington Department of Fish and Wildlife used GAP data to

develop a breeding bird atlas and an atlas of mammals of Washington State.

• The Washington Department of Fish and Wildlife uses GAP data to operate

an integrated landscape management program.

• The Washington Department of Fish and Wildlife uses GAP data from

eastern Washington to assist with an innovative program that brings the

forest products industry, state agency biologists, nongovernment

organizations, and tribal biologists together in the field to jointly determine

the appropriate management practices for any particular site of concern

(Timber, Fish & Wildlife Program).

• The Idaho Department of Fish and Game used GAP data to evaluate the

impact from expanded military training activities on public lands in

southern Idaho.

• The Idaho Department of Fish and Game uses GAP data for regional

planning efforts on a regular basis.
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• The Puerto Rico Department of Natural and Environmental Resources is

incorporating Puerto Rico GAP data in publicly accessible, interactive

mapping databases with land ownership and land use information.

Statewide Planning

Biodiversity planning programs or projects are now underway in Arizona, California, Colorado, Maine, Missouri, Nevada, Oregon, Puerto Rico, and Tennessee. It is

likely that similar efforts will develop in other states. In some cases, these efforts

grew out of the state GAP project; however, in most cases, the GAP data are being

used to meet a previously defined need. In all cases, GAP data are central to their

development and operations.

Federal Agency Applications

Examples of applications of GAP data by federal agencies:

• GAP data are being supplied to all military installations in the Great Basin

ecoregion for integrated management of the natural resources. These

installations constitute a very large amount of land area. Much of it is of

high value for native species.

• The Ouachita National Forest used the Arkansas GAP data to help them

develop an ecosystem management plan.

• The Wyoming GAP data were used by The National Aeronautics and Space

Agency to calibrate a model that predicts vegetation types based on climate

and soil variables.

• The potential contributions to biodiversity conservation of four options

proposed for new wilderness designation in Idaho were quantified by the

Idaho Cooperative Fish and Wildlife Research Unit in cooperation with the

Park Studies Unit.

• The potential contributions to biodiversity conservation of four options

proposed for new national park designation in Idaho were quantified by the

Idaho Cooperative Park Studies Unit.

• The U.S. Forest Service in Booneville, Arkansas, used the Arkansas GAP

data land cover maps in a 3-dimensional presentation to provide the public

with a visual representation of the region and to enhance the public’s

involvement with the national forest planning process.

• The U.S. Fish and Wildlife Service regularly uses the GAP data for

southern California for habitat evaluation and management.
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GENERAL TECHNICAL REPORT IITF-GTR-39

• The U.S. Forest Service, Bureau of Land Management, and National Park

Service are using GAP data for a wide variety of natural resource management operations in Utah. For example, the entire Utah GAP database is

directly linked with existing National Park Service databases for use by

national parks.

• The Bureau of Land Management uses the Wyoming GAP data for

managing the Buffalo Resource Area.

• The U.S. Forest Service used the Utah GAP data to help assist them in

evaluating human-induced impacts to forested lands surrounding ski resorts

in central Utah.

• The U.S. Fish and Wildlife Service in Delaware used GAP data to help

identify potential habitat for the federally endangered Delmarva fox

squirrel (Sciurus niger cinereus). These maps were displayed and served as

a catalyst for bringing together people with a stake in the issue.

• The U.S. Fish and Wildlife Service used the Indiana GAP data as part of a

biological assessment for the base closure of the Jefferson Proving Grounds

and its conversion to a National Wildlife Refuge. This 58,000-acre installation has restricted human access due to unexploded ordinance and contains

some of the highest quality natural habitat in Indiana.

• The U.S. Fish and Wildlife Service in Louisiana used GAP data to avoid

conflict over the designation of critical habitat of the federally endangered

Louisiana black bear (Ursus americanus luteolus).

• The National Oceanic and Atmospheric Administration Coastal Marine

Sanctuary in Washington State uses GAP data for an educational display.

• In Washington and New Mexico, digital land cover maps have been

distributed to all national forests.

• The U.S. Natural Resources Conservation Service in New Mexico is using

a GAP clustered imagery as a base for their land cover mapping activities.

• The Department of Defense (DoD) is funding the development of an

electronic environmental information system for the Mojave ecoregion,

which would use GAP data as a foundation or base layer of information.

The system will link 29 DoD installations to a common source of

environmental information.
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The Puerto Rico Gap Analysis Project

Appendix 2. Terrestrial Vertebrate Species of Puerto RicoThis table includes the scientific name, English and Spanish common names, species identification code (species ID), class, order, family, and taxonomicauthority for 436 terrestrial vertebrate species found in Puerto Rico. The 177 resident, endemic, or endangered terrestrial vertebrate species addressed inthe Puerto Rico Gap Analysis Project are in bold.Table 24—Terrestrial vertebrate species of Puerto RicoScientific name English common name Spanish common name Species ID Class Order Family AuthorityBirds (328 species)

Accipiter striatus venator Puerto Rican sharp-shinned Halcón de sierra ABNKC12021 Aves Falconiformes Accipitridae Wetmore, 1914

hawk

Actitis macularia Spotted sandpiper Playero coleador ABNNF04020 Aves Charadriiformes Scolopacidae Linnaeus, 1766

Agelaius xanthomus Yellow-shouldered blackbird Mariquita de Puerto Rico ABPBXB0040 Aves Passeriformes Icteridae Sclater, 1862Aix sponsa Wood duck Pato joyuyo ABNJB09010 Aves Anseriformes Anatidae Linnaeus, 1758Ajaia ajaja Roseate spoonbill Espátula rosada ABNGE05010 Aves Ciconiiformes Threskiornithidae Linnaeus, 1758Amandava amandava Red avadavat Gorrión fresa ABPCB03010 Aves Passeriformes Estrildidae Linnaeus, 1758Amazona amazonica Orange-winged parrot Cotorra alianaranjada LANRA78220 Aves Psittaciformes Psittacidae Linnaeus, 1766Amazona ochrocephala Yellow-headed parrot Cotorra cabeciamarilla ABNQA15120 Aves Psittaciformes Psittacidae Ridgway, 1887

Amazona ventralis Hispaniolan parrot Cotorra de La Española ABNQA15050 Aves Psittaciformes Psittacidae Muller, 1776Amazona viridigenalis Red-crowned parrot Cotorra coronirroja ABNQA15080 Aves Psittaciformes Psittacidae Cassin, 1853Amazona vittata Puerto Rican parrot Cotorra Puertorriqueña ABNQA15060 Aves Psittaciformes Psittacidae Boddaert, 1783Ammodramus savannarum Grasshopper sparrow Gorrión chicharra ABPBXA0020 Aves Passeriformes Emberizidae Gmelin, 1789Anas acuta Northern pintail Pato pescuecilargo ABNJB10110 Aves Anseriformes Anatidae Linnaeus, 1758Anas americana American wigeon Pato cabeciblanco ABNJB10180 Aves Anseriformes Anatidae Gmelin, 1789

Americano

Anas bahamensis White-cheeked pintail Pato quijada colorada ABNJB10100 Aves Anseriformes Anatidae Linnaeus, 1758

Anas clypeata Northern shoveler Pato cuchareta ABNJB10150 Aves Anseriformes Anatidae Linnaeus, 1758

Anas crecca Green-winged teal Pato aliverde ABNJB10010 Aves Anseriformes Anatidae Linnaeus, 1758Anas cyanoptera Cinnamon teal Pato colorado ABNJB10140 Aves Anseriformes Anatidae Vieillot, 1816Anas discors Blue-winged teal Pato zarcel ABNJB10130 Aves Anseriformes Anatidae Linnaeus, 1766Anas penelope Eurasian wigeon Pato cabeciblanco ABNJB10170 Aves Anseriformes Anatidae Linnaeus, 1758

Europeo

Anas platyrhynchos Mallard Pato cabeciverde ABNJB10060 Aves Anseriformes Anatidae Linnaeus, 1758Anas querquedula Garganey Pato carretón ABNJB10120 Aves Anseriformes Anatidae Linnaeus, 1758Anas rubripes American black duck Pato oscuro ABNJB10040 Aves Anseriformes Anatidae Brewster, 1902Anas strepera Gadwall Pato grís ABNJB10160 Aves Anseriformes Anatidae Linnaeus, 1758Anous minutus Black noddy Cervera negra ABNNM11020 Aves Charadriiformes Laridae Boie, 1844Anous stolidus Brown noddy Cervera ABNNM11010 Aves Charadriiformes Laridae Linnaeus, 1758
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GENERAL TECHNICAL REPORT IITF-GTR-39 Table 24—Terrestrial vertebrate species of Puerto Rico (continued) Scientific name English common name Spanish common name Species ID Class Order Family Authority Anthracothorax dominicus Antillean mango Zumbador dorado ABNUC11040 Aves Apodiformes Trochilidae Linnaeus, 1766 Anthracothorax viridis Green mango Zumbador verde ABNUC11050 Aves Apodiformes Trochilidae Audebert and Viellot, 1801 Apus melba Alpine swift Vencejo real ABNUA05030 Aves Apodiformes Apodidae Linnaeus, 1758 Aramus guarauna Limpkin Carrao ABNMJ01010 Aves Gruiformes Aramidae Linnaeus, 1766 Aratinga canicularis Orange-fronted parakeet Periquito ABNQA06070 Aves Psittaciformes Psittacidae Linnaeus, 1758 frentianaranjado Aratinga chloroptera Hispaniolan parakeet Periquito de la Española ABNQA06040 Aves Psittaciformes Psittacidae Souance, 1856 Aratinga pertinax Brown-throated parakeet Periquito de curazao ABNQA06080 Aves Psittaciformes Psittacidae Linnaeus, 1758 Archilochus colubris Ruby-throated hummingbird Zumbadorcito gorgirrojo ABNUC45010 Aves Apodiformes Trochilidae Linnaeus, 1758 Ardea alba Great egret Garzón blanco ABNGA04040 Aves Ciconiiformes Ardeidae Linnaeus, 1758

Ardea herodias Great blue heron Garzón cenizo ABNGA04010 Aves Ciconiiformes Ardeidae Linnaeus, 1758Arenaria interpres Ruddy turnstone Playero turco ABNNF09010 Aves Charadriiformes Scolopacidae Linnaeus, 1758Asio flammeus Short-eared owl Múcaro real ABNSB13040 Aves Strigiformes Strigidae Pontoppidan,

1763

Aythya affinis Lesser scaup Pato pechiblanco ABNJB11070 Aves Anseriformes Anatidae Eyton, 1838Aythya collaris Ring-necked duck Pato acollarado ABNJB11040 Aves Anseriformes Anatidae Donovan, 1809Aythya marila Greater scaup Pato boludo ABNJB11060 Aves Anseriformes Anatidae Linnaeus, 1761

Aythya valisineria Canvasback Pato lomiblanco ABNJB11020 Aves Anseriformes Anatidae Wilson, 1814

Bartramia longicauda Upland sandpiper Ganga ABNNF06010 Aves Charadriiformes Scolopacidae Bechstein, 1812Bombycilla cedrorum Cedar waxwing Ampelis Americano ABPBN01020 Aves Passeriformes Bombycillidae Vieillot, 1808

Botaurus lentiginosus American bittern Yaboa Americana ABNGA01020 Aves Ciconiiformes Ardeidae Rackett, 1813Branta bernicla Brant goose Ganso carinegra ABNJB05010 Aves Anseriformes Anatidae Linnaeus, 1758Branta canadensis Canada goose Ganso canadiense ABNJB05030 Aves Anseriformes Anatidae Linnaeus, 1758Brotogeris versicolurus White-winged parakeet Pequito aliamarillo ABNQA11020 Aves Psittaciformes Psittacidae Muller, 1776Bubulcus ibis Cattle egret Garza del ganado ABNGA07010 Aves Ciconiiformes Ardeidae Linnaeus, 1758Bucephala albeola Bufflehead Pato pínto ABNJB18030 Aves Anseriformes Anatidae Linnaeus, 1758Buteo jamaicensis Red-tailed hawk Guaraguao colirrojo ABNKC19110 Aves Falconiformes Accipitridae Gmelin, 1788Buteo platypterus Puerto Rican broad- Guaraguao de bosque ABNKC19050 Aves Falconiformes Accipitridae Vieillot, 1823

brunnescens winged hawk

Buteo ridgwayi Ridgway’s hawk Guaraguaito de la Española ABNKC19040 Aves Falconiformes Accipitridae Cory, 1883Buteogallus anthracinus Common black-hawk Guaraguano negro ABNKC15010 Aves Falconiformes Accipitridae Deppe, 1830

Butorides virescens Green heron Martinete verde ABNGA08010 Aves Ciconiiformes Ardeidae Linnaeus, 1758Cairina moschata Muscovy duck Pato criollo ABNJB07010 Aves Anseriformes Anatidae Linnaeus, 1758Calidris alba Sanderling Playero arenero ABNNF11030 Aves Charadriiformes Scolopacidae Pallas, 1764

Calidris alpina Dunlin Playero espalda colorada ABNNF11170 Aves Charadriiformes Scolopacidae Linnaeus, 1758Calidris canutus Red knot Playero gordo ABNNF11020 Aves Charadriiformes Scolopacidae Linnaeus, 1758
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Table 24—Terrestrial vertebrate species of Puerto Rico (continued)Scientific name English common name Spanish common name Species ID Class Order Family Authority

Calidris ferruginea Curlew sandpiper Playero zarapatin ABNNF11180 Aves Charadriiformes Scolopacidae Pontoppidan,

1763

Calidris fuscicollis White-rumped sandpiper Playero de rabadilla blanca ABNNF11110 Aves Charadriiformes Scolopacidae Vieillot, 1819

Calidris himantopus Stilt sandpiper Playero patilargo ABNNF11190 Aves Charadriiformes Scolopacidae Bonaparte,

1826

Calidris mauri Western sandpiper Playero occidental ABNNF11050 Aves Charadriiformes Scolopacidae Cabanis, 1857Calidris melanotos Pectoral sandpiper Playero pectoral ABNNF11130 Aves Charadriiformes Scolopacidae Vieillot, 1819

Calidris minutilla Least sandpiper Playerito menudillo ABNNF11100 Aves Charadriiformes Scolopacidae Vieillot, 1819Calidris pusilla Semipalmated sandpiper Playero gracioso ABNNF11040 Aves Charadriiformes Scolopacidae Linnaeus, 1766Calonectris diomedea Cory’s shearwater Pampero ceniciento ABNDB06020 Aves Procellariiformes Procellariidae Scopoli, 1769

