U.S. Fish and Wildlife 

Service Refuges

And Other Nearby Reserves 

In Southwestern Puerto Rico

Peter L. Weaver and Joseph J. Schwagerl

United States

Department of

Agriculture

Forest Service

International Institute 

of Tropical Forestry

General Technical

Report IITF-40
Front cover

Top, Cabo Rojo National Wildlife Refuge. The Cabo Rojo Salt flats, critical wildlife habitat 

at the southwestern tip of Puerto Rico, boast a long history of wildlife use and human activity 

(Photo by Jorge Salivia). Bottom, Laguna Cartagena National Wildlife Refuge. Farming and 

water diversion measures around the Cartagena Lagoon in southwestern Puerto Rico have 

modified its water quality and regimen (Photo by Peter L. Weaver) .

Top, Wildlife refuge. Welcome to the U.S. Fish and Wildlife Service’s Cabo Rojo National 

Wildlife Refuge via Mariano Rodríguez Lane (Photo by Peter L. Weaver). Bottom, Headquarters. 

First constructed to monitor regional communications, the original headquarters was the 

home of the U.S. Fish and Wildlife Service for 35 years (Photo by Peter L. Weaver). 
U.S. Fish and Wildlife 

Service Refuges

And Other Nearby Reserves 

In Southwestern Puerto Rico

Peter L. Weaver

International Institute of Tropical Forestry

U.S. Department of Agriculture Forest Service

Río Piedras, Puerto Rico

and

Joseph J. Schwagerl

U.S. Fish and Wildlife Service

U.S. Department of the Interior

Cabo Rojo, Puerto Rico

June 2009

International Institute of Tropical Forestry

Jardín Botánico Sur

1201 Calle Ceiba

San Juan, PR 00926-1119

iii

Contents

 Page

Abstract ............................................................................................................................... v

List of Figures ..................................................................................................................... vi

List of Tables ....................................................................................................................... vii

Background Information ................................................................................................. 1

Introduction ........................................................................................................................ 1

Objectives and Parameters .................................................................................................. 1

Administrative Details ........................................................................................................ 2

Historical Perspective ......................................................................................................... 2

Environmental Conditions of the Refuges ................................................................... 6

Geology .............................................................................................................................. 6

Soils ................................................................................................................................... 9

Physiography and Hydrology ............................................................................................. 13

Climate ................................................................................................................................ 14

Flora ...................................................................................................................................... 17

Major Forest Types in the Southwest ................................................................................. 18

Other Vegetation Types ....................................................................................................... 19

Vegetation Types at the Refuges ......................................................................................... 19

Fauna on the Refuges ....................................................................................................... 21

Mammals ............................................................................................................................ 21

Birds .................................................................................................................................. 25

Reptiles and Amphibians .................................................................................................... 35

Fish ..................................................................................................................................... 36

Other Flora and Fauna ........................................................................................................ 37

Environmental Issues ....................................................................................................... 37

Endangered Species .......................................................................................................... 39

Management Activities ..................................................................................................... 41

Fire and Habitat Restoration ............................................................................................... 41

Cartagena Lagoon ............................................................................................................... 46

Structures and Improvements ............................................................................................. 48

Collaborative Activities ...................................................................................................... 50

Public Use of Refuges ....................................................................................................... 51

Visitors ................................................................................................................................ 51

Special Use permits ........................................................................................................... 52

Other Major Protected Areas in the Southwest .......................................................... 52

Boquerón Commonwealth Forest ....................................................................................... 52

Boquerón Wildlife Refuge .................................................................................................. 55

Guánica Commonwealth Forest ......................................................................................... 57
iv

 Page

Los Morrillos de Cabo Rojo Lighthouse ............................................................................ 58

Parguera Natural Reserve ................................................................................................... 59

Punta Guaniquilla Natural Reserve .................................................................................... 60

General Observations and Research in Southwestern Puerto Rico ....................... 61

Flora—Mangroves and Phenology ..................................................................................... 62

Fauna—Birds ...................................................................................................................... 62

Cabo Rojo National Wildlife Refuge Studies—Headquarters Tract 

and Municipality ................................................................................................................ 62

Flora—Tree Genetics, Mistletoe, and Restoration ............................................................. 62

Fauna—Birds ...................................................................................................................... 63

Other Fauna ........................................................................................................................ 64

Cabo Rojo National Wildlife Refuge Studies—Salinas Tract ................................. 64

Lagoon System and Its Wildlife ......................................................................................... 64

Fauna—Shorebirds and Wading Birds ................................................................................ 65

Fauna—Sea Turtles ............................................................................................................. 66

Laguna Cartagena National Wildlife Refuge Studies—Lagoon Tract .................. 66

Water Quality, Insects, and Earthworms ............................................................................. 66

Native Bird Species ............................................................................................................ 66

Laguna Cartagena National Wildlife Refuge Studies—Tinaja Tract ................... 67

Secondary Succession and Restoration .............................................................................. 67

Boquerón Commonwealth Forest .................................................................................. 67

Fauna—Bird Species .......................................................................................................... 67

Guánica Commonwealth Forest .................................................................................... 69

Climatic Influences ............................................................................................................. 69

Flora—Forest and Tree Species .......................................................................................... 69

Fauna—Bird Monitoring Studies ....................................................................................... 70

Fauna—Individual Bird Species ......................................................................................... 71

Fauna—Lizards and Geckos ............................................................................................... 72

Future Activities ................................................................................................................ 73

Acknowledgments ............................................................................................................. 75

Endnotes .............................................................................................................................. 75

Literature Cited .................................................................................................................. 77

Appendix A—Alphabetical listing by scientific name of all tree species that 

were mentioned in the text or tables, and of other plant species mentioned only 

in text. Ecological information is provided only for tree species .......................................... 89