Caprimulgus carolinensis Chuck-will’s-widow Guabairo de la Carolina ABNTA07010 Aves Caprimulgiformes Caprimulgidae Gmelin, 1789Caprimulgus cayennensis White-tailed nightjar Guabairo coliblanco ABNTA07100 Aves Caprimulgiformes Caprimulgidae J.F. Gmelin,

1789

Caprimulgus noctitherus Puerto Rican nightjar Guabairo de Puerto Rico ABNTA07080 Aves Caprimulgiformes Caprimulgidae Wetmore, 1919Carduelis cucullata Red siskin Cardenalito ABPBY06070 Aves Passeriformes Fringillidae Swainson, 1820Catharacta skua Great skua Págalo grande ABNNM01040 Aves Charadriiformes Laridae Brunnich, 1764Cathartes aura Turkey vulture Aura tiñosa ABNKA02010 Aves Ciconiiformes Cathartidae Linnaeus, 1758Catharus bicknelli Bicknell’s thrush Zorzal de Bicknell ABPBJ18120 Aves Passeriformes Turdidae Ridgway, 1882

Catharus minimus Gray-cheeked thrush Zorzal carigris ABPBJ18090 Aves Passeriformes Turdidae Lafresnaye, 1848

Catoptrophorus Willet Playero aliblanco ABNNF02010 Aves Charadriiformes Scolopacidae Gmelin, 1789

semipalmatus

Ceryle alcyon Belted kingfisher Martín pescador norteño ABNXD01020 Aves Coraciiformes Alcedinidae Linnaeus, 1758Ceryle torquata Ringed kingfisher Martín pescador anillado ABNXD01010 Aves Coraciiformes Alcedinidae Linnaeus, 1776Chaetura brachyura Short-tailed swift Vencejo pequeño ABNUA03040 Aves Apodiformes Apodidae Jardine, 1846Charadrius alexandrinus Snowy plover Chorlo blanco ABNNB03030 Aves Charadriiformes Charadriidae Linnaeus, 1758Charadrius melodus Piping plover Chorlo melódico ABNNB03070 Aves Charadriiformes Charadriidae Ord, 1824Charadrius semipalmatus Semipalmated plover Chorlo acollarado ABNNB03060 Aves Charadriiformes Charadriidae Bonaparte, 1825Charadrius vociferus Killdeer Chorlo sabanero ABNNB03090 Aves Charadriiformes Charadriidae Linnaeus, 1758Charadrius wilsonia Wilson’s plover Chorlo marítimo ABNNB03040 Aves Charadriiformes Charadriidae Ord, 1814Chen caerulescens Snow goose Ganso blanco ABNJB04010 Aves Anseriformes Anatidae Linnaeus, 1758Chlidonias niger Black tern Gharrán ceniza ABNNM10020 Aves Charadriiformes Laridae Linnaeus, 1758Chlorostilbon maugaeus Puerto Rican emerald Zumbadorcito de ABNUC18050 Aves Apodiformes Trochilidae Audebert and

Puerto Rico Vieillot, 1801

Chordeiles gundlachii Antillean nighthawk Querequeqúe Antillano ABNTA02030 Aves Caprimulgiformes Caprimulgidae Lawrence, 1857Chordeiles minor Common nighthawk Querequeqúe menor ABNTA02020 Aves Caprimulgiformes Caprimulgidae Forster, 1771Circus cyaneus Northern harrier Gavilán de ciénaga ABNKC11010 Aves Falconiformes Accipitridae Linnaeus, 1758Coccyzus americanus Yellow-billed cuckoo Pájaro bobo pechiblanco ABNRB02020 Aves Cuculiformes Cuculidae Linnaeus, 1758
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GENERAL TECHNICAL REPORT IITF-GTR-39 Table 24—Terrestrial vertebrate species of Puerto Rico (continued) Scientific name English common name Spanish common name Species ID Class Order Family Authority Coccyzus erythropthalmus Black-billed cuckoo Pájaro bobo piquinegro ABNRB02010 Aves Cuculiformes Cuculidae Wilson, 1811 Coccyzus minor Mangrove cuckoo Pájaro bobo menor ABNRB02030 Aves Cuculiformes Cuculidae Gmelin, 1788 Coereba flaveola Bananaquit Reinita común ABPBX28010 Aves Passeriformes Emberizidae Linnaeus, 1758 Colinus virginianus Northern bobwhite Codorniz de Virginia ABNLC21020 Aves Galliformes Odontophoridae Linnaeus, 1758 Columba livia Rock dove Paloma doméstica ABNPB01010 Aves Columbiformes Columbidae Gmelin, 1789 Columbina passerina Common ground-dove Rolita ABNPB06020 Aves Columbiformes Columbidae Linnaeus, 1758 Contopus hispaniolensis Hispaniolan pewee Bobito de la Española LAP0946120 Aves Passeriformes Tyrannidae Bryant, 1867 Contopus latirostris Lesser Antillean pewee Bobito Antíllano Menor ABPAE32090 Aves Passeriformes Tyrannidae Verreaux, 1866 Corvus leucognaphalus White-necked crow Cuervo pescueciblanco ABPAV10050 Aves Passeriformes Corvidae Daudin, 1800 Crotophaga ani Smooth-billed ani Garrapatero ABNRB11020 Aves Cuculiformes Cuculidae Linnaeus, 1758 Cygnus columbianus Tundra swan Cisne de tundra ABNJB02010 Aves Anseriformes Anatidae Ord, 1815

Cypseloides niger Black swift Vencejo negro ABNUA01010 Aves Apodiformes Apodidae Gmelin, 1789

Dendrocygna arborea West Indian whistling-duck Chiriría caribeña ABNJB01020 Aves Anseriformes Anatidae Linnaeus, 1758Dendrocygna autumnalis Black-bellied whistling-duck Chiriría pinta ABNJB01040 Aves Anseriformes Anatidae Linnaeus, 1758Dendrocygna bicolor Fulvous whistling-duck Chiriría bicolor ABNJB01010 Aves Anseriformes Anatidae Vieillot, 1816Dendroica adelaidae Adelaide’s warbler Reinita mariposera ABPBX03150 Aves Passeriformes Parulidae Baird, 1865

Dendroica angelae Elfin-woods warbler Reinita de bosque enaño ABPBX03270 Aves Passeriformes Parulidae Kepler and

Parkes, 1972

Dendroica caerulescens Black-throated blue warbler Reinita azul ABPBX03050 Aves Passeriformes Parulidae Gmelin, 1789Dendroica castanea Bay-breasted warbler Reinita castaña ABPBX03220 Aves Passeriformes Parulidae Wilson, 1810

Dendroica cerulea Cerulean warbler Reinita cerúlea ABPBX03240 Aves Passeriformes Parulidae Wilson, 1810Dendroica coronata Yellow-rumped warbler Reinita coronada ABPBX03060 Aves Passeriformes Parulidae Linnaeus, 1766Dendroica discolor Prairie warbler Reinita galana ABPBX03190 Aves Passeriformes Parulidae Vieillot, 1809

Dendroica dominica Yellow-throated warbler Reinita gargantiamarilla ABPBX03130 Aves Passeriformes Parulidae Linnaeus, 1776Dendroica fusca Blackburnian warbler Reinita de fuego ABPBX03120 Aves Passeriformes Parulidae Muller, 1776Dendroica magnolia Magnolia warbler Reinita manchada ABPBX03030 Aves Passeriformes Parulidae Wilson, 1811

Dendroica palmarum Palm warbler Reinita palmera ABPBX03210 Aves Passeriformes Parulidae Gmelin, 1789Dendroica pensylvanica Chestnut-sided warbler Reinita flanquicastaña ABPBX03020 Aves Passeriformes Parulidae Linnaeus, 1766Dendroica petechia Yellow warbler Reinita amarilla ABPBX03010 Aves Passeriformes Parulidae Linnaeus, 1766Dendroica pinus Pine warbler Reinita de pinos ABPBX03170 Aves Passeriformes Parulidae Wilson, 1811

Dendroica striata Blackpoll warbler Reinita rayada ABPBX03230 Aves Passeriformes Parulidae Forster, 1772Dendroica tigrina Cape May warbler Reinita tigre ABPBX03040 Aves Passeriformes Parulidae Gmelin, 1789Dendroica virens Black-throated green warbler Reinita verdosa ABPBX03100 Aves Passeriformes Parulidae Gmelin, 1789

Dolichonyx oryzivorus Bobolink Chambergo ABPBXA9010 Aves Passeriformes Icteridae Linnaeus, 1758Dumetella carolinensis Gray catbird Mauyador gris ABPBK01010 Aves Passeriformes Mimidae Linnaeus, 1766Egretta caerulea Little blue heron Garza azul ABNGA06040 Aves Ciconiiformes Ardeidae Linnaeus, 1758Egretta garzetta Little egret Garza común ABNGA06020 Aves Ciconiiformes Ardeidae Linnaeus, 1766
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Table 24—Terrestrial vertebrate species of Puerto Rico (continued)Scientific name English common name Spanish common name Species ID Class Order Family Authority

Egretta gularis Western reef heron Garza dimorfa ABNGA06070 Aves Ciconiiformes Ardeidae Bosc, 1792Egretta rufescens Reddish egret Garza rojiza ABNGA06060 Aves Ciconiiformes Ardeidae Gmelin, 1789Egretta thula Snowy egret Garza blanca ABNGA06030 Aves Ciconiiformes Ardeidae Molina, 1782Egretta tricolor Tricolored heron Garza pechiblanca ABNGA06050 Aves Ciconiiformes Ardeidae Muller, 1776Elaenia martinica Caribbean elaenia Juí blanco ABPAE09010 Aves Passeriformes Tyrannidae Linnaeus, 1766

Estrilda melpoda Orange-cheeked waxbill Veterano ABPCB02020 Aves Passeriformes Estrildidae Vieillot, 1817Estrilda troglodytes Black-rumped waxbill Veterano orejicolorado ABPCB02030 Aves Passeriformes Estrildidae Lichtenstein, 1823

Eudocimus albus White ibis Ibis blanco ABNGE01010 Aves Ciconiiformes Threskiornithidae Linnaeus, 1758Eulampis holosericeus Green-throated carib Zumbador de pecho azul ABNUC12020 Aves Apodiformes Trochilidae Linnaeus, 1758

Euphonia musica Antillean Euphonia Jilguero ABPBX35060 Aves Passeriformes Emberizidae J. F. Gmelin, 1789Euplectes afer Yellow-crowned bishop Napoleon tejedor ABPCA02020 Aves Passeriformes Ploceidae J.F. Gmelin, 1789Euplectes franciscanus Orange bishop Obispo colorado ABPCA02030 Aves Passeriformes Ploceidae Isert, 1789Falco columbarius Merlin Halcón migratorio ABNKD06030 Aves Falconiformes Falconidae Linnaeus, 1758Falco peregrinus anatum American peregrine falcon Halcón peregrino ABNKD06071 Aves Falconiformes Falconidae 

Falco sparverius American kestrel Halcón común ABNKD06020 Aves Falconiformes Falconidae Linnaeus, 1758Fregata magnificens Magnificent frigatebird Fragata magnifica, tijereta ABNFF01010 Aves Pelecaniformes Fregatidae Mathews, 1914

Fulica americana American coot Gallinazo Americano ABNME14020 Aves Gruiformes Rallidae Gmelin, 1789Fulica caribaea Caribbean coot Gallinazo Caribeño ABNME14030 Aves Gruiformes Rallidae Ridgway, 1884Fulmarus glacialis Northern fulmar Fulmar boreal ABNDB01010 Aves Procellariiformes Procellariidae Linnaeus, 1761Gallinago gallinago Common snipe Becasina común ABNNF18010 Aves Charadriiformes Scolopacidae Linnaeus, 1758Gallinula chloropus Common moorhen Gallareta común ABNME13010 Aves Gruiformes Rallidae Linnaeus, 1758Gallus gallus Red junglefowl Gallina, gallo ABNLC06010 Aves Galliformes Phasianidae Linnaeus, 1758Geothlypis trichas Common yellowthroat Reinita pica tierra ABPBX12010 Aves Passeriformes Parulidae Linnaeus, 1766Geotrygon chrysia Key West quail-dove Paloma perdíz áurea ABNPB09020 Aves Columbiformes Columbidae Bonaparte, 1855Geotrygon montana Ruddy quail-dove Paloma perdíz rojiza ABNPB09110 Aves Columbiformes Columbidae Linnaeus, 1758Geotrygon mystacea Bridled quail-dove Paloma perdíz de ABNPB09030 Aves Columbiformes Columbidae Temminck, 1811

Martinica

Gracula religiosa Hill myna Maina de colinas ABPBT03010 Aves Passeriformes Columbidae Linnaeus, 1758Haematopus palliatus American oystercatcher Ostrero Americano ABNNC01010 Aves Charadriiformes Haematopodidae Temminck, 1820Helmitheros vermivorus Worm-eating warbler Reinita gusanera ABPBX08010 Aves Passeriformes Parulidae Gmelin, 1789Himantopus mexicanus Black-necked stilt Viuda Mexicana ABNND01010 Aves Charadriiformes Recurvirostridae Muller, 1776Hirundo rustica Barn swallow Golondrina horquilla ABPAU09030 Aves Passeriformes Hirundinidae Linnaeus, 1758Hylocichla mustelina Wood thrush Zorzal pecho manchado ABPBJ19010 Aves Passeriformes Turdidae Gmelin, 1789Icterus dominicensis Greater Antillean oriole Calandria ABPBXB9010 Aves Passeriformes Icteridae Linnaeus, 1766

Icterus galbula Baltimore oriole Calandria del norte ABPBXB9190 Aves Passeriformes Icteridae Linnaeus, 1758Icterus icterus Venezuelan troupial Turpial ABPBXB9120 Aves Passeriformes Icteridae Linnaeus, 1766
134

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 24—Terrestrial vertebrate species of Puerto Rico (continued) Scientific name English common name Spanish common name Species ID Class Order Family Authority Ixobrychus exilis Least bittern Martinetito ABNGA02010 Aves Ciconiiformes Ardeidae Gmelin, 1789 Jacana spinosa Northern jacana Jacana Centroamericana ABNNE01010 Aves Charadriiformes Jacanidae Linnaeus, 1758 Junco hyemalis Dark-eyed junco Junco ojioscuro ABPBXA5020 Aves Passeriformes Emberizidae Linnaeus, 1758 Larus argentatus Herring gull Gaviota argéntea ABNNM03120 Aves Charadriiformes Laridae Pontoppidan, 1763 Larus atricilla Laughing gull Gaviota cabecinegra ABNNM03010 Aves Charadriiformes Laridae Linnaeus, 1758 Larus delawarensis Ring-billed gull Gaviota piquianillada ABNNM03100 Aves Charadriiformes Laridae Ord, 1815 Larus fuscus Lesser Black-backed gull Gaviota sombriá ABNNM03150 Aves Charadriiformes Laridae Linnaeus, 1758 Larus marinus Great Black-backed gull Gaviota marina ABNNM03210 Aves Charadriiformes Laridae Linnaeus, 1758 Larus philadelphia Bonaparte’s gull Gaviota de Bonaparte ABNNM03050 Aves Charadriiformes Laridae Ord, 1815 Larus pipixcan Franklin’s gull Gaviota de Franklin ABNNM03020 Aves Charadriiformes Laridae Wagler, 1831 Larus ridibundus Black-headed gull Gaviota cabecinegra ABNNM03040 Aves Charadriiformes Laridae Linnaeus, 1766

forastera

Laterallus jamaicensis Black rail Gallito negro ABNME03040 Aves Gruiformes Rallidae Gmelin, 1789

Limnodromus griseus Short-billed dowitcher Agujeta pico corto ABNNF16010 Aves Charadriiformes Scolopacidae Gmelin, 1789

Limnodromus scolopaceus Long-billed dowitcher Agujeta pico largo ABNNF16020 Aves Charadriiformes Scolopacidae Say, 1823Limnothlypis swainsonii Swainson’s warbler Reinita de Swainson ABPBX09010 Aves Passeriformes Parulidae Audubon, 1834Limosa fedoa Marbled godwit Barga canela ABNNF08040 Aves Charadriiformes Scolopacidae Linnaeus, 1758

Limosa haemastica Hudsonian godwit Barga aliblanca ABNNF08020 Aves Charadriiformes Scolopacidae Linnaeus, 1758

Lonchura cucullata Bronze mannikin Diablito L000000192 Aves Passeriformes Estrildidae Swainson, 1837

Lonchura malabarica Warbling silverbill Gorrión picoplata ABPCB04010 Aves Passeriformes Estrildidae Linnaeus, 1758Lonchura malacca Chestnut mannikin Monjita tricolor ABPCB04040 Aves Passeriformes Estrildidae Linnaeus, 1766Lonchura punctulata Nutmeg mannikin Gorrión canela ABPCB04030 Aves Passeriformes Estrildidae Linnaeus, 1758Lophodytes cucullatus Hooded merganser Mergansa encapuchada ABNJB20010 Aves Anseriformes Anatidae Linnaeus, 1758Loxigilla noctis Lesser Antillean bullfinch Antillano menor ABPBX82030 Aves Passeriformes Emberizidae Linnaeus, 1766Loxigilla portoricensis Puerto Rican bullfinch Gallito ABPBX82010 Aves Passeriformes Emberizidae Daudin, 1800

Margarops fuscatus Pearly-eyed thrasher Zorzal pardo ABPBK09020 Aves Passeriformes Mimidae Vieillot, 1808

Megascops nudipes Puerto Rican screech-owl Múcaro común ABNSB01120 Aves Strigiformes Strigidae Daudin, 1800

Melanerpes portoricensis Puerto Rican woodpecker Carpintero ABNYF04030 Aves Piciformes Picidae Daudin, 1803Mellisuga minima Vervain hummingbird Zumbadorcito menor ABNUC46010 Aves Apodiformes Trochilidae Linnaeus, 1758Melopsittacus undulatus Budgerigar Periquito Australiano ABNQA01010 Aves Psittaciformes Psittacidae Shaw, 1805Melospiza lincolnii Lincoln’s sparrow Gorrion de Linclon ABPBXA3020 Aves Passeriformes Emberizidae Audubon, 1834Mergus serrator Red-breasted merganser Mergansa de pico largo ABNJB21020 Aves Anseriformes Anatidae Linnaeus, 1758

Mimus polyglottos Northern mockingbird Ruisenor ABPBK03010 Aves Passeriformes Mimidae Linnaeus, 1758Mniotilta varia Black-and-white warbler Reinita trepadora ABPBX05010 Aves Passeriformes Parulidae Linnaeus, 1766Molothrus bonariensis Shiny cowbird Tordo lustrosso ABPBXB7010 Aves Passeriformes Icteridae Gmelin, 1789Myiarchus antillarum Puerto Rican flycatcher Juí de Puerto Rico ABPAE43130 Aves Passeriformes Tyrannidae Bryant, 1866Myiarchus crinitus Great crested flycatcher Juí crestado ABPAE43070 Aves Passeriformes Tyrannidae Linnaeus, 1758Myiarchus stolidus Stolid flycatcher Manuelito ABPAE43120 Aves Passeriformes Tyrannidae Gosse, 1847
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The Puerto Rico Gap Analysis Project

Table 24—Terrestrial vertebrate species of Puerto Rico (continued)Scientific name English common name Spanish common name Species ID Class Order Family Authority

Myiopsitta monachus Monk parakeet Perico monje ABNQA04010 Aves Psittaciformes Psittacidea Boddaert, 1783Nandayus nenday Black-hooded Parakeet Periquito nanday ABNQA15190 Aves Psittaciformes Psittacidea Vieillot, 1823Nesospingus speculiferus Puerto Rican tanager Llorosa de Puerto Rico ABPBX52010 Aves Passeriformes Emberizidae Lawrene, 1875

Nomonyx dominicus Masked duck Pato Dominico ABNJB22020 Aves Anseriformes Anatidae Linnaeus, 1758Numenius americanus Long-billed curlew Playero Americano ABNNF07070 Aves Charadriiformes Scolopacidae Bechstein, 1812Numenius borealis Eskimo curlew Playero picocorvo ABNNF07010 Aves Charadriiformes Scolopacidae Forster, 1772

esquimal

Numenius phaeopus Whimbrel Playero trinador ABNNF07020 Aves Charadriiformes Scolopacidae Linnaeus, 1758Numida meleagris Helmeted guineafowl Guinea torcaz ABNLC25010 Aves Galliformes Phasianidae Linnaeus, 1758Nyctanassa violacea Yellow-crowned night-heron Yaboa común ABNGA13010 Aves Ciconiiformes Ardeidae Linnaeus, 1758Nyctibius jamaicensis Northern potoo Nictibio Jamaicano LANUC01060 Aves Caprimulgifores Nyctibiidae J.F. Gmelin, 1789

Nycticorax nycticorax Black-crowned night-heron Yaboa real ABNGA11010 Aves Ciconiiformes Ardeidae Linnaeus, 1758Oceanites oceanicus Wilson’s storm-petrel Petrel de Wilson ABNDC01010 Aves Procellariiformes Hydrobatidae Kuhl, 1820Oceanodroma leucorhoa Leach’s storm-petrel Pamperito rabo ABNDC04020 Aves Procellariiformes Hydrobatidae Vieillot, 1818

horquillado

Oenanthe oenanthe Northern wheatear Zorzal gris ABPBJ14010 Aves Passeriformes Turdidae Linnaeus, 1758

Oporornis agilis Connecticut warbler Reinita de Connecticut ABPBX11020 Aves Passeriformes Parulidae Wilson, 1812

Oporornis formosus Kentucky warbler Reinita de Kentucky ABPBX11010 Aves Passeriformes Parulidae Wilson, 1811Oporornis philadelphia Mourning warbler Reinita triste ABPBX11030 Aves Passeriformes Parulidae Wilson, 1810

Orthorhyncus cristatus Antillean crested hummingbird Zumbador crestado ABNUC13010 Aves Apodiformes Trochilidae Linnaeus, 1758Oxyura jamaicensis Ruddy duck Pato chorizo ABNJB22010 Aves Anseriformes Anatidae Gmelin, 1789Padda oryzivora Java sparrow Gorrión arrocero ABPCB05010 Aves Passeriformes Estrildidae Linnaeus, 1758Pandion haliaetus Osprey Aguila pescadora ABNKC01010 Aves Falconiformes Accipitridae Linnaeus, 1758Parula americana Northern parula Reinita pechidorada ABPBX02010 Aves Passeriformes Parulidae Linnaeus, 1758Passer domesticus House sparrow Gorrión doméstico ABPBZ01010 Aves Passeriformes Passeridae Linnaeus, 1758Passerina caerulea Blue grosbeak Picogrueso azul ABPBX64010 Aves Passeriformes Cardinalidae Linnaeus, 1758Passerina ciris Painted bunting Mariposa ABPBX64060 Aves Passeriformes Cardinalidae Linnaeus, 1758Passerina cyanea Indigo bunting Gorrión azul ABPBX64030 Aves Passeriformes Cardinalidae Linnaeus, 1766Patagioenas inornata Puerto Rican plain pigeon Paloma sabanera ABNPB01071 Aves Columbiformes Columbidae Vigors, 1827

Patagioenas leucocephala White-crowned pigeon Paloma cabeciblanca ABNPB01050 Aves Columbiformes Columbidae Linnaeus, 1758Patagioenas squamosa Scaly-naped pigeon Paloma turca ABNPB01040 Aves Columbiformes Columbidae Bonnaterre, 1792Pelecanus erythrorhynchos American White pelican Pelícano blanco ABNFC01010 Aves Pelecaniformes Pelecanidae Gmelin, 1789Pelecanus occidentalis Brown pelican Pelícano pardo ABNFC01020 Aves Pelecaniformes Pelecanidae Linnaeus, 1766

Petrochelidon fulva Cave swallow Golondrina de cuevas ABPAU09020 Aves Passeriformes Hirundinidae Vieillot, 1808

Petrochelidon pyrrhonota Cliff swallow Golondrina de Peñasco ABPAU09010 Aves Passeriformes Hirundinidae Vieillot, 1817

Phaethon aethereus Red-billed tropicbird Rabijunco piquirrojo ABNFA01020 Aves Pelecaniformes Phaethontidae Linnaeus, 1758Phaethon lepturus White-tailed tropicbird Rabijunco coliblanco ABNFA01010 Aves Pelecaniformes Phaethontidae Daudin, 1802
136

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 24—Terrestrial vertebrate species of Puerto Rico (continued) Scientific name English common name Spanish common name Species ID Class Order Family Authority Phalacrocorax auritus Double-crested cormorant Cormorán crestado ABNFD01020 Aves Pelecaniformes Phalacrocoracidae Lesson, 1831 Phalacrocorax brasilianus Neotropic cormorant Cormorán neotropical ABNFD01030 Aves Pelecaniformes Phalacrocoracidae Gmelin, 1789 Phalaropus lobatus Red-necked phalarope Falaropo picofíno ABNNF20020 Aves Charadriiformes Scolopacidae Linnaeus, 1758 Phalaropus tricolor Wilson’s phalarope Falaropo tricolor ABNNF20010 Aves Charadriiformes Scolopacidae Vieillot, 1819 Pheucticus ludovicianus Rose-breasted grosbeak Piquegrueso pechirrojo ABPBX61030 Aves Passeriformes Cardinalidae Linnaeus, 1766 Philomachus pugnax Ruff Playero combatiente ABNNF15010 Aves Charadriiformes Scolopacidae Linnaeus, 1758 Phoenicopterus ruber Greater flamingo Flamenco ABNHA01010 Aves Phoenicopteriformes Phoenicopteridae Linnaeus, 1758 Picoides villosus Hairy woodpecker Carpintero velloso ABNYF07040 Aves Piciformes Picidae Linnaeus, 1766 Piranga olivacea Scarlet tanager Tangara escarlata ABPBX45040 Aves Passeriformes Thraupidae Gmelin, 1789 Plegadis falcinellus Glossy ibis Ibis lustroso ABNGE02010 Aves Ciconiiformes Threskiornithidae Linnaeus, 1766 Pluvialis dominica American golden-plover Chorlo dorado ABNNB02030 Aves Charadriiformes Charadriidae Muller, 1776

Pluvialis squatarola Black-bellied plover Chorlo cabezón ABNNB02010 Aves Charadriiformes Charadriidae Linnaeus, 1758Podilymbus podiceps Pied-billed grebe Zaramago ABNCA02010 Aves Podicipediformes Podicipedidae Linnaeus, 1758Porphyrio martinica Purple gallinule Gallareta azul ABNME12010 Aves Gruiformes Rallidae Linnaeus, 1766

Porzana carolina Sora Gallito sora ABNME08020 Aves Gruiformes Rallidae Linnaeus, 1758

Porzana flaviventer Yellow-breasted crake Gallito amarillo ABNME08030 Aves Gruiformes Rallidae Boddaert, 1783Progne dominicensis Caribbean martin Golondrina de iglesias ABPAU01030 Aves Passeriformes Hirundinidae J.F. Gmelin,

1789

Progne subis Purple martin Golondrina azul Americana ABPAU01010 Aves Passeriformes Hirundinidae Linnaeus, 1758Protonotaria citrea Prothonotary warbler Reinita protonotaria ABPBX07010 Aves Passeriformes Parulidae Boddaert, 1783Pterodroma arminjoniana Herald petrel Petrel de la Trindade ABNDB03080 Aves Ciconiiformes Procellariidae Giglioli and

Salvadori, 1869

Pterodroma hasitata Black-capped petrel Petrel Antíllano ABNDB03010 Aves Procellariiformes Procellariidae Kuhl, 1820

Puffinus assimilis Little shearwater Pampero chico ABNDB07120 Aves Ciconiiformes Procellaridae Gould, 1838Puffinus gravis Greater shearwater Pampero capirotado ABNDB07030 Aves Procellariiformes Procellariidae O’Reilly, 1818

Puffinus griseus Sooty shearwater Pardela gris ABNDB07060 Aves Procellariiformes Procellariidae J.F. Gmelin, 1789

Puffinus iherminieri Audubon’s shearwater Pampero de Audubon ABNDB07130 Aves Procellariiformes Procellariidae Lesson, 1839Puffinus puffinus Manx shearwater Pampero pichoneta ABNDB07090 Aves Procellariiformes Procellariidae Brunnich, 1764Quiscalus niger Greater Antillean grackle Mozambique ABPBXB6030 Aves Passeriformes Icteridae Boddaert, 1783Rallus limicola Virginia rail Pollo de Virginia ABNME05030 Aves Gruiformes Rallidae Vieillot, 1819