Appendix B—Plant species at the Cabo Rojo and Laguna Cartagena National 

Wildlife Refuges .................................................................................................................... 95

Contents (continued)
v

Abstract

The main purpose of this paper is to summarize information about the Cabo Rojo and 

Laguna Cartagena National Wildlife Refuges in southwestern Puerto Rico. The Cabo 

Rojo (Headquarters and Salinas tracts) and Laguna Cartagena (Lagoon and Tinaja tracts) 

occupy 1174 ha and are part of the 10 231-ha Caribbean system of nine refuges. Their 

major geologic and physiographic features are Sierra Bermeja, the oldest mountain range 

in Puerto Rico, and the floodplains of the Lajas Valley. Together, the refuges combined 

contain 9 geologic features and 26 soils’ types. Rainfall averages about 1000 mm/year 

and hurricanes occur periodically. Among the several vegetation types, including are 

mangroves, salt flats, littoral woodland, mesquite and semievergreen woodland, coastal 

shrub, deciduous woodland, and pasture. At least 644 plant species grow on the refuges. 

Eight of these species are considered rare and endangered, and have formal recovery plans. 

Refuge mammals include two species of native bats and six introduced species—two 

monkeys, a mongoose, two rats, and a mouse. Both the Cabo Rojo Salt Flats and Cartagena 

Lagoon are renowned as wintering habitat for migratory birds where >200 resident and 

migratory birds have been observed, including 12 island endemics. Moreover, several other 

species are listed as rare and endangered, or of conservation concern, or as target species 

for management. Among the reptiles are seven lizards, two geckos, and three turtles (one 

terrestrial and two marine). Five amphibians (one toad and four frogs) and 18 species of 

fish (fresh and brackish water) are also present. Six of the wildlife species have formal 

recovery plans. 

The major management activities on the refuges include protection against fires, habitat 

restoration, public education, special uses, and water management at the Cartagena Lagoon 

and the Salinas salt flats. Habitat recovery has been stimulated through fire control, through 

livestock removal, and by planting >9,000 trees of nearly 80 species. Public education 

encompasses nearly 20,000 visits annually to interpretative centers at the Headquarters 

and Salinas tracts; moreover, another 2,600 visitors frequent the refuges to observe 

wildlife and take nature photographs, and nearly 4,900 others participate in educational 

and interpretative programs. Special use permits have been issued to 174 users since 1979, 

mostly for administrative matters or environmental research. Finally, water management at 

the Cartegena Lagoon remains a problem, partly because of historical events and partly due 

to current difficulties. 

In addition to the U.S. Fish and Wildlife Service (F&WS) refuges, six other major 

protected areas occupy about 6800 ha along the southwestern coast: Punta Guaniquilla 

Natural Reserve, Boquerón Wildlife Refuge, Los Morrillos de Cabo Rojo, Boquerón 

Forest, Parguera Natural Reserve, and Guánica Forest. An additional 23 bird species not 

recorded on the refuges have been observed in these protected areas. The F&WS refuges 

and the protected areas provide critical wildlife habitat in Puerto Rico’s southwest—an 

area that is experiencing a rapid increase in human population. 

Keywords: Cabo Rojo Wildlife Refuge, Laguna Cartagena Wildlife Refuge, fauna, flora, 

climate, geology, soils, management.
vi

List of Figures 

Figure 1—Places mentioned within southwestern Puerto Rico in text .............................. 5

Figure 2—Population in the Cabo Rojo municipality: 1750–2025 (Junta de 

Planificación 1995, 2005; Ramos y Ramírez de Arellano and others 1985) ........................ 5

Figure 3—Geology and topography map of the Cabo Rojo National Wildlife 

Refuge (Bawiec 2001, Volckmann 1984a)............................................................................ 8

Figure 4—Geology and topography map of the Laguna Cartagena National 

Wildlife Refuge (Bawiec 2001, Volckmann 1984a) ............................................................ 10

Figure 5—Soils map of the Cabo Rojo National Wildlife Refuge (Carter 1965).............. 11

Figure 6—Soils map of the Laguna Cartagena National Wildlife Refuge 

(Carter 1965)........................................................................................................................ 12

Figure 7—Rainfall and temperature diagram (1971–2000): (A) Lajas, 

(B) Magueyes....................................................................................................................... 15

Figure 8—Hurricane trajectories over Puerto Rico since 1700 ......................................... 16

Figure 9—Christmas bird counts: (A) number of species, (B) number of individual 

birds, and (C) number of participants. ................................................................................. 35

Figure 10—Tree plantings by refuge tract and year since 1980........................................ 45
vii

List of Tables

Table 1—U.S. Fish and Wildlife Service refuges in the Caribbean Basin and 

Commonwealth protected areas in southwestern Puerto Rico ............................................. 3

Table 2—Geology and soils on the Headquarters and Salinas tracts of the Cabo 

Rojo National Wildlife Refuge ............................................................................................ 7

Table 3—Geology and soils on the Lagoon and Tinaja tracts of the Laguna 

Cartagena National Wildlife Refuge .................................................................................... 9

Table 4—Mean annual rainfall at the U.S. Fish and Wildlife Service refuges, 

1991–2005 ............................................................................................................................ 14

Table 5—Animal species other than birds at the Cabo Rojo and Laguna 

Cartagena National Wildlife Refuges, and Commonwealth protected areas in 

southwestern Puerto Rico .................................................................................................... 22

Table 6—Bird species at the Cabo Rojo and Laguna Cartagena National Wildlife 

Refuges and Commonwealth protected areas in southwestern Puerto Rico ........................ 26

Table 7—Butterfly species found on the Cabo Rojo and Laguna Cartagena 

National Wildlife Refuges ................................................................................................... 38