Rallus longirostris Clapper rail Pollo de mangle ABNME05010 Aves Gruiformes Rallidae Boddaert, 1783Recurvirostra americana American avocet Avoceta Americana ABNND02010 Aves Charadriiformes Recurvirostridae Gmelin, 1789Riparia riparia Bank swallow Golondrina parda ABPAU08010 Aves Passeriformes Hirundinidae Linnaeus, 1758

Rynchops niger Black skimmer Rayador Americano ABNNM14010 Aves Charadriiformes Laridae Linnaeus, 1758Saurothera vieilloti Puerto Rican lizard-cuckoo Pájaro bobo mayor ABNRB03020 Aves Cuculiformes Cuculidae Bonaparte, 1850Seiurus aurocapillus Ovenbird Pizpita dorada ABPBX10010 Aves Passeriformes Parulidae Linnaeus, 1766Seiurus motacilla Louisiana waterthrush Pizpita de río ABPBX10030 Aves Passeriformes Parulidae Vieillot, 1809Seiurus noveboracensis Northern waterthrush Pizpita de mangle ABPBX10020 Aves Passeriformes Parulidae Gmelin, 1789
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The Puerto Rico Gap Analysis Project

Table 24—Terrestrial vertebrate species of Puerto Rico (continued)Scientific name English common name Spanish common name Species ID Class Order Family Authority

Serinus mozambicus Yellow-fronted canary Canario cantador ABPBY07010 Aves Passeriformes Fringillidae Muller, 1776Setophaga ruticilla American redstart Reinita candelita ABPBX06010 Aves Passeriformes Parulidae Linnaeus, 1758Sicalis flaveola Saffron finch Pinzón azafrán ABPBX89010 Aves Passeriformes Emberizidae Linnaeus, 1766Sphyrapicus varius Yellow-bellied sapsucker Carpintero de paso ABNYF05010 Aves Piciformes Picidae Linnaeus, 1766Spindalis portoricensis Puerto Rican spindalis Reina mora L000000251 Aves Passeriformes Thraupidae Bryant,H, 1866Spiza americana Dickcissel Sabanero Americano ABPBX65010 Aves Passeriformes Cardinalidae Gmelin, 1789Stercorarius pomarinus Pomarine jaeger Págalo pomarino ABNNM01010 Aves Charadriiformes Laridae Temminck, 1815Sterna anaethetus Bridled tern Gaviota monja ABNNM08140 Aves Charadriiformes Laridae Scopoli, 1786Sterna antillarum Least tern Gaviota chica ABNNM08100 Aves Charadriiformes Laridae Lesson, 1847Sterna caspia Caspian tern Gaviota de Caspia ABNNM08020 Aves Charadriiformes Laridae Pallas, 1770Sterna dougallii Roseate tern Palometa ABNNM08060 Aves Charadriiformes Laridae Montagu, 1813Sterna forsteri Forster’s tern Gaviota de Forster ABNNM08090 Aves Charadriiformes Laridae Nuttall, 1834Sterna fuscata Sooty tern Gaviota oscura ABNNM08150 Aves Charadriiformes Laridae Linnaeus, 1766Sterna hirundo Common tern Gaviota común ABNNM08070 Aves Charadriiformes Laridae Linnaeus, 1758Sterna maxima Royal tern Gaviota real ABNNM08030 Aves Charadriiformes Laridae Boddaert, 1783Sterna nilotica Gull-billed tern Gaviota piquigorda ABNNM08010 Aves Charadriiformes Laridae Gmelin, 1789

Sterna paradisaea Arctic tern Gaviota del Artíco ABNNM08080 Aves Charadriiformes Laridae Pontoppidan, 1763Sterna sandvicensis Sandwich tern Gaviota piquiaguda ABNNM08050 Aves Charadriiformes Laridae Latham, 1787

Streptopelia decaocto Eurasian collared-dove Tórtola turca ABNPB02030 Aves Columbiformes Columbidae Frivaldszky, 1838

Streptopelia risoria Ringed turtle-dove Tórtola ABNPB02010 Aves Columbiformes Columbidae Linnaeus, 1758Streptoprocne zonaris White-collared swift Vencejo acollarado ABNUA02010 Aves Apodiformes Apodidae Shaw, 1796

Sturnus vulgaris European starling Estornino pinto ABPBT01010 Aves Passeriformes Sturnidae Linnaeus, 1758Sula dactylatra Masked booby Boba enmascarada ABNFB01010 Aves Pelecaniformes Sulidae Lesson, 1831Sula leucogaster Brown booby Boba parda ABNFB01030 Aves Pelecaniformes Sulidae Boddaert, 1783Sula sula Red-footed booby Boba patirroja ABNFB01040 Aves Pelecaniformes Sulidae Linnaeus, 1766Tachornis phoenicobia Antillean palm swift Vencejo Antíllano ABNUA08010 Aves Apodiformes Apodidae Gosse, 1847Tachybaptus dominicus Least grebe Tigua ABNCA01010 Aves Podicipediformes Podicipedidae Linnaeus, 1766Tachycineta bicolor Tree swallow Golondrina bicolor ABPAU03010 Aves Passeriformes Hirundinidae Vieillot, 1808

Tiaris bicolor Black-faced grassquit Gorrión negro ABPBX80030 Aves Passeriformes Emberizidae Linnaeus, 1766Tiaris olivacea Yellow-faced grassquit Gorrion barba amarilla ABPBX80020 Aves Passeriformes Emberizidae Linnaeus, 1766Todus mexicanus Puerto Rican tody San Pedrito ABNXB01050 Aves Coraciiformes Todidae Lesson, 1838

Tringa flavipes Lesser yellowlegs Playero guineilla pequeño ABNNF01030 Aves Charadriiformes Scolopacidae Gmelin, 1789Tringa melanoleuca Greater yellowlegs Playero guineilla grande ABNNF01020 Aves Charadriiformes Scolopacidae Gmelin, 1789Tringa nebularia Common greenshank Playero claro ABNNF01010 Aves Charadriiformes Scolopacidae Gunnerus, 1767Tringa solitaria Solitary sandpiper Playero solitario ABNNF01070 Aves Charadriiformes Scolopacidae Wilson, 1813Tryngites subruficollis Buff-breasted sandpiper Playero canela ABNNF14010 Aves Charadriiformes Scolopacidae Vieillot, 1819

Turdus migratorius American robin Zorzal petirrojo ABPBJ20170 Aves Passeriformes Turdidae Linnaeus, 1766
138

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 24—Terrestrial vertebrate species of Puerto Rico (continued) Scientific name English common name Spanish common name Species ID Class Order Family Authority Turdus plumbeus Red-legged thrush Zorzal de patas coloradas ABPBJ20210 Aves Passeriformes Turdidae Linnaeus, 1758 Tyrannus caudifasciatus Loggerhead kingbird Clerigo ABPAE52080 Aves Passeriformes Tyrannidae d’Orbigny, 1839 Tyrannus dominicensis Gray kingbird Pitirre gris ABPAE52070 Aves Passeriformes Tyrannidae Gmelin, 1788 Tyrannus forficatus Scissor-tailed flycatcher Pitirre rabotijera ABPAE52100 Aves Passeriformes Tyrannidae Gmelin, 1789 Tyrannus tyrannus Eastern kingbird Pitirre norteño ABPAE52060 Aves Passeriformes Tyrannidae Linnaeus, 1758 Tyto alba Barn owl Lechuza común ABNSA01010 Aves Strigiformes Tytonidae Scopoli, 1769 Vanellus vanellus Northern lapwing Avefría ABNNB01010 Aves Charadriiformes Charadriidae Linnaeus, 1758 Vermivora chrysoptera Golden-winged warbler Reinita alidorada ABPBX01030 Aves Passeriformes Parulidae Linnaeus, 1766 Vermivora ruficapilla Nashville warbler Chipe cachetigris ABPBX01060 Aves Passeriformes Parulidae Wilson, 1811 Vidua macroura Pin-tailed whydah Viuda colicinta ABPCB06010 Aves Passeriformes Estrildidae Pallas, 1764 Vireo altiloquus Black-whiskered vireo Julian Chivi ABPBW01250 Aves Passeriformes Vireonidae Vieillot, 1808

Vireo flavifrons Yellow-throated vireo Vireo gargantiamarillo ABPBW01170 Aves Passeriformes Vireonidae Vieillot, 1808Vireo griseus White-eyed vireo Julian Chivi ojiblanco ABPBW01020 Aves Passeriformes Vireonidae Boddaert, 1783Vireo latimeri Puerto Rican vireo Bienteveo de Puerto Rico ABPBW01090 Aves Passeriformes Vireonidae Baird, 1866

Vireo olivaceus Red-eyed vireo Vireo de ojo rojo ABPBW01240 Aves Passeriformes Vireonidae Linnaeus, 1766

Wilsonia canadensis Canada warbler Reinita de Canada ABPBX16030 Aves Passeriformes Parulidae Linnaeus, 1766Wilsonia citrina Hooded warbler Reinita viuda ABPBX16010 Aves Passeriformes Parulidae Boddaert, 1783

Wilsonia pusilla Wilson’s warbler Reinita gorrinegra ABPBX16020 Aves Passeriformes Parulidae Wilson, 1811

Zenaida asiatica White-winged dove Tórtola aliblanca ABNPB04010 Aves Columbiformes Columbidae Linnaeus, 1758

Zenaida aurita Zenaida dove Tórtola cardosantera ABNPB04020 Aves Columbiformes Columbidae Linnaeus, 1758Zenaida macroura Mourning dove Tórtola rabilarga ABNPB04040 Aves Columbiformes Columbidae Linnaeus, 1758

Amphibians (24 species)

Bufo lemur Puerto Rican crested toad Sapo concho AAABB02010 Amphibia Anura Bufonidae Cope, 1869

Bufo marinus Giant toad Sapo común AAABB01100 Amphibia Anura Bufonidae Linnaeus, 1758Eleutherodactylus

antillensis Antillean frog Churí AAABD04250 Amphibia Anura Leptodactylidae Reinhardt and

Lutken, 1863

Eleutherodactylus brittoni Grass coqui Coquí de las hierbas AAABD04090 Amphibia Anura Leptodactylidae Schmidt, 1920Eleutherodactylus cochranae Whistling frog Coquí pitito AAABD04180 Amphibia Anura Leptodactylidae Grant, 1932Eleutherodactylus cooki Rock coqui Coquí guajón AAABD04010 Amphibia Anura Leptodactylidae Grant, 1931Eleutherodactylus coqui Common coqui Coquí de las yerbas AAABD04100 Amphibia Anura Leptodactylidae Thomas, 1966Eleutherodactylus eneidae Eneida’s coqui Coquí de Eneida AAABD04060 Amphibia Anura Leptodactylidae Rivero, 1959

Eleutherodactylus gryllus Cricket coqui Coquí grillo AAABD04110 Amphibia Anura Leptodactylidae Schmidt, 1920

Eleutherodactylus hedricki Tree-hole frog Coquí de Hedrick AAABD04020 Amphibia Anura Leptodactylidae Rivero, 1963

Eleutherodactylus jasperi Golden coqui Coquí dorado AAABD04030 Amphibia Anura Leptodactylidae Drewry and

Jones, 1976
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The Puerto Rico Gap Analysis Project

Table 24—Terrestrial vertebrate species of Puerto Rico (continued)Scientific name English common name Spanish common name Species ID Class Order Family Authority

Eleutherodactylus karlschmidti Web-footed coqui Coquí palmeado AAABD04040 Amphibia Anura Leptodactylidae Grant, 1931Eleutherodactylus locustus Warty coqui Coquí martillito AAABD04120 Amphibia Anura Leptodactylidae Schmidt, 1920Eleutherodactylus monensis Mona coqui Coquí de la Mona AAABD04130 Amphibia Anura Leptodactylidae Meerwarth, 1920Eleutherodactylus Puerto Rican coqui Coquí de la montaña AAABD04140 Amphibia Anura Leptodactylidae Schmidt, 1920

portoricensis

Eleutherodactylus richmondi Ground coqui Coquí caoba AAABD04150 Amphibia Anura Leptodactylidae Stejneger, 1904

Eleutherodactylus unicolor Burrowing coqui Coquí duende AAABD04050 Amphibia Anura Leptodactylidae Stejneger, 1904Eleutherodactylus wightmanaeWrinkled frog Coquí melodioso AAABD04160 Amphibia Anura Leptodactylidae Schmidt, 1920Hyla cinerea Green treefrog Hila verde AAABC02060 Amphibia Anura Hylidae Schneider, 1799Leptodactylus albilabris White-lipped frog Ranita de blanco AAABD02020 Amphibia Anura Leptodactylidae Günther, 1859Osteopilus septentrionalis Cuban treefrog Hila platanera AAABC04010 Amphibia Anura Hylidae Duméril and

Bibron, 1841

Rana catesbeiana Bullfrog Rana toro AAABH01070 Amphibia Anura Ranidae Shaw, 1802Rana grylio Pig frog Rana AAABH01110 Amphibia Anura Ranidae Stejneger, 1901Scinax rubra Treefrog Hila inquieta AAABC08010 Amphibia Anura Hylidae Laurenti, 1768

Mammals (27 species)

Artibeus jamaicensis Jamaican fruit-eating bat Muciérlago frutero AMACB07010 Mammalia Chiroptera Phyllostomidae Leach, 1821

Brachyphylla cavernarum Antillean fruit-eating bat Murciélago cavernícola AMACB08010 Mammalia Chiroptera Phyllostomidae Gray, 1834Capra hircus Goat Cabra AMALE07010 Mammalia Artiodactyla Bovidae Linnaeus, 1758Eptesicus fuscus Big brown bat Murciélago marrón mayor AMACC04010 Mammalia Chiroptera Vespertilionidae Palisot de

Beauvois, 1796

Erophylla sezekorni Buffy flower bat Murciélago las flores AMACB09010 Mammalia Chiroptera Phyllostomidae Gundlach, 1860Felis catus Feral cat Gatto AMAJH01070 Mammalia Carnivora Felidae Linnaeus, 1775

Herpestes javanicus Small Indian mongoose Mangosta AMAJJ01010 Mammalia Carnivora Herpestidae E. Geoffroy SaintHilaire, 1818

Isolobodon portoricensis Puerto Rican hutia Jutía AMAFL01010 Mammalia Rodentia Capromyidae Allen, 1916

Lasiurus borealis Eastern red bat Murciélago rojo AMACC05010 Mammalia Chiroptera Vespertilionidae Muller, 1776Macaca mulatta Rhesus monkey Macaco AMAMB02010 Mammalia Primates Cercopithecidae Zimmerman, 1780Macrotus waterhousii Waterhouse’s leaf-nosed bat Murciélago orejudo LMGK025020 Mammalia Chiroptera Phyllostomidae Gray, 1843

Molossus molossus Velvety free-tailed bat Murciélago casero AMACD03010 Mammalia Chiroptera Molossidae Pallas, 1766

Monachus tropicalis West Indian monk seal AMAJG04010 Mammalia Carnivora Phocidae Gray, 1850Monophyllus plethodon Insular long-tongued bat Murciélago lenguilargo AMACB06010 Mammalia Chiroptera Phyllostomidae Miller, 1900

Monophyllus redmani Puerto Rican long-tongued bat Murciélago AMACB06021 Mammalia Chiroptera Phyllostomidae Leach, 1821

portoricensis

Mormoops blainvillii Antillean ghost-faced bat Murciélago barbicacho AMACA01020 Mammalia Chiroptera Mormoopidae Leach, 1821

Mus musculus House mouse Ratón casero AMAFF22010 Mammalia Rodentia Muridae Linnaeus, 1758Noctilio leporinus Greater bulldog bat Murciélago pescador AMACE01010 Mammalia Chiroptera Noctilionidae Linnaeus, 1758Odocoileus virginianus White-tailed deer Venado rabiblanco AMALC02020 Mammalia Artiodactyla Cervidae Zimmermann, 1780
140

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 24—Terrestrial vertebrate species of Puerto Rico (continued) Scientific name English common name Spanish common name Species ID Class Order Family Authority Pteronotus parnellii Parnell’s mustached bat Murciélago bigotudo mayor AMACA02011 Mammalia Chiroptera Mormoopidae Miller, 1902 portoricensis Pteronotus quadridens Sooty mustached bat Murciélago bigotudo menor AMACA02020 Mammalia Chiroptera Mormoopidae Gundlach, 1840 Rattus norvegicus Norway rat Rata de Noruega AMAFF21020 Mammalia Rodentia Muridae Berkenhout, 1769 Rattus rattus Black rat Rata negra AMAFF21010 Mammalia Rodentia Muridae Linnaeus, 1758 Stenoderma rufum Desmarest’s fig-eating bat Murciélago rojo frutero AMACB05010 Mammalia Chiroptera Phyllostomidae Desmarest, 1820 Sus scrofa Feral hog Cerdo AMALA01010 Mammalia Artiodactyla Suidae Linnaeus, 1758 Tadarida brasiliensis Brazilian free-tailed bat Murciélago de cola libre AMACD01010 Mammalia Chiroptera Molossidae I. Geoffroy, 1824 Trichechus manatus West Indian manatee Manati Antíllano AMAKA01010 Mammalia Sirenia Trichechidae Linnaeus, 1758 Reptiles (57 species)

Alsophis portoricensis Puerto Rican racer Culebra Corredora ARADB40010 Reptilia Squamata Colubridae Reinhardt and

Lütken, 1862

Ameiva alboguttata Mona ground lizard Siguan de la Mona ARACJ01042 Reptilia Squamata Teiidae Boulenger, 1896

Ameiva desechensis Desecheo ground lizard Siguana de Desecheo ARACJ01043 Reptilia Squamata Iguanidae Heatwole and

Torres, 1967

Ameiva exsul Puerto Rican ground lizard Siguana común ARACJ01040 Reptilia Squamata Teiidae Cope, 1862Ameiva wetmorei Blue-tailed ground lizard Siguana de rabo azul ARACJ01030 Reptilia Squamata Teiidae Stejneger, 1913Amphisbaena bakeri Baker’s legless lizard Culebrita ciega de Baker ARACA02010 Reptilia Squamata Amphisbaenidae Stejneger, 1904

Amphisbaena caeca Puerto Rican worm lizard Culebrita ciega común ARACA02020 Reptilia Squamata Amphisbaenidae Cuvier, 1829Amphisbaena schmidti Schmidt’s worm lizard Culebrita ciega de Schmidt ARACA02030 Reptilia Squamata Amphisbaenidae Gans, 1964Amphisbaena xera North American worm lizard Culebrita ciega del seco ARACA02040 Reptilia Squamata Amphisbaenidae Thomas, 1966Anolis cooki Cook’s anole Lagartijo del seco ARACF01100 Reptilia Squamata Iguanidae Grant, 1931Anolis cristatellus Crested anole Lagartijo común ARACF01020 Reptilia Squamata Iguanidae Duméril and

Bibron, 1837

Anolis cuvieri Puerto Rican giant anole Lagarto verde ARACF01110 Reptilia Squamata Iguanidae Merrem, 1820

Anolis desechensis Desecheo anole Lagartijo de Desecheo ARACF01120 Reptilia Squamata Iguanidae Heatwole, 1976

Anolis evermanni Emerald anole Lagartijo verde ARACF01130 Reptilia Squamata Iguanidae Stejneger, 1904Anolis gundlachi Yellow-bearded anole Lagartijo barba amarilla ARACF01140 Reptilia Squamata Iguanidae Peters, 1876Anolis krugi Upland grass anole Lagartijo jardinero de ARACF01150 Reptilia Squamata Iguanidae Peters, 1876

montana

Anolis monensis Mona anole Lagartijo de la Mona ARACF01160 Reptilia Squamata Iguanidae Stejneger, 1904

Anolis occultus Puerto Rican pygmy anole Lagartijo pigmeo ARACF01080 Reptilia Squamata Iguanidae Williams and

Rivero, 1965

Anolis poncensis Dryland grass anole Lagartijo jardinero de Ponce ARACF01170 Reptilia Squamata Iguanidae Stejneger, 1904

Anolis pulchellus Common grass anole Lagartijo jardinero ARACF01210 Reptilia Squamata Iguanidae Duméril and

Bibron, 1837
141

The Puerto Rico Gap Analysis Project

Table 24—Terrestrial vertebrate species of Puerto Rico (continued)

Scientific name English common name Spanish common name Species ID Class Order Family Authority

Anolis roosevelti Culebra Island giant anole Lagartijo gigante de ARACF01070 Reptilia Squamata Iguanidae Grant, 1931

Culebra

Anolis stratulus Barred anole Lagartijo manchado ARACF01200 Reptilia Squamata Iguanidae Cope, 1861Arrhyton exiguum Culebra garden snake Culebra de jardín ARADB41010 Reptilia Squamata Colubridae Cope, 1862Caiman crocodilus Spectacled caiman Baba ARABA02010 Reptilia Crocodila Alligatoridae Linnaeus, 1758Caretta caretta Loggerhead sea turtle Cabezón ARAAA01010 Reptilia Testudines Cheloniidae Linnaeus, 1758Chelonia mydas Green sea turtle Pejeblanco ARAAA02010 Reptilia Testudines Cheloniidae Linnaeus, 1758Cyclura nubila Cuban ground iguana Iguana Cubana ARACF18030 Reptilia Squamata Iguanidae Gray, 1831Cyclura stejnegeri Mona Island rock iguana Iguana de la Mona ARACF18010 Reptilia Squamata Iguanidae Barbour and

Noble, 1916

Dermochelys coriacea Leatherback sea turtle Tinglar ARAAC01010 Reptilia Testudines dermochelyidae Vandelli, 1761Diploglossus pleei Puerto Rican galliwasp Celestus ARACB03010 Reptilia Squamata Anguidae Duméril and

Bibron, 1839

Epicrates inornatus Puerto Rican boa Culebrón ARADA03020 Reptilia Squamata Boidae Reinhardt, 1843Epicrates monensis granti Virgin Islands tree boa Culebrón de la Isla Virgin ARADA03011 Reptilia Squamata BoidaeEpicrates monensis monensis Mona boa Culebrón de la Mona ARADA03012 Reptilia Squamata Boidae Eretmochelys imbricata Hawksbill Carey ARAAA03010 Reptilia Testudines Cheloniidae Linnaeus, 1766

Hemidactylus brookii Brook’s house gecko Salamanquesa ARACD03040 Reptilia Squamata Gekkonidae Gray, 1845Hemidactylus mabouia Cosmopolitan house gecko Salamanquesa ARACD03050 Reptilia Squamata Gekkonidae Moreau de Jonnés,

1818

Iguana iguana Green iguana Iguana común ARACF09010 Reptilia Squamata Iguanidae Linnaeus, 1758Lepidochelys olivacea Olive Ridley sea turtle Tortuga bastarda ARAAA04020 Reptilia Testudines Cheloniidae Eschscholtz, 1829Mabuya mabouya sloanei Slippery-backed mabuya Lucía ARACH04011 Reptilia Squamata Scincidae Daudin, 1803Phyllodactylus wirshingi Greater Antillean leaf-toed gecko Salamanquesa bandeada ARACD04020 Reptilia Squamata Gekkonidae Kerster and

Smith, 1955

Sphaerodactylus gaigeae Gaige’s dwarf gecko Salamanquita de Pandura ARACD05050 Reptilia Squamata Gekkonidae Grant, 1932Sphaerodactylus klauberi Klauber’s dwarf gecko Salamanquita negra ARACD05060 Reptilia Squamata Gekkonidae Grant, 1931Sphaerodactylus levinsi Desecheo gecko Salamanquita de Desecheo ARACD05070 Reptilia Squamata Gekkonidae Heatwole, 1968

Sphaerodactylus macrolepis Common dwarf gecko Salamanquita común ARACD05110 Reptilia Squamata Gekkonidae Gunther, 1859Sphaerodactylus Monito gecko Salamanquita de Monito ARACD05040 Reptilia Squamata Gekkonidae Schwartz, 1977

micropithecus

Sphaerodactylus monensis Mona dwarf gecko Salamanquita de la Mona ARACD05080 Reptilia Squamata Gekkonidae Meerwarth, 1901Sphaerodactylus nicholsi Nichol’s dwarf gecko Salamanquita pigmea ARACD05090 Reptilia Squamata Gekkonidae Grant, 1931Sphaerodactylus roosevelti Roosevelt’s dwarf gecko Salamanquita de Roosevelt ARACD05010 Reptilia Squamata Gekkonidae Grant, 1931Sphaerodactylus townsendi Townsend’s dwarf gecko Salamanquita del sureste ARACD05120 Reptilia Squamata Gekkonidae Grant, 1931

Thamnophis sirtalis Common garter snake Culebra listonada común ARADB36130 Reptilia Squamata Colubridae Linnaeus, 1758
142

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 24—Terrestrial vertebrate species of Puerto Rico (continued) Scientific name English common name Spanish common name Species ID Class Order Family Authority Trachemys stejnegeri stejnegeri Puerto Rican slider Hicotea ARAAD09030 Reptilia Testudines Emydidae Schmidt, 1928 Typhlops granti Grant’s blind snake Vibora de Grant ARADG02010 Reptilia Squamata Typhlopidae Ruthven and Gaige, 1935 Typhlops hypomethes Blind snake Vibora Universitaria ARADG02060 Reptilia Squamata Typhlopidae Hedges and Thomas, 1991 Typhlops monensis Mona blind snake Vibora de la Mona ARADG02020 Reptilia Squamata Typhlopidae Schmidt, 1926 Typhlops platycephalus Blind snake Vibora de cabeza aplastada ARADG02050 Reptilia Squamata Typhlopidae Duméril and Bibron, 1844 Typhlops richardii Richard’s blind snake Vibora común ARADG02030 Reptilia Squamata Typhlopidae Duméril and Bibron, 1844 Typhlops rostellatus Puerto Rican wetland blind snake Vibora de pico ARADG02040 Reptilia Squamata Typhlopidae Stejneger, 1904
143

The Puerto Rico Gap Analysis Project

Appendix 3. Land Cover Vegetation ClassificationIdentification codes (values) from the raster image land cover map, unit name, physiography, moisture class, substrate, forest age if applicable, landforms on which class typically occurs, formation of vegetation type (from Dansereau 1966 or Borhidi 1996 when noted; otherwise from Ahern et al.1999, and Areces-Mallea et al. 1999), area in hectares (ha) and percentage of Puerto Rico for 70 land cover units in the PRGAP land cover map anddatabase.Table 25—Puerto Rico land cover type site featuresValue Class name Physiognomy Moisture Substrate Age Land form Formation Area Percentage

Years Hectares

1 Mature secondary lowland Closed forest Dry Alluvial > 25 Plains and Closed, broadleaf mixed, 1548 0.2

dry alluvial semideciduous valleys lowland subtropical semiforest deciduous forest.

2 Young secondary lowland dry Closed forest Dry Alluvial < 25 Plains and Closed, broadleaf mixed, 2881 0.3

alluvial semideciduous forest valleys lowland subtropical semideciduous forest.

3 Lowland dry alluvial Open forest, Dry Alluvial NA Plains and Open, broadleaf, lowland 4472 0.5

shrubland and woodland woodland, or valleys subtropical semideciduous

shrubland woodland and shrubland.

46 Lowland dry riparian forest Closed forest Dry Alluvial NA Valleys Closed, broadleaf, lowland 806 0.1

subtropical seasonal evergreen forest.

47 Lowland dry riparian Open forest, Dry Alluvial NA Valleys Open, broadleaf, lowland 423 0.1

shrubland and woodland woodland or subtropical semideciduous

shrubland woodland and shrubland.

4 Mature secondary lowland Closed forest Dry Calcareous, > 25 Ridges, slopes, Closed, broadleaf, lowland 982 0.1

dry limestone evergreen limestone valleys, and subtropical seasonalforest plains evergreen forest.

5 Mature secondary lowland Closed forest Dry Calcareous, > 25 Ridges, slopes, Closed, broadleaf, mixed 10 669 1.2

dry limestone limestone valleys, and lowland subtropical semisemideciduous forest plains deciduous forest.