Table 8—U.S. Fish and Wildlife Service species recovery plans applicable to 

species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges ...................... 40
viii

List of Tables (continued)

Table 9—Chronology of activities that influenced the Cabo Rojo National 

Wildlife Refuge (Headquarters and Las Salinas National Wildlife tracts) .......................... 42

Table 10—Tree species planted at the Cabo Rojo and Laguna Cartagena 

National Wildlife Refuges, 1980–2003 ................................................................................ 43

Table 11—Chronology of activities that influenced the Laguna Cartagena 

National Wildlife Refuge (Lagoon and Tinaja tracts) ......................................................... 47

Table 12—Structures and improvements at the Cabo Rojo and Laguna 

Cartagena National Wildlife Refuges ................................................................................ 49

Table 13—Special use permits for the Cabo Rojo and Laguna Cartagena 

National Wildlife Refuges .................................................................................................. 53
Background Information

Introduction

The U.S. Fish and Wildlife Service (F&WS) in 

the U.S. Department of the Interior administers the 

national wildlife refuge system, which grew from a 

single 1.2-ha refuge in 1903 to 390 refuges totaling 

137380 km2

 by 1979 (Doyle 1979). The first refuge, 

Pelican Island in eastern Florida, was set aside by 

President Theodore Roosevelt to protect the brown 

pelican. In 1909, when Roosevelt left office, the 

wildlife refuge system had 51 reservations in 21 States 

and 3 territories (Clark 1993). When the Salinas tract 

was added to the Cabo Rojo National Wildlife Refuge 

(NWR) in October 1999, wildlife refuges numbered 

more than 500 and occupied nearly 380000 km2 

(Meretsky and others 2006 ; U.S. Fish and Wildlife 

Service 1999, see endnotes). Today, refuges are 

scattered in all States throughout the continental 

United States and in Alaska, Hawaii, Puerto Rico, 

the U.S. Virgin Islands (Ogburn 1979), and Navassa 

Island near Haiti’s southwestern shores (U.S. Fish and 

Wildlife Service 2002).

Initially, the purpose of the refuge system was 

to protect wildlife, especially migratory waterfowl, 

from the devastating impacts of a growing society 

(Doyle 1979). Because migratory bird species move 

across State, provincial, and national borders, the 

Migratory Bird Treaty Act of 1918 recognized them 

as an international resource. Management of the 

wildlife and their habitats are the responsibility of 

the Secretary of the Interior. In 1924, a provision 

allowed for hunting and fishing. In 1934, during the 

Great Depression, an effort was made to purchase 

and restore habitats. Moreover, Congress passed the 

Migratory Bird Hunting and Conservation Stamp Act, 

which was, in essence, a tax on those who hunted 

migratory waterfowl. In 1956, the Fish and Wildlife 

Act expanded the role of refuges beyond waterfowl to 

all endangered wildlife. At the same time, the refuges 

faced pressure for greater public use, including 

recreation. In 1973, Congress passed the Endangered 

Species Act, subsequently amended, to protect 

endangered and threatened species. Today, refuge 

lands are mainly used for environmental education, 

fishing, wildlife observation, hiking, and photography 

rather than hunting. 

The stated mission of the F&WS is “to administer 

a national network of lands and waters for the 

conservation, management, and where appropriate, 

restoration of the fish, wildlife and plant resources and 

their habitats within the United States for the benefit 

of present and future generations of Americans” 

(Meretsky and others 2006). Since many important 

migratory bird habitats in the eastern Caribbean have 

been lost through development, refuges in Puerto 

Rico and the Virgin Islands have become increasingly 

important in the conservation of migratory wildlife.

Objectives and Parameters

The purpose of this report is to compile 

information on the Cabo Rojo and Laguna 

Cartagena NWRs into a convenient summary to

 serve as a baseline for future research and 

management activities. This includes all known 

vascular plants and vertebrate animal species tallied 

within the refuges. The topics include administrative 

details; history; research, i.e., environmental data, 

flora, and fauna; management; and public use. This 

review of research topics highlights the importance 

of the refuges in the conservation of dry forest habitat 

and its wildlife, in particular the avifauna of Puerto 

Rico’s southwest. Studies conducted within the 

refuges have been mentioned or briefly summarized 

along with relevant work carried out at nearby sites, 

including the Commonwealth forest reserve units 

at Boquerón and Guánica, the Boquerón Wildlife 

Refuge, the Parguera and Punta Guaniquilla Natural 

Reserves, and Los Morrillos de Cabo Rojo (Cabo 

Rojo Lighthouse area) (Departamento de Resursos 

Naturales 1979, Ventosa-Febles and others 2005). 

Inclusion of other sites, notably Guánica, is important 

because of the amount of relevant research that has 

been carried out there. Environmental issues and 

suggestions for future research and management 

activities have been outlined. 

The Cartagena Lagoon, on the Laguna Cartagena 

NWR wildlife refuge, has a long record of 

human activity, and many observations have been 

summarized in unpublished reports over the years. 

This report does cite some gray literature, i.e., 

unpublished lists of plants and birds, trip reports, 

short-term field investigations, and student projects, 

mainly related to the refuge sites. Lists of nonvascular 
2

plants and invertebrates, or details of water 

management for the Cartagena Lagoon, however, are 

topics beyond the scope of this review. Finally, the 

text relies heavily on the use of scientific names for 

the flora and fauna. To facilitate species identification, 

common names have been tabulated in English and 

Spanish. Also, English common names have been 

included when bird species are first mentioned 

in text or when they are discussed as separate 

research projects. 