6 Young secondary lowland Closed forest Dry Calcareous, < 25 Ridges, slopes, Closed, broadleaf, mixed 3922 0.4

dry limestone semideciduous limestone valleys, and lowland subtropical semiforest plains deciduous forest

7 Lowland dry limestone Open forest, Dry Calcareous, NA Ridges, slopes, Open, broadleaf, lowland 8181 0.9

woodland and shrubland woodland, or limestone valleys, and subtropical semideciduous

shrubland plains woodland and shrubland.
144

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 25—Puerto Rico land cover type site features (continued) Value Class name Physiognomy Moisture Substrate Age Land form Formation Area Percentage Years Hectares 8 Lowland dry limestone Shrubland Dry Calcareous, NA Ridges, slopes, Closed, broadleaf, lowland 4694 0.5 shrubland limestone valleys, and subtropical semideciduous plains shrubland. 9 Lowland dry cactus Shrubland Dry Calcareous, > 25 Ridges, slopes, Semidesert scrubs with 47 0.0 shrubland limestone valleys, and columnar cacti (Borhidi plains 1996). 10 Coastal dwarf woodland and Shrubland Dry Calcareous, > 25 Ridges, slopes, Open, broadleaf lowland 103 0.0 shrubland limestone valleys, and subtropical semideciduous plains woodland and shrubland. 11 Lowland dry limestone Closed forest Dry Calcareous, > 25 Steep slopes Closed, broadleaf, mixed 12 0.0

cliffside semideciduous limestone lowland subtropical semiforest deciduous forest.

12 Lowland dry limestone Open forest, Dry Calcareous, > 25 Steep slopes Open, broadleaf, lowland 49 0.0

cliffside shrubland and woodland, or limestone subtropical semideciduous

woodland shrubland woodland and shrubland.

19 Abandoned dry forest Closed forest Dry Calcareous, > 25 Ridges, slopes, Closed, broadleaf, lowland 81 0.0

plantation limestone valleys, and subtropical seasonal

plains evergreen forest.

13 Mature secondary lowland Closed forest Dry Noncalcar- > 25 Ridges, slopes, Closed, broadleaf mixed, 7062 0.8

dry noncalcareous eous, valleys, and lowland subtropical semisemideciduous forest volcanic plains deciduous forest.

14 Young secondary lowland Closed forest Dry Noncalcar- < 25 Ridges, slopes, Closed, broadleaf mixed, 3645 0.4

dry noncalcareous eous, valleys, and lowland subtropical semisemideciduous forest volcanic plains deciduous forest.

15 Lowland dry noncalcareous Open forest, Dry Noncalcar- NA Ridges, slopes, Open, broadleaf, lowland 13 307 1.5

shrubland and woodland woodland, or eous, valleys, and subtropical semideciduous

shrubland volcanic plains woodland and shrubland.

16 Mature secondary dry and Closed forest Dry Ultramafic, > 25 Ridges, slopes, Closed, broadleaf mixed, 1839 0.2

moist serpentine serpentine valleys, and lowland subtropical semisemideciduous forest plains deciduous forest.

17 Young secondary dry and Closed forest Dry Ultramafic, < 25 Ridges, slopes, Closed, broadleaf mixed, 1960 0.2

moist serpentine serpentine valleys, and lowland subtropical semisemideciduous forest plains deciduous forest.

18 Dry and moist serpentine Open forest, Dry Ultramafic, NA Ridges, slopes, Arid lowland serpentine

woodland and shrubland woodland, serpentine valleys, and shrubwoods (Borhidi 1996). 841 0.1

or shrubland plains

20 Mature secondary lowland Closed forest Moist Alluvial > 25 Plains and Closed, broadleaf evergreen, 2256 0.3

moist alluvial evergreen valleys lowland subtropical seasonalforest evergreen forest. Lowland

rain forest (Dansereau 1996).
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The Puerto Rico Gap Analysis Project

Table 25—Puerto Rico land cover type site features (continued)Value Class name Physiognomy Moisture Substrate Age Land form Formation Area Percentage

Years Hectares

21 Young secondary lowland Closed forest Moist Alluvial < 25 Plains and Closed, broadleaf evergreen, 6675 0.8

moist alluvial evergreen valleys lowland subtropical seasonalforest evergreen forest.

22 Lowland moist alluvial Open forest, Moist Alluvial NA Plains and Open, broadleaf evergreen, 5141 0.6

shrubland and woodland woodland, or valleys lowland subtropical seasonal

shrubland evergreen woodland and

shrubland.

68 Lowland moist riparian forest Closed forest Moist Alluvial > 25 Valleys Closed, broadleaf evergreen, 916 0.1

lowland subtropical seasonalevergreen forest. Lowlandrainforest (Dansereau 1996).

69 Lowland moist riparian Open forest, Moist Alluvial NA Valleys Open, broadleaf evergreen, 508 0.1

shrubland and woodland woodland, or lowland subtropical seasonal

shrubland evergreen woodland and

shrubland.

23 Mature secondary moist Closed forest Moist Calcareous, > 25 Ridges, slopes, Closed, broadleaf mixed, 46 423 5.2

limestone evergreen and limestone and valleys lowland subtropicalsemideciduous forest semideciduous forests.

Karstic forests (Borhidi1996).

24 Young secondary moist Closed forest Moist Calcareous, < 25 Ridges, slopes, Closed, broadleaf mixed, 15 494 1.7

limestone evergreen and limestone and valleys lowland subtropicalsemideciduous forest semideciduous forests.

Karstic forests (Borhidi1996).

25 Moist limestone shrubland Open forest, Moist Calcareous, NA Ridges, slopes, Open, broadleaf mixed, low- 10 338 1.2

and woodland woodland, or limestone and valleys land subtropical semishrubland deciduous forests.

26 Mature secondary lowland Closed forest Moist Noncalcar- > 25 Ridges, slopes, Closed, broadleaf evergreen, 40 446 4.5

moist noncalcareous eous, valleys, and lowland subtropical seasonal

evergreen forest volcanic plains evergreen forest. Lowland

rain forest (Dansereau 1996).

27 Young secondary lowland Closed forest Moist Noncalcar- < 25 Ridges, slopes, Closed, broadleaf evergreen, 52 981 5.9

moist noncalcareous eous, valleys, and lowland subtropical seasonal

evergreen forest volcanic plains evergreen forest.

28 Lowland moist noncalcareous Open forest, Moist Noncalcar- NA Ridges, slopes, Open, broadleaf evergreen, 44 523 5.0

shrubland and woodland woodland, or eous, valleys, and lowland subtropical seasonal

shrubland volcanic plains evergreen woodland and

shrubland.
146

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 25—Puerto Rico land cover type site features (continued) Value Class name Physiognomy Moisture Substrate Age Land form Formation Area Percentage Years Hectares 29 Lowland moist abandoned Closed forest Moist Noncalcar- > 25 Ridges, slopes, Closed, broadleaf evergreen, 11 068 1.2 and active coffee eous, valleys, and lowland subtropical seasonal plantations volcanic plains evergreen forest. 30 Mature secondary montane Closed forest Wet Alluvial > 25 Plains and Closed, broadleaf evergreen, 613 0.1 wet alluvial evergreen valleys montane subtropical rain alluvial forest forest. Montane rain forests (Borhidi 1966). 31 Young secondary montane Closed forest Wet Alluvial < 25 Plains and Closed, broadleaf evergreen, 988 0.1 wet alluvial evergreen forest valleys montane subtropical rain forest. Montane rain forests (Borhidi 1966).

32 Montane wet alluvial Open forest, Wet Alluvial NA Plains and Open, broadleaf evergreen, 900 0.1

shrubland and woodland woodland, or valleys montane subtropical

shrubland woodland and shrubland.

33 Mature secondary montane Closed forest Wet Noncalcar- > 25 Ridges, slopes, Closed, broadleaf evergreen, 14 968 1.7

wet noncalcareous evergreen eous, and valleys montane subtropical rain

forest volcanic forest. Montane rain forests

(Borhidi 1966).

43 Montane wet evergreen Closed forest Wet Noncalcar- > 25 Ridges, slopes, Closed, broadleaf evergreen, 54 859 6.1

abandoned and active eous, and valleys montane subtropical rain

coffee plantation volcanic forest.

34 Mature primary and Closed forest Wet Noncalcar- > 25 Ridges, slopes, Closed, broadleaf evergreen, 8856 1.0

secondary montane wet eous, and valleys montane subtropical rain

noncalcareous evergreen volcanic forest. Montane rain foreststabonuco forest of the Caribbean

(Borhidi 1966).

35 Mature primary and Closed forest Wet Noncalcar- > 25 Ridges, slopes, Closed, broadleaf evergreen, 15 583 1.7

secondary montane wet eous, and valleys montane subtropical cloud

noncalcareous evergreen volcanic forest. Cloud forests of thepalo colorado cloud forest Antillean high mountains

(Borhidi 1966).

36 Mature primary and Closed forest Wet Noncalcar- > 25 Ridges, slopes, Closed, broadleaf evergreen, 8781 1.0

secondary montane wet eous, and valleys montane subtropical rain

noncalcareous evergreen volcanic forest. Montane rain forestssierra palm forest of the Caribbean (Borhidi

1966).

37 Mature primary and Closed forest Wet Noncalcar- > 25 Ridges and Closed, broadleaf evergreen, 2936 0.3

secondary montane wet eous, slopes montane subtropical cloudnoncalcareous evergreen volcanic forest. Cloud forests of theelfin woodland cloud forest Antillean high mountains

(Borhidi 1966).
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The Puerto Rico Gap Analysis Project

Table 25—Puerto Rico land cover type site features (continued)Value Class name Physiognomy Moisture Substrate Age Land form Formation Area Percentage

Years Hectares

38 Young secondary montane Closed forest Wet Noncalcar- < 25 Ridges, slopes, Closed, broadleaf evergreen, 21 651 2.4

wet noncalcareous evergreen eous, and valleys montane subtropical rain

forest volcanic forest. Montane rain forests

of the Caribbean (Borhidi1966).

39 Montane wet noncalcareous Open forest, Wet Noncalcar- NA Ridges, slopes, Open, broadleaf evergreen, 24 090 2.7

evergreen shrubland and woodland, or eous, and valleys montane subtropical

woodland shrubland volcanic woodland and shrubland.

Moist, high montaneshrubwoods (Borhidi 1996).

40 Mature secondary montane Closed forest Wet Ultramafic, > 25 Ridges, slopes, Closed, broadleaf evergreen, 3379 0.4

wet serpentine evergreen serpentine and valleys montane subtropical rain

forest forest.

41 Young secondary montane Closed forest Wet Ultramafic, < 25 Ridges, slopes, Closed, broadleaf evergreen, 651 0.1

wet serpentine evergreen serpentine and valleys montane subtropical rain

forest forest.

42 Wet serpentine shrubland Open forest, Wet Ultramafic, NA Ridges, slopes, Open, broadleaf evergreen, 358 0.0

and woodland woodland, or serpentine and valleys montane subtropical

shrubland woodland and shrubland.

44 Mangrove forest and Mixed closed Flooded Marine > 25 Plains Mixed closed and open, 8699 1.0

shrubland and open sediments flooded, tidally flooded

forest evergreen sclerophyllus

forest. Mangrove (Dansereau1966, Borhidi 1996)

45 Freshwater Pterocarpus Closed forest Flooded Alluvial > 25 Plains and Closed, flooded, subtropical 261 0.0

swamp deposits valleys seasonally flooded rain forest.

48 Dry grasslands and pastures Grassland Dry NA Plains, slopes, Subtropical grassland with a 42 176 4.7

and valleys shrub layer.

49 Dry cactus grassland and Grassland Dry NA Plains, slopes, Subtropical grassland with a 213 0.0

shrubland and valleys shrub layer.

50 Moist grasslands and pastures Grassland Moist NA Plains, slopes, Subtropical grassland with a 218 945 24.5

and valleys shrub layer.

53 Seasonally flooded Grassland Seasonally Nonsaline NA Plains, Seasonally or temporarily 18 225 2.0

herbaceous nonsaline flooded alluvial depressions, flooded nonsaline subtropical

wetlands deposits and valleys grassland with a shrub layer.
148

GENERAL TECHNICAL REPORT IITF-GTR-39 Table 25—Puerto Rico land cover type site features (continued) Value Class name Physiognomy Moisture Substrate Age Land form Formation Area Percentage Years Hectares 54 Seasonally flooded Grassland Seasonally Saline NA Plains, Seasonally or temporarily 4865 0.5 herbaceous saline wetlands flooded alluvial depressions, flooded saline subtropical deposits and valleys grassland with a shrub layer. and marine sediments 51 Emergent herbaceous Herbaceous Flooded Nonsaline NA Plains, Perennially flooded nonsaline 1102 0.1 nonsaline wetlands alluvial depressions, aquatic herbaceous deposits and valleys vegetation. 52 Emergent herbaceous saline Herbaceous Flooded Saline NA Plains, Perennially flooded nonsaline 882 0.1 wetlands alluvial depressions, aquatic herbaceous

deposits and valleys vegetation.and marinesediments

55 Hay and row crops Herbaceous Dry and Alluvial NA Plains, Herbaceous agriculture. 25 764 2.9

moist deposits slopes,

and valleys

56 Woody agriculture and Closed forest Dry and Alluvial NA Plains Orchards. 492 0.1

plantations moist deposits

57 Rocky cliffs and shelves Barrens NA NA Steep slopes NA 397 0.0

and plains

58 Gravel beaches and stony Barrens NA NA Plains, NA 82 0.0

shoreline depressions,

and valleys

59 Fine to coarse sandy beaches, Barrens NA NA Plains, NA 1155 0.1

mixed sand and gravel beaches depressions,

and valleys

60 Riparian and other natural barrens Barrens NA NA Plains, NA 449 0.1

depressions,and valleys

61 Salt and mudflats Barrens NA NA Plains, NA 1493 0.2

depressions,and valleys

65 Artificial barrens Barrens NA NA Ridges, slopes, NA 8674 1.0

valleys, andplains
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The Puerto Rico Gap Analysis Project

Table 25—Puerto Rico land cover type site features (continued)Value Class name Physiognomy Moisture Substrate Age Land form Formation Area Percentage

Years Hectares

62 Salt production Barrens NA NA Plains, NA 20 0.0

depressions,and valleys

63 High-density urban Developed NA NA Ridges, slopes, NA 52 335 5.9

development valleys, and

plains

64 Low-density urban Developed NA NA Ridges, slopes, NA 37 237 4.2

development valleys, and

plains

66 Freshwater Water NA NA Plains, NA 4434 0.5

depressions,and valleys

67 Saline water Water NA NA Plains and NA 4005 0.5

depressions

70 Aquaculture Water NA NA Plains NA 101 0.0
150

GENERAL TECHNICAL REPORT IITF-GTR-39 Appendix 4. Puerto Rico Land Cover Descriptions Identification codes (values) from the raster image land cover map, unit name, and general description for 70 land cover units in the PRGAP land cover map and database. Table 26—Descriptions of land cover types in Puerto Rico Value Land cover type General description 1 Mature secondary lowland dry Mature dry forest on alluvial material of the coastal plain on the northeast and south coast of the main island and alluvial semideciduous forest on Vieques, Culebra, and Mona. These remnants are in relatively undisturbed areas, often along riparian corridors. Much of this extensive substrate has been converted to agriculture and pasture. Conditions are somewhat more moist than surroundings because of the riparian location. 2 Young secondary lowland dry Young secondary dry forest on alluvial material of the coastal plain on the northeast and south coast of the main alluvial semideciduous forest island. This is often disturbed by fire and grazing and developed on abandoned agricultural lands.

3 Lowland dry alluvial shrubland Open forest and shrubland developing on abandoned pasture of the southern coastal plain. This is often

and woodland disturbed by fire and grazing.

46 Lowland dry riparian forest Semideciduous and evergreen forests along the riparian corridors in the southwestern part of the island. Often

remnants of formerly extensive southern coastal plain forests on alluvial material. More evergreen speciesbecause of greater available moisture along the riparian corridor.

47 Lowland dry riparian shrubland Open forest and shrubland along riparian corridors of the southern coastal plain. This is often disturbed by fire

and woodland and grazing.

4 Mature secondary lowland dry Evergreen forests in arroyos, sinkholes, and narrow valleys on limestone substrates in the southwestern part of the

limestone evergreen forest main island and in the bajuras (sinkholes) of Mona Island.

5 Mature secondary lowland dry Mature semideciduous forests on mid and upper slopes of limestone hills on the southwestern part of the main

limestone semideciduous forest island and Caja de Muertos and on the limestone plateau of Mona Island.

6 Young secondary lowland dry Young secondary semideciduous forests on mid and upper slopes of limestone hills on the southwestern part of

limestone semideciduous forest the main island. These often on abandoned agricultural land.

7 Lowland dry limestone woodland Open semideciduous forests and shrublands on limestone hills in the southwestern part of the main island, Caja

and shrubland de Muertos, and Mona Island. These often on abandoned agricultural land or semi-active pastures.

8 Lowland dry limestone shrubland Dense thorny shrublands on limestone substrates in the southwestern part of the island, primarily abandoned

pasture, and on the eastern edge of Mona Island, strongly influenced by salt-laden winds restricting tree growth.

9 Lowland dry cactus shrubland Open shrubland with cactus on limestone substrates, often with poor soil development and much exposed

bedrock.

10 Coastal dwarf woodland and Open forest of dwarfed trees in wind-exposed coastal sites on the southwest side of the island, somewhat easterly

shrubland exposure. Very old but gnarled and twisted trees with horizontal stem growth on leeward side. Little soil

development and much exposed bedrock.

11 Lowland dry limestone cliffside Narrow bands of closed semideciduous forest on limestone cliff edges and along the base, primarily on Mona

semideciduous forest Island.
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The Puerto Rico Gap Analysis Project

Table 26—Descriptions of landcover types in Puerto Rico (continued)Value Land cover type General description

12 Lowland dry limestone cliffside Narrow bands of open semideciduous forest or woodland on limestone cliff edges and along the base, primarily

shrubland and woodland on Mona Island.

19 Abandoned dry forest plantation Abandoned plantations of Casuarina and Swietenia on limestone substrates on Mona Island, primarily on the

coastal plain.

13 Mature secondary lowland dry Mature dry forest on substrates derived from volcanic bedrock. These are found on coastal hills in northeastern

noncalcareous semideciduous Puerto Rico and in the Sierra Bermeja range in southwestern Puerto Rico. It occurs on noncalcareous substrates

forest in the lowland subtropical dry Holdridge life zone (Ewel and Whitmore 1973).

14 Young secondary lowland dry Young secondary dry forest on substrates derived from volcanic bedrock. These are found on coastal hills in

noncalcareous semideciduous northeastern Puerto Rico and in the Sierra Bermeja range in southwestern Puerto Rico. They are typicallyforest pastures abandoned in the last 10 to 25 years.

15 Lowland dry noncalcareous Open dry forest (woodland) and shrubland on substrates derived from volcanic bedrock. These are found on

shrubland and woodland coastal hills in northeastern Puerto Rico and in the Sierra Bermeja range in southwestern Puerto Rico.

They are typically recently abandoned or marginally active pastures.

16 Mature secondary dry and moist Mature forests on ultramafic (high pH) serpentine soils in the wetter Maricao and drier Susua areas of Puerto Rico.

serpentine semideciduous forest

17 Young secondary dry and moist Young secondary forests on ultramafic (high pH) serpentine soils in the wetter Maricao and drier Susua areas of

serpentine semideciduous forest Puerto Rico.

18 Dry and moist serpentine Open forests (woodland) on ultramafic (high pH) serpentine soils in the wetter Maricao and drier Susua areas of

woodland and shrubland Puerto Rico.

20 Mature secondary lowland moist Mature lowland moist forests on alluvial material in the northern and eastern coastal plains and lower valleys,

alluvial evergreen forest often associated with riparian corridors.

21 Young secondary lowland moist Young secondary lowland moist forests on alluvial material in the northern and eastern coastal plains and lower

alluvial evergreen forest valleys, often developed on abandoned pasture or former sugar cane fields.

22 Lowland moist alluvial shrubland Lowland moist woodland and shrubland on alluvial material in the northern and eastern coastal plains and lower

and woodland valleys, often developed on abandoned or semi-active pasture.

68 Lowland moist riparian forest Riparian forests along rivers in the northern and eastern coastal plains, either relicts of formerly extensive forests

or, more commonly, regenerating forests on abandoned agricultural land.

69 Lowland moist riparian shrubland Riparian woodland and shrubland along rivers in the northern and eastern coastal plains on abandoned or semiand woodland active pastures. Often disturbed by grazing and flooding.

23 Mature secondary moist limestone A matrix of mature forest types occurring along a topographic sequence in the haystack hills or “mogotes” of the

evergreen and semideciduous northern karst region of Puerto Rico. These are mature forest types with drier conditions on the ridges and steepforest slopes and more mesic conditions on lower slopes and valley bottoms. Much have regenerated on longabandoned (> 25 years) agricultural fields, either sugar cane, shade coffee, or pasture.
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GENERAL TECHNICAL REPORT IITF-GTR-39 Table 26—Descriptions of landcover types in Puerto Rico (continued) Value Land cover type General description 24 Young secondary moist limestone A matrix of young secondary forest types occurring along a topographic sequence in the haystack hills or evergreen and semideciduous “mogotes” of the northern karst region of Puerto Rico. These are early successional forest types with drier forest conditions on the ridges and steep slopes and more mesic conditions on lower slopes and valley bottoms. Much have regenerated on recently abandoned (10 to 25 years) agricultural fields, either sugar cane, shade coffee, or pasture. 25 Moist limestone shrubland and Shrubland and woodland in the haystack hills or “mogotes” of the northern karst region of Puerto Rico. Most are woodland abandoned or semi-active pasture. 26 Mature secondary lowland moist Mature lowland moist forest on substrates primarily derived from volcanic material on the northern coastal plain, noncalcareous evergreen forest lower elevations of the central and Luquillo Mountains, the Caguas Valley, and southeastern Puerto Rico. These occur primarily on long-abandoned (> 25 years) sugar cane, shade coffee, and pasture lands but contain remnants

of older lowland moist forests and harbor a majority of native tree species. It is likely similar to formerlyextensive moist lowland forests converted to agriculture in the preceding century. It occurs in the lowlandsubtropical moist Holdridge life zone (Ewel and Whitmore).

27 Young secondary lowland moist Young secondary lowland moist forest on substrates primarily derived from volcanic material on the northern

noncalcareous evergreen forest coastal plain, lower elevations of the central and Luquillo Mountains, the Caguas Valley, and southeastern

Puerto Rico. These occur primarily on recently abandoned (10 to 25 years) pasture lands and have a number ofestablished exotic species.

28 Lowland moist noncalcareous Young secondary lowland moist forest on substrates primarily derived from volcanic material on the northern

shrubland and woodland coastal plain, lower elevations of the central and Luquillo Mountains, the Caguas Valley, and southeastern

Puerto Rico. These occur primarily on recently abandoned or semi-active pasture lands.

29 Lowland moist abandoned and A mix of active sun coffee plantations and abandoned shade coffee plantations in the lower elevations of the

active coffee plantations central mountains.

30 Mature secondary montane wet Mature secondary wet evergreen forests on small patches of alluvial material in the upper central and Luquillo

alluvial evergreen forest Mountains. These are areas with less steep terrain and better soil development, and deeper soils than surrounding

volcanic substrates.

31 Young secondary montane wet Young secondary wet evergreen forests on small patches of alluvial material in the upper central and Luquillo

alluvial evergreen forest Mountains. These are areas with less steep terrain and better soil development, and deeper soils than surrounding

volcanic substrates. Forests have developed on recently abandoned agricultural lands.

32 Montane wet alluvial shrubland Montane wet shrublands and open forests (woodland) on small patches of alluvial material in the upper central

and woodland and Luquillo Mountains. Developing on very recently abandoned agricultural lands.

33 Mature secondary montane wet Mature secondary wet evergreen forests on volcanic substrates in the upper central and Cayey Mountains. These

noncalcareous evergreen forest are on areas that were not likely shade coffee but may have had been cleared for other reasons.

43 Montane wet evergreen A mix of active sun coffee plantations and abandoned shade coffee plantations in the upper elevations of the

abandoned and active coffee central mountains.plantation
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Table 26—Descriptions of landcover types in Puerto Rico (continued)Value Land cover type General description

34 Mature primary and secondary Mature primary tabonuco forests in the Luquillo Mountains and other mature secondary forests in the central

montane wet noncalcareous mountains. This is a closed broad-leaved forest occurring in the subtropical wet Holdridge life zone (Ewel &evergreen tabonuco forest Whitmore 1973.

35 Mature primary and secondary Mature primary palo colorado tall cloud forests in the Luquillo Mountains and other mature cloud secondary

montane wet noncalcareous forests in the central mountains. Occurs in the lower montane wet Holdridge life zone and is considered theevergreen palo colorado cloud zonal vegetation of this life zone (Ewel and Whitmore 1973).forest

36 Mature primary and secondary Mature primary sierra palm forests in the Luquillo Mountains and other mature secondary forests in the central

montane wet noncalcareous mountains. This is a closed broad-leaved evergreen forest (Weaver 1991, 1994) and classified within theevergreen sierra palm forest montane subtropical rain forest formation. The Sierra palm forest type is found in the lower montane wet and

subtropical rain Holdridge life zones (Ewel & Whitmore 1973) on azonal (although common) steeply slopingsites.

37 Mature primary and secondary Mature primary elfin woodland in the Luquillo Mountains and other mature secondary cloud forests in the

montane wet noncalcareous central mountains. This is a closed broad-leaved evergreen forest (Ahern et al. 1999) classified in the elfinevergreen elfin woodland cloud woodland forest type or alliance (Weaver 1994). The community is found within both the lower montane wetforest and rain Holdridge life zones (Ewel & Whitmore 1973) on azonal sites including exposed slopes and ridges

(Weaver 1990, 1991,1994; Weaver et al. 1986).

38 Young secondary montane Young secondary wet evergreen forests on volcanic substrates in the upper Luquillo, central, and Cayey

wet noncalcareous evergreen mountains. Regenerating on recently (10 to 25 years) abandoned pastures.forest

39 Montane wet noncalcareous Montane wet shrubland and open woodland on volcanic substrates in the upper Luquillo, central, and Cayey

evergreen shrubland and woodland mountains. Regenerating on abandoned or semi-active pastures or disturbed areas (landslides, hurricane, forest

clearing).

40 Mature secondary montane wet Mature secondary wet evergreen forests and some forest plantations on ultramafic (high pH) serpentine substrates

serpentine evergreen forest in the Maricao area of the central mountains.

41 Young secondary montane wet Young secondary wet evergreen forests on ultramafic (high pH) serpentine substrates in the Maricao area of the

serpentine evergreen forest central mountains. Regenerating on recently abandoned (10 to 25 years) agricultural land.

42 Wet serpentine shrubland and Montane wet shrubland and open woodland on ultramafic (high pH) serpentine substrates in the Maricao area of

woodland the central mountains. Regenerating on abandoned or semi-active pastures or disturbed areas (landslides,

hurricane, forest clearing).

44 Mangrove forest and shrubland Flooded forest and shrublands in coastal areas periodically inundated with saline tide waters of the Caribbean

and Atlantic oceans. Mangroves may be riparian, estuarine, or littoral, typically with some freshwater inputs aswell as connectivity with the ocean. May be very old and stable communities (up to 4,000 years), or relativelyyoung communities invading land formerly drained for agriculture.
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GENERAL TECHNICAL REPORT IITF-GTR-39 Table 26—Descriptions of landcover types in Puerto Rico (continued) Value Land cover type General description 45 Freshwater Pterocarpus swamp Flooded freshwater swamps on the coastal plain and along riparian areas. It occurs in relict patches within the Lowland subtropical moist Holdridge life zone (Ewel and Whitmore 1973) and was likely much more extensive on the coastal plain and riparian corridors prior to deforestation for agriculture (Alvárez 1982, Eusse and Aide 1999). It is found on undisturbed freshwater seasonally flooded noncalcareous alluvial substrates. 48 Dry grasslands and pastures Active and abandoned pastures in the subtropical dry Holdridge life zone on the southwestern side of Puerto Rico and the smaller islands. Maintained as grassland by fire and grazing. 49 Dry cactus grassland and Rocky, often calcareous, slopes on the southwest side of the island and the Mona plateau. shrubland 50 Moist grasslands and pastures Active and abandoned pastures in the moist and wet Holdridge life zones on the north and central areas of Puerto Rico. Maintained as grassland by fire, grazing, or other disturbances.

53 Seasonally flooded herbaceous Grasslands in lowland areas of the coastal plain frequently flooded by winter and spring rains and overflowing

nonsaline wetlands rivers. Often used for pasture or hay when not flooded.

54 Seasonally flooded herbaceous Grasslands in lowland areas of the coastal plain frequently flooded by saline and brackish water during storms.

saline wetlands Often adjacent to mudflats and mangroves but with some drainage and often drying out.

51 Emergent herbaceous nonsaline Mixed herbs or grasses emerging from brackish water in perennially flooded swamp lands. May included floating

wetlands mats of herbaceous vegetation.