Administrative Details

The F&WS manages nine refuges in the Caribbean 

Basin that total about 10230 ha of terrestrial habitat 

(table 1). Puerto Rico and its offshore islands contain 

five of the refuges; the U.S. Virgin Islands, three; 

and Navassa Island, about 60 km west of Haiti, is the 

remaining refuge. The refuges range in size from 6 ha 

on Green Cay to 7 566 ha on Vieques. The two most 

distant, Navassa and Sandy Point, are separated by 

1100 km; all except Navassa, however, are within 250 

km of the Headquarters tract at Cabo Rojo. 

The Cabo Rojo and the Laguna Cartagena NWRs, 

situated in southwestern Puerto Rico, together occupy 

1174 ha (Ventosa-Febles and others 2005) (fig. 1). 

The Headquarters and Salinas (salt flats) tracts are 

in the Cabo Rojo NWR, which also includes the 

current F&WS headquarters in the Caribbean. The 

Headquarters tract was transferred from the Foreign 

Broadcast Information Service, i.e., the Central 

Intelligence Agency, in June 1974 to help implement 

the national migratory bird program according to 

16 U.S.C. 667b, an act authorizing the transfer of 

real property for wildlife or other purposes. The 

Salinas tract, numerically the most important area for 

shorebirds in Puerto Rico and the Caribbean region 

(Collazo and others 1995, see endnotes), was acquired 

in 1999 to protect important shore bird habitat.

The Laguna Cartagena NWR includes the Lagoon 

and Tinaja tracts. The six parcels in the Lagoon tract, 

previously owned by four individuals, were appraised 

in 1985 (Waight 1985, see endnotes). The Lagoon 

tract, established through a lease agreement with the 

Commonwealth of Puerto Rico in August of 1989, 

was for the conservation and management of fish and 

wildlife resources according to 16 U.S.C. 742f(a)(4), 

and for the benefit of the F&WS (Earsom and others 

2004). The Tinaja tract was acquired in 1996 via a fee 

title from the U.S. Department of Agriculture Farm 

Service Agency. About one-half of the Lagoon 

tract is occupied by the Cartagena Lagoon and 

surrounding wetlands. 

The major purpose for acquiring refuge lands in 

Puerto Rico was to assist the F&WS in carrying out 

its mission, which includes the conservation and 

protection of fish and wildlife resources, and carrying 

out the national migratory bird management program. 

Others have provided more specific details (Raffaele 

and others 1977; Schaffner 1995a and 1995b, see 

endnotes; Woodbury and others 1975): 

•	 Increase, improve, and protect nesting and

foraging habitat for native forest birds, shore birds, 

and wetland species, including threatened and 

endangered species.

•	 Provide wintering habitat for neotropical migratory

forest bird species.

•	 Increase native plant diversity and protect

threatened and endangered species. 

•	 Restore fish, wildlife, plant resources, and their

habitats.

•	 Reduce threats from exotic plant and animal

species.

•	 Provide environmental education and interpretative

opportunities for the public. 

Historical Perspective 

Puerto Rico’s southwest, including the salt flats of 

Cabo Rojo and the Lajas Valley, has a long and rich 

history (Ibern Fleytas 1960). Before discovery, Cabo 

Rojo had numerous Indian inhabitants as evidenced 

by shell mounds and excavated sites. The Indians 

survived by collecting snails and other littoral fauna, 

fishing, hunting, and cultivating subsistence crops, 

including yuca. It is unlikely that their activities had a 

major impact on the island’s vegetation although they 

used fire to clear land for their crops. 

Some contend that in November 1493, Columbus 

and his crew first set foot on Puerto Rico in the 

vicinity of the town of Boquerón while looking for 

water (Ibern Fleytas 1960). Most, however, suggest 

it was most probably farther north near Añasco or 

Aguada (Brau 1969). As early as 1511, the Cabo Rojo 

Salt Flats were being exploited and San Germán was 

the first settlement dedicated to marketing salt (GilBermejo García 1970, Sued Badillo 1985). In 1519, 
3

Table 1—U.S. Fish and Wildlife Service refuges in the Caribbean Basin and Commonwealth protected areas in 

southwestern Puerto Rico a

Location and property name

Date(s)

established Area

Range in

elevation

Number of 

species

Plants Habitat description c Birds

ha b m

U.S. Fish and Wildlife Service

refuges in the Caribbean

Puerto Rico’s southwest

Cabo Rojo

Headquarters tract d 1974 238 3–35 245 107 Scattered trees, shrubs, pasture

Salinas tract e 1999 513 0–3 ~147 118 Salt flats, lagoons; mangrove,

upland scrub

Laguna Cartagena

Lagoon tract f 1989 317 11–40 189 146 Freshwater lagoon; grass, 

scattered trees

Tinaja tract g 1996 106 20–300 196 17+ Abandoned pasture and

secondary forest

Subtotal 1974–99 1,174 0–300

Elsewhere in Puerto Rico

Desecheo h 1976 146 0–206 161 51 Secondry forest and scrub,

heavily grazed

Culebra and cays i 1909–2004 607 0–107 410 120 Secondry forest and scrub;

mangrove

Vieques j 2001–2003 7,566 0–301 781 107 Secondary subtropical

moist and dry forest

U.S. Virgin Islands

Buck Island, St. Thomas k 1969/81/04 18 0–33 43+ 10 Secondary scrub

Green Cay, St. Croix l 1977 6 0–19 69 24 Secondary scrub

Sandy Point, St. Croix m 1984/03 170 0–3 225 99 Secondary scrub

Navassa Island n 1999 544 0–60 147 49 Secondary forest

Subtotal 1909–2004 9,057 0–301

Total 1909–2004 10,231 0–301

Reserves of the Commonwealth

in southwestern Puerto Rico

Boquerón Forest o 1918 1,876 0–5 ? >50 Mangrove wetlands, salinas, 

and scrub

Boquerón Wildlife Refuge p 1963 252 0–1 ? 130 Mangrove wetlands, salinas, 

and scrub

Los Morrillos de Cabo Rojo q 1882 105 0–28 147 52 Secondary scrub

Punta Guanaquilla Natural 1976 152 0–46 253 81 Coastal thicket, secondary Reserve r scrub, wetlands

Guanica Forest s 1919 4,016 0–228 500 154 Late secondary subtropical

dry forest

Parguera Natural Reserve t 1979 408 0–95 184 64 Mangrove wetlands, salinas,

and scrub

Total 1882–1980 6,809 0–228

continued
4

Table 1—U.S. Fish and Wildlife Service refuges in the Caribbean Basin and Commonwealth protected 

areas in southwestern Puerto Rico a (continued)

a Except for parts of Vieques, all areas are located in subtropical dry forest. 

b Terrestrial areas (marine areas omitted) rounded to the nearest hectare.

c Flowering plants and ferns.

d Plants (McKenzie 1986, see endnotes); birds (Del Llano and others 1986, U.S. Fish and Wildlife Service 2002).

e Source: Ventosa-Febles and others (2005). Plants (no tally available; probably similar to Los Morillos de Cabo Rojo); birds 

(Collazo and others 1995, U.S. Fish and Wildllife Service 2004). 

f Plants (Diaz-Soltero 1990); birds (U.S. Fish and Wildlife 2004).

g Plants (Proctor 1996, see endnotes; Weaver and Chinea 2003; Weaver and Schwagerl 2004, 2005); birds (U.S. Fish and Wildlife 

Service 2004). 

h Plants (Breckon 2000, Woodbury and others 1971); birds (Anon. 1987, see endnotes; Struthers 1927; Wetmore 1918).

i Plants (Woodbury 1981, see endnotes); birds (U.S. Fish and Wildlife Service 1996); plants and birds (Metzen 1981, 

see endnotes). 

j Plants (Proctor 1994); birds (Sorrie 1975).

k Plants and birds (Lombard 2004, see endnotes).

l Plants (Woodbury and Vivaldi 1982, see endnotes; Joe Schwagerl, personal observation.); birds (Wiley 2003, see endnotes).

m Plants (Anon., no date, see endnotes); birds (U.S. Fish and Wildlife Service 1993a).

n Plants (Zanoni and Buck 1999); birds (Earsom and others, in press). o Source: Department of Natural Resources (1976); Ventosa-Febles and others (2005). Boquerón Forest is divided in 12 separate 

segments located in the municipalities of Lajas, Boquerón, and Mayaguez, as follows (from Bahia Montalva west to Cabo Rojo, 

then north toMayaguez): Bahía Montalva, Pitahaya, Los Molinos, lands near El Faro, Boquerón, Guaniquilla (mangroves at Villa 

Taína), Villa de la Mela mangroves, Puerto Real, Barrio Miradero (Punta Ostiones), Cayo Ratones, Joyuda Lagoon mangroves, 

and Guanajibo mangroves. Plants are mainly typical of mangroves and salinas.

p Source: Ventosa-Febles and others (2005). q Sources: Cerame Vivas (1988); Municipio Autonomo de Cabo Rojo (1998).

r Sources: Fuentes Santiago and Quevedo Bonilla (2002); Vazquez and Kolterman (1998).

s Sources: Canals Mora (1990), Departamento de Recursos Naturales (1985), Quevedo and others (1990), Velez Rodriguez 

(no date).

t Sources: Departamento de Recursos Naturales (1981); Departamento de Recursos Naturales y Ambientales (2000a); VentosaFebles and others (2005).

the Spanish directed that sugar production be pursued 

in Puerto Rico. Other crops that were grown at one 

time or another around Cabo Rojo included sugar 

cane, cacao, coffee, cotton, and tobacco (Gil-Bermejo 

García 1970). 

By 1559, the town of Cabo Rojo was settled. 

Apparently, the settlement was moved from an earlier 

site along the coast where a Spanish vessel was sunk 

by the French; later, the French also attacked San 

Germán. Cofresí (Roberto Couphersein y Ramírez 

de Arellano) was a Puerto Rican pirate born in Cabo 

Rojo in 1791. He attacked foreign, i.e., non-Spanish, 

ships and shared the booty with friends. Subsequently, 

he became a local legend and tales of his character 

and exploits were greatly enhanced. 

In 1898, when Puerto Rico was occupied by the 

United States, the island’s population was about 

800,000 and the dominant industries were agriculture 

and lumbering (Morris 1901). The principal crop was 

coffee, followed by sugar, both accounting for more 

than 90 percent of the exports. Salt works situated 

along the south coast, including those at Cabo Rojo, 

constituted the principal mineral industry. At that 

time, the population of the town of Cabo Rojo was 

about 16,150; since the mid-1970s, the population of 

the municipality has climbed dramatically (Ramos y 

Ramírez de Arellano and others 1985, see endnotes) 

(fig. 2). 

A socioeconomic study carried out during the early 

1940s in the Lajas Valley region, i.e., from Boquerón 

to Guánica, showed the major “industrial” activities 

were salt production (13,600 tons per year), fishing 

(9.1 tons per week), charcoal production, and bark 

stripping of Rhizorphora mangle for tannin production 

in the Boquerón Forest (Hernández-Ramírez 1947). 

The principal farming activities (percentages of 
5

Figure 1—Places mentioned within southwestern Puerto Rico in text.

Population 1775-1995 Projections 1995 Projections 2005

Municipality - Cabo Rojo

0

10

20

30

40

50

60

1750 1800 1850 1900 1950 2000 2050

Year

Population (thousands)

Figure 2—Population in the Cabo Rojo municipality: 1750–2025 

(Junta de Planifi cación 1995, 2005; Ramos y Ramírez de Arellano 

and others 1985).

r

r

r

r

r

r

r

r r

r

r r

r

r

r

r r

r

r

r

r r

r

r

r

r

r r

r

r

r

r

r

r r r

r

r

r

r

r

r

r r r

r

r

r

r

r

r

r

r

r

r r r

r

r

r

r

r

r

r

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9

7

6 5

5

5

8

4

1

3

2

49

48

43

37

35

32

33

29

29

29 20

20

18

28

30

38

23

31

31 31

41

21

21

36

44

26

40

40 40 40

50

50

15

19

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19 47

22

39

27

45

12

13

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10 17

42

50

25

24 46

16

34 11

Legend

r Place names

Select water courses

Drained areas

Municipalities

Primary roads

Secondary roads

Tertiary roads

Management agency

Department of Natural and Environmental Resources

Conservation Trust of Puerto Rico

US Fish and Wildlife Service

Key Map items

 Protected areas: 

 1 Cabo Rojo Refuge

 2 Laguna Cartagena Refuge

 3 Boquerón Forest

 4 Boquerón Wildlife Refuge 

 5 Guánica Forest

 6 Los Morrillos de Cabo Rojo

 7 Parguera Natural Reserve

 8 Punta Guaniquilla Natural Reserve

 

 Towns and highways:

 9 Boquerón

 10 Cabo Rojo

 11 Ensenada

 12 Guánica

Key Map items

 Towns and highways: 

 13 Guayanilla

 14 Lajas

 15 Maguayo

 16 Parguera 

 17 San Germán

 18 Yauco

 19 PR Route 101

 20 PR Route 301

 21 PR Route 303 

 Geographic features:

 22 Bahía Boquerón

 23 Bahía Fosforescente

 24 Bahía Salinas

Key Map items

 Geographic features: 

 25 Bahía Sucia

 26 Cabo Rojo

 27 Cerro Mariquita

 28 Ciénaga El Anegado 

 29 Drainage Canal

 30 Hacienda La Romana

 31 Irrigation Canal

 32 Isla Cueva

 33 Isla Guayacán

 34 Isla Magueyes

 35 Lago Loco 

 36 Laguna Candelaria

 37 Laguna Fraternidad

 38 Laguna Guánica

Key Map items

 Geographic features: 

 39 Laguna Rincón

 40 Lajas Valley

 41 Magara Canal

 42 Mona Passage

 43 Playa Combate

 44 Punta Aguila

 45 Punta Guaniquilla

 46 Punta Pitahaya

 47 Quebrada Boquerón

 48 Quebrada Los Llanos

 49 Río Guanajibo

 50 Sierra Bermeja

0 1 2 4 6 8

km

farmland in parentheses) were: sugar cane production 

(57), corn (17), cotton and cowpeas (12), and several 

other crops (14). Pasturing of livestock included dairy 

cows, goats, sheep, and work animals.

Both refuge sites were essentially abandoned farms 

when acquired by the F&WS. The principal use of the 

Cabo Rojo Headquarters tract at the time of purchase 

was to graze cattle. Cattle grazing and the harvest 

of forage followed for several years. At Salinas, the 

major activity was salt production along with some 

wood harvest and grazing. 

During the 1920s, the Cartagena Lagoon and 

adjacent lands were partitioned into estates for the 

production of sugar cane (Danforth 1926). At that 

time, owners considered draining the lagoon to 
6

Growing population. The municipality of Cabo 

Rojo has experienced a substantial increase in 

population during the past 20 years adding to 

the pressures on the region’s wildlife. 

(Photo by Peter L. Weaver) 

increase the planting area but needed the water to 

irrigate nearby fields during the summer months. 

Later, the lagoon and surrounding lands were acquired 

by the Puerto Rican Government. 

Past use of the Tinaja tract included timber harvest 

for fuelwood and fence posts with recurrent fires 

used to clear the land for cattle grazing. Use of the 

steep upper slopes was abandoned first and reverted 

to scrub with scattered grass and barren patches. 

Grazing by goats, however, continued for a period. 

The midelevation slopes of Tinaja have had a mixture 

of secondary native and exotic tree species since 

the mid-1930s. On the lower slopes, where the last 

fire occurred in 1996, grazing by cattle and horses 

continued for a longer period. During the past decade, 

grazing and fire have been eliminated on the entire 

tract through improved fencing and regular F&WS 

patrol. A combination of factors—a dry climate, 

past sugar cane production, heavy grazing, recurrent 

fires, and later, a dense grass cover, have resulted in 

a slow recovery of woody vegetation on the refuge 

properties.

Both refuge properties are an integral part of 

public landholdings in southwestern Puerto Rico. 

Moreover, several other reserves contain about 6,800 

ha of critical wildlife habitat managed for resident and 

migratory bird species (table 1). Land use changes 

in and around the Guánica Forest during the past 50 

years showed that forest cover and urban areas had 

increased at the expense of agriculture, confirming 

a transition from an agrarian to urban society (Lugo 

and others 1996). Moreover, since 1990, the human 

population in the municipality of 

Cabo Rojo has grown in excess of 

projections because several coastal 

areas have been developed as summer 

homes (Junta de Planificación 

1995, 2005) (fig. 2). These changes 

highlight the importance of protecting 

and managing critical wildlife habitat. 

Environmental Conditions 

of the Refuges

The national wildlife refuge properties in 

southwestern Puerto Rico total 1174 ha and include 

four tracts of land ranging in size from 106 to 513 ha 

(table 1). 

Geology

The southwestern part of Puerto Rico is 

characterized by long ridges such as the Sierra 

Bermeja separated by parallel valleys like Lajas that 

extend eastward for many kilometers from the Mona 

Passage (Meyerhoff 1933) (fig.1). The ridges contain 

rocks of marine Cretaceous sediments such as ashy 

shales, massive limestone, and agglomerates. The 

valleys, in contrast, are partly covered with alluvial 

deposits of recent origin underlain by consolidated 

carbonate and clastic strata (sedimentary rock) of 

Cretaceous and Tertiary age (Bonnet and Tirado 

Sulsona 1950, Graves 1991). These, in turn, are 

underlain by igneous rocks at depths of 165 to 265 m. 

The 1500-ha Sierra Bermeja is the oldest and 

among the most interesting mountain ranges on the 

Caribbean plate. The Bermeja Complex of rocks is 

composed of basalt (volcanic origin), amphibolites 

and serpentinite (metamorphic rocks), and chert 

(formed by organisms in deep marine waters). The 

complex contains radiolarian (amoeboid protozoan) 

fossils in chert that date to 195 million years ago 

(Montgomery and others 1994a, 1994b; Pindell 
7

and Barrett 1990). The Lower Jurassic radiolarian 

signature is older than the Caribbean Sea and 

establishes the Pacific origin of the Caribbean plate. 

The Yauco-Boquerón anticlinal valley (Lajas 

Valley) is about 30 km long by 5 km at its widest 

point (Graves 1991). During the Tertiary period, 

the Lajas Valley was eroded by a large stream with 

its source in the mountains near the town of Yauco. 

Subsequently, the Río Guanajibo extended its 

headwaters and pirated streams flowing into the Lajas 

Valley (Mitchell 1922).

Nearly three-quarters the Cabo Rojo NWR 

consists of beach deposits and quartz sand deposits; 

also, Ponce limestone and alluvium cover nearly 

one-quarter of the tract (Bawiec 2001, Volckmann 

1984a) (table 2, fig. 3). The remaining lands, in 

descending areal coverage, are occupied by pyroxene 

olivine basalt, Mariquita chert, mangrove swamp, 

and Parguera limestone. Brief descriptions of the 

geological features at Cabo Rojo NWR (Headquarters 

and Salinas tracts) are listed below with map 

designations (table 2, fig. 3):

Table 2—Geology and soils on the Headquarters and the Salinas tracts of the Cabo Rojo 

National Wildlife Refuge

Geology/soils

Map designation and unit name a Area Refuge

ha percent

Geology b

Kjm—Mariquita chert, Lower Cretaceous-Upper Jurassic 1.60 0.21

Kp—Parguera limestone, Upper Cretaceous 0.14 0.02

Kpob—Pyroxene Olivine basalt, Upper Cretaceous? 10.71 1.41

Qa—Alluvium, Holocene-Pleistocene 80.31 10.60

Qb—Beach deposits, Holocene 255.62 33.73

Qs—Mangrove swamp, Holocene 1.54 0.20

Qts—Quartz sand deposits, Tertiary 306.52 40.45

Tpo—Ponce limestone, Miocene-Oligocene 101.37 13.38

Total c 757.81 100.00

Soils d 

AcD—Aguilita clay, 5 to 20 percent slopes 11.34 1.50

AcE—Aguilita clay, 20 to 50 percent slopes 1.71 0.23

AmC2—Amelia-Magüayo gravelly clay loams, 5 to 12 percent slopes 17.86 2.36

AsB—Americus fine sand, 2 to 5 percent slopes 86.77 11.45

Ca—Cartagena clay 19.34 2.55

Co—Coastal beach 24.09 3.18

FrB—Fraternidad clay, 2 to 5 percent slopes 89.89 11.86

Gf—Guaynabo fine sand 28.20 3.72

JcB—Jacana clay, 2 to 5 percent slopes 0.58 0.08

Lr—Limestone rock land 24.46 3.23

PzB—Pozo blanco gravelly clay loam, 0 to 5 percent slopes 1.47 0.19

PzC—Pozo blanco gravelly clay loam, 5 to 12 percent slopes 14.20 1.87

SoA—Sosa loamy sand, 0 to 5 percent slopes 105.91 13.97

Tf—Tidal flats 107.43 14.17

Ts—Tidal swamp 1.25 0.16

W—Salt water lagoons 223.41 29.48

Total c 757.91 100.00

a Refer to geology map (fig. 3) and soils map (fig. 5).

b Sources: Bawiec (2001); Volckmann (1984a, 1984b).

c Totals slightly different due to map error.

d Sources: Bawiec (2001); Carter (1965).
8

Geology (area [ha])

KJm (1.60)

Kp (0.14)

Kpob (10.71)

Qa (80.31)

Qb (255.62)

Qs (1.54)

QTs (306.52)

Tpo (101.37)

0 0.250.5 1 1.5 2

km

0 0.25 0.5 1 1.5 2

Miles

Figure 3—Geology and topography map of the Cabo Rojo National Wildlife Refuge 

(Bawiec 2001, Volckmann 1984a). 

•	 KJm—Mariquita chert, laminated, fine grained,

and dark gray in color containing radiolarian 

(amoeboid protozoa) and locally Foraminifera 

(amoeboid protists) 

•	 Kp—Parguera limestone, basal conglomerate that

grades upward into limestone composed of coarse 

skeletal clasts of fossils

•	 Kpob—Porphyritic, i.e., embedded crystals.

augite hypersthenes, i.e., lustrous silicate of iron/

magnesium-olivine basalt 

•	 Qa—Quaternary alluvium, unconsolidated sand

and gravel with some silt and clay; locally includes 

fluvial terrace deposits

•	 Qb—Beach deposits or poorly consolidated beach

sands; locally, sand dune deposits

•	 Qs—Swamp deposits, including sandy and silty

clay, locally ranging to peat or peaty muck 

•	 Qts—Friable quartz-rich sand deposits

•	 Tpo—Ponce limestone and Juana Diaz formation

with poorly cemented, somewhat friable 

calcirudite (skeletal limestone) and calcarenite 

(dune limestone), including oyster shells, minor 

corals, and gastropods 

More than 80 percent of the Laguna Cartagena 

NWR is covered by alluvium and nearly 20 percent by 

Mariquita chert; the remainder is Magüayo porphyry 

(Bawiec 2001, Volckmann 1984b) (table 3, fig. 4). 

Alluvium dominates in the valley, merging with 

colluvium at lowest levels along the valley walls. 

Mariquita chert covers the refuge from mid-to-high 

elevations. At highest elevations, Magüayo porphyry, 

the only geological feature not discussed above, 

covers a small area (table 3, fig. 4). 

•	 TKh—Magüayo porphyry, a quartz-bearing

inrusive rock occurring in isolated bodies
9

Table 3—Geology and soils on the Lagoon and Tinaja tracts of the Laguna Cartagena 

National Wildlife Refuge

Geology/soils

Map designation—unit name a Area Refuge

ha percent

Geology b

Kjm—Mariquita chert, Lower Cretaceous-Upper Jurassic 2.61 0.62

Qa—Alluvium, Holocene-Pleistocene 76.43 18.05

TKh—Magüayo porphyry, Lower Cretaceous 344.37 81.33

Total 423.41 100.00

Soils c 

Ag—Aguirre clay 19.73 4.66

AmB—Amelia-Magüayo gravelly clay loam, 2 to 5 percent slopes 37.85 8.94

AmC2—Amelia-Magüayo gravelly clay loams, 5 to 12 percent slopes 55.93 13.21

Ca—Cartagena clay 6.86 1.62

Cc—Cartagena silty clay loam, acid variant 23.50 5.55

Gc—Guánica clay 84.43 19.94

GuD—Guayama cherty clay loam, 5 to 20 percent slopes 0.70 0.16

GuF—Guayama cherty clay loam, 20 to 60 percent slopes 65.21 15.40

Sc—San Anton silty clay, moderately deep 0.92 0.22

Sn—Santa Isabel clay 20.54 4.85

Va—Vayas silty clay 7.77 1.84

Vo—Volcanic rock land 12.29 2.90

W—Fresh water lagoon 87.68 20.71

Total 423.41 100.00

a Refer to geology map (fig. 4) and soils map (fig. 6).

b Sources: Bawiec (2001); Volckmann (1984a, 1984b).

c Sources: Bawiec (2001); Carter (1965).

Soils

Soils in the Lajas Valley exhibit certain 

characteristics in common (Lugo-López and others 

1959). They are very deep with high, nearly uniform 

clay content. The top soil is usually 30 cm, but in 

places approaches 60 cm deep. The small pores 

conduct water rather well in the top soil but show very 

slow hydraulic conductivity in the subsoil. The soils 

are low in organic matter and nitrogen and generally 

high in soluble salts and exchangeable sodium, 

notably below 60 cm. The taxonomy of Puerto Rican 

soils, including those of the Lajas Valley, was outlined 

according to the new soils classification system (7th

approximation) (Lugo-López and Rivera 1976, 1977).

At the Cabo Rojo NWR, the soils of the 

Headquarters and Salinas tracts are distinct (table 

2). For the most part, the Headquarters tract is an 

upland site and Salinas is a coastal one. The 15 soil 

types on the refuge are separated into 5 types of clay; 

3 gravelly clay loams; 1 loamy sand; and 3 types of 

sand, limestone rock, tidal flats, and tidal swamps. 

Salt water lagoons cover about 40 percent of the 

Salinas tract, i.e., 30 percent of the entire refuge. 

Brief descriptions of the soils at Cabo Rojo NWR 

(Headquarters and Salinas) are listed below with map 

designations (Carter 1965) (table 2, fig. 5):
10

KJm (76.43)

Qa (344.37)

TKh (2.61)

0 0.10.2 0.4 0.6 0.8

km

0 0.1 0.2 0.4 0.6 0.8

Miles

100

2

100 00

200

200

Geology (area ha ) [ ]

Figure 4—Geology and topography map of the Laguna Cartagena National Wildlife Refuge 

(Bawiec 2001, Volckmann 1984a).

•	 AcD—Aguilita clays are sloping; well drainedto-excessively drained, shallow soils on hills. 

They are highly calcareous and of medium natural 

fertility.

•	 AcE—Aguilita clays are a steeper variant of the

above. 

•	 AmC2—Amelia-Magüayo gravelly clay loams are

a series that occur together. They are characterized 

by deep, well-drained acid soils in alluvium and 

colluvium, occurring on eroded, gentle foot slopes 

of hills. The mixture is about 60 to 70 percent 

Amelia, and 30 to 40 percent Magüayo.

•	 AsB—Americus fine sands are deep, gently

sloping, excessively drained soils on the Coastal 

Plain. They are strongly acid and low in fertility. 

•	 Ca—Cartagena clays are level to nearly level,

somewhat poorly drained, and slightly saline-tomoderately saline soils on alluvial fans. When wet, 

they are slightly sticky and plastic throughout the 

profile. They have a perched water table at depths 

of 60 cm or greater. 

•	 Co—Coastal beaches are made up of light-colored,

gently rolling beach sands along the coast. They 

are an unconsolidated mixture of coral, shell, 

and quartz and are calcareous, excessively well 

drained, and low in natural fertility.

•	 FrB—Fraternidad clays are deep, nearly level or

gently sloping and moderately well-drained soils. 

They are neutral to calcareous, heavy, sticky, 
11

shrink and swell with moisture changes, and are 

high in natural fertility. 

•	 Gf—Guayabo fine sands are gently sloping, deep,

excessively drained soils on the Coastal Plain. 

They are acid and low in natural fertility. 

•	 JcB—Jácana clays are gently to strongly sloping,

well-drained alluvial and colluvial soils in valleys. 