52 Emergent herbaceous saline Mixed herbs or grasses emerging from freshwater in perennially flooded swamp lands. Often adjacent to mudflats

wetlands and mangroves or developing on disturbed saline swamp lands.

55 Hay and row crops Hay crops, pineapple, bananas, plantains, or other herbaceous row crops.56 Woody agriculture and plantations Primarily coconut plantations but may include citrus, mango, pine, and mahogany plantations.57 Rocky cliffs and shelves Coastal cliffs and rocky shelves and shorelines in the intertidal and supratidal zones. Typically < 20 percent

vegetation cover, vegetation halophytic. Often periodically flooded with storm surge.

58 Gravel beaches and stony Stoney and gravelly shorelines in the intertidal and supratidal zones along the coast. Typically < 20 percent

shoreline vegetation cover, vegetation halophytic. Often periodically flooded with storm surge.

59 Fine to coarse sandy beaches, Fine to coarse sandy beaches, mixed sand and gravel beaches in the intertidal and supratidal zones along the

mixed sand and gravel beaches coast. Typically < 20 percent vegetation cover, vegetation halophytic. Often periodically flooded with storm

surge. Some beaches subject to artificial maintenance.

60 Riparian and other natural barrens Typically < 20 percent vegetation cover, often periodically flooded if riparian.61 Salt and mudflats Typically < 20 percent vegetation cover, often periodically flooded. Salt flats in low-lying coastal areas

seasonally flooded with brackish water, part of a mangrove habitat complex when associated with mangrovetrees.

65 Artificial barrens Recent developments cleared for construction (ephemeral, likely to have changed within recent years), landfills,

and quarries.
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Table 26—Descriptions of landcover types in Puerto Rico (continued)Value Land cover type General description

62 Salt production Commercial salt production.63 High-density urban development Developed pixels that exist within a matrix of > 20 percent developed pixels (< 80 percent vegetated, water or

natural barrens) within a 1-km2 area surrounding each pixel (Martinuzzi et al. 2007).

64 Low-density urban development Developed pixels that exist within a matrix of < 20 percent developed pixels (> 80 percent vegetated, water or

natural barrens) within a 1-km2 area surrounding each pixel (Martinuzzi et al. 2007).

66 Freshwater Lakes, ponds and rivers that are detectable in Landsat imagery. A number of small streams, small ponds, and

ephemeral ponds and streams are not included.

67 Saline water Coastal lagoons, brackish canals, and estuaries. Often bordered by mangrove and mudflats. These form part of

the mangrove habitat complex when adjacent to mangroves.

70 Aquaculture Shrimp farms.
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Glossary

algorithm—A procedure to solve a problem or model a solution. In the gap

analysis program (GAP) this typically refers to a geographic information system

(GIS) procedure used to model animal distributions.

alliance level—A land unit made up of a group of natural communities that have

the same dominant or co-dominant plant species or, in the absence of vegetation, by

the dominant land cover typically described according to the Anderson land cover

classification (Grossman et al. 1994).

alpha diversity—A single within-habitat measure of species diversity regardless of

internal pattern, generally over an area of 0.1 to 1000 ha (Whittaker 1960, 1977).

anthropogenic—Caused by humans.

assemblages—A group of ecologically interrelated plant and animal species.

band, spectral—A segment of the electromagnetic spectrum defined by a range of

wavelengths (e.g., blue, green, red, near infrared, far infrared) that compose the

satellite imagery (e.g., Landsat ETM+ imagery).

beta diversity—The change in species diversity among different natural communities of a landscape; an index of between-habitat diversity (Whittaker 1960, 1977).

biodiversity—Generally, the variety of life and its interrelated processes.

biogeographic—Relating to the geographical distribution of plants and animals

biological diversity—See biodiversity.

cartographic—Pertaining to the art or technique of making maps or charts.

coarse filter—The general conservation activities that conserve the common

elements of the landscape matrix, as opposed to the “fine filter” conservation

activities that are aimed at special cases such as rare elements (Jenkins 1985).

community—A group of coexisting plants and animals.

cover type—A nontechnical high-level floristic and structural description of

vegetation cover.

cross-walking—Matching equivalent land cover categories between two or more

classification systems.

delineate—Identifying the boundaries between more or less homogenous areas; for

remotely sensed images it is based on detectable differences in tone and texture.
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delta diversity—The change in species diversity between landscapes along major

climatic or physiographic gradients (Whittaker 1977).

digitization—Entering spatial data digitally into a geographic information system.

ecoregion—A large region, usually spanning several million hectares, characterized

by having similar biota, climate, and physiography (topography, hydrology, relief).

ecosystem—A biological community, its physical environment, and the processes

through which matter and energy are transferred among the components.

edge-matching—The process of connecting polygons at the boundary between two

independently created maps, either between thematic mapper scenes or between

state GAP data sets.

element—A plant community or animal species mapped by GAP. May also be

referred to as “element of biodiversity.”

error of commission—The occurrence of a species (or other map category) is

erroneously predicted in an area where it is, in fact, absent.

error of omission—Failure of a model to predict the occurrence of a species that is

actually present in an area.

extinction—Disappearance of a species throughout its entire range.

extirpation—Disappearance of a species from part of its range.

fine filter—See “coarse filter.”

floristic—Pertaining to the plant species that make up the vegetation of a given

area.

formation level—The level of land cover categorization between “group” and

“alliance” describing the structural attributes of a land unit, for example, “evergreen coniferous woodlands with rounded crowns” (Jennings 1993).

gamma diversity—The species diversity of a landscape, generally covering 1000

to 1 000 000 ha, made up of more than one kind of natural community (Whittaker

1977).

gap analysis—A comparison of the distribution of elements of biodiversity with

that of areas managed for their long-term viability to identify elements with

inadequate representation.

geographic information systems (GIS)—Computer hardware and software for

storing, retrieving, manipulating, and analyzing spatial data.
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GENERAL TECHNICAL REPORT IITF-GTR-39

global positioning system (GPS)—An instrument that uses satellite signals to

pinpoint its location on the Earth’s surface.

ground truthing—Verifying maps by checking the actual occurrence of plant and

animal species in the field at representative sample locations.

habitat—The physical structure, vegetation composition, and physiognomy of an

area, the characteristics of which determine its suitability for particular animal or

plant species.

hectare—A metric unit of area of 10 000 m2

 and equal to 2.47 acres.

metadata—Information about data, e.g., their source, lineage, content, structure,

and availability.

minimum mapping unit—The smallest area that is depicted on a map.

Neotropics—The biogeographic region stretching southward from the tropic of

Cancer and including southern Mexico, Central and South America, and the West

Indies.

phenology—The study of periodic biological phenomena, such as flowering,

breeding, and migration, especially as related to climate.

phenotype—The environmentally and genetically determined observable appearance of an organism, especially as considered with respect to all possible genetically influenced expressions of one specific character.

physiognomic—Based on physical features.

physiographic province—A region having a pattern of relief features or land

forms that differ significantly from that of adjacent regions.

pixel—The smallest spatial unit in a raster data structure.

polygon—An area enclosed by lines in a vector-based GIS data layer or a region of

contiguous homogeneous pixels in a raster system.

preprocessing—Those operations that prepare data for subsequent analysis, usually

by attempts to correct or compensate for systematic, radiometric, and geometric

errors.

range—The geographic limit of the species.

range unit—A spatial, geographic unit to record and display species geographic

range.
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reach—A stream or river segment between inflowing tributaries.

registration, spatial—Matching different images to each other by finding points

on the images that can be matched to known points on the ground.

remote sensing—Deriving information about the Earth’s surface from images

acquired at a distance, usually relying on measurement of electromagnetic radiation

reflected or emitted from the feature of interest.

resolution, spatial—The ability of a remote sensing system to record and display

fine detail in a distinguishable manner or the smallest feature that can be distinguished or resolved on a map or image, such as a thematic mapper pixel.

scale, map—The ratio of distance on a map to distance in the real word, expressed

as a fraction; the smaller the denominator, the larger the scale, e.g. 1:24,000 is

larger than 1:100,000.

species richness—The number of species of a particular interest group found in a

given area.

supervised classification—The process of classifying thematic mapper pixels of

unknown identity by using samples of known identity (i.e., pixels already assigned

to informational classes by ground truthing or registration with known land cover)

as training data.

synoptic—Constituting a brief statement or outline of a subject; presenting a

summary.

tessellation—The division of a map into areas of equal and uniform shape such as

the EPA-EMAP hexagon.

Thematic Mapper—A sensor on LANDSAT 4 and 5 satellites that records information in seven spectral bands, has a spatial resolution of about 30 by 30 m, and

represents digital values in 256 levels of brightness per band.

universal transverse mercator—One of several map projections or systems of

transformations that enables locations on the spherical Earth to be represented

systematically on a flat map.

universal transverse mercator grid—A geographic reference system used as the

basis for worldwide locational coding of information in a GIS or on a map.
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GENERAL TECHNICAL REPORT IITF-GTR-39

unsupervised classification—The definition, identification, labeling, and mapping

of natural groups, or classes, of spectral values within a scene. These spectral

classes are reasonably uniform in brightness in several spectral channels.

vector format—A data structure that uses polygons, arcs (lines), and points as

fundamental units for analysis and manipulation in a GIS.

wildlife habitat relationship model—A method of linking patterns of known

habitat use by animal species with maps of existing vegetation, thereby identifying

the spatial extent of important habitat features for use in conservation and management
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Glossary of Acronyms

ACSM American Congress on Surveying and Mapping

ADAMAS Aquatic Database Management System

ADEM Alabama Department of Environmental Management

AML ARC/INFO Macro Language

ASPRS American Society for Photogrammetry & Remote Sensing

AVHRR Advanced Very High Resolution Radiometer (satellite

system)

BBS Breeding Bird Survey

BEST Biomonitoring of Environmental Status and Trends

BLM Bureau of Land Management

BRD Biological Resources Division of the USGS

CAFF Conservation of Arctic Flora and Fauna

C-CAP Coastwatch Change Analysis Program (NOAA)

CDC Conservation Data Center

CEC Council on Environmental Cooperation

CENR Committee on Environment and Natural Resources

CERES California Environmental Resources Evaluation System

CIESIN Consortium for International Earth Science Information

Network

CODA Conservation Options and Decision Analysis (software)

CRMP Coordinated Resource Management Plan

CRUC Cooperative Research Unit Center

DLG-E Digital line graph–enhanced

DNER Department of Natural and Environmental Resources

DOI Department of the Interior

EDC EROS Data Center
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GENERAL TECHNICAL REPORT IITF-GTR-39

ECOMAP The National Hierarchical Framework of Ecological Units

mapping project of the USDA Forest Service

EMAP Environmental Monitoring & Assessment Program

EMAP-LC EMAP-Landscape Characterization (USEPA)

EMSL Environmental Monitoring & Systems Laboratory

(USEPA)

EMTC Environmental Management Technical Center (NBS)

EOS Earth Observing System

EOSAT Earth Observation Satellite Company (the commercial

operator of the Landsat satellite system)

EOSDIS EOS Data and Information System

ERDAS Imagery processing software

ERL Environmental Research Laboratory, Corvallis (USEPA)

EROS Earth Resources Observation Systems (USGS)

ESRI Environmental Systems Research Institute

ETM+ Enhanced Thematic Mapper plus

FGDC Federal Geographic Data Committee

FIA Forest Inventory and Analysis

FIPS Federal Information Processing Standard

FTP file transfer protocol

FY Fiscal Year

GAO General Accounting Office (Congress)

GAP Gap Analysis Program

GCDIS Global Change Data and Information System

GIS Geographic information system

GLIS Global Land Information System (USGS)
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GLOBE Global Learning and Observations to Benefit the

Environment

GPS Global positioning system

GRASS Geographic Resources Analysis Support System

GRIS Geographic Resource Information Systems

HRMSI High-Resolution Multispectral Stereo Imager

IALE International Association of Landscape Ecology

IDRISI GIS developed by Clark University

IITF International Institute of Tropical Forestry

IKONOS Commercial Earth observation satellite

LAPS Land Acquisition Priority System

LC/LU Land cover/land use (USGS)

MIPS Map and Image Processing System

MOU Memorandum of understanding

MMU Minimum mapping unit

MRLC Multi-Resolution Land Characteristics Consortium

MSS Multi-spectral scanner

MTPE Mission to Planet Earth

NAD North American Datum

NAFTA North American Free Trade Agreement

NALC North American Landscape Characterization

(USEPA, USGS)

NAWQA National Water Quality Assessment (USGS)

NBII National Biological Information Infrastructure

NBS National Biological Service

NCCP Natural Communities Conservation Planning program

(in CA)
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GENERAL TECHNICAL REPORT IITF-GTR-39

NDCDB National Digital Cartographic Data Base

NDVI Normalized Difference of Vegetation Index

NERC National Ecology Research Center (Fort Collins, CO)

NMD National Mapping Division

NOAA National Oceanic and Atmospheric Administration

NPS National Park Service

NSDI National Spatial Data Infrastructure

NSTC National Science and Technology Council

NVCS National Vegetation Classification System

NWI National Wetlands Inventory (USFWS)

OMB Office of Management and Budget (Administration)

OSIS Oregon Species Information System

PA Protected area

PARC Public Access Resource Center

PI Principal investigator

PRGAP Puerto Rico Gap Analysis Project

PRUSVIGAP Puerto Rico and the United States Virgin Islands Gap

Analysis Project

SAB Science Advisory Board (USEPA)

SCICOLL Scientific Collections Permit Database

SDTS Spatial data transfer standard

SGID State Geographic Information Database

SMMR Scanning Multichannel Microwave Radiometer

SNEP Sierra Nevada Ecosystem Project

SOPI Sociedad Ornitológica Puertorriqueña

SOFIA Southern Forest Inventory and Analysis
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SPOT Systáme Pour l’Observation de la Terre

RMSE Root mean square error

TIGER Topologically Integrated Geographic Encoding and

Referencing system (used for U.S. census)

TM Thematic Mapper

TNC The Nature Conservancy

UA Urban area

UNESCO United Nations Educational, Scientific, and Cultural

Organization

URISA Urban and Regional Information Systems Association.

URL Universal resource locator

USEPA U.S. Environmental Protection Agency

USGS U.S. Geological Service

USFS U.S. Forest Service

USFWS U.S. Fish & Wildlife Service

UTM Universal Transverse Mercator

WHRM Wildlife/habitat relationship model
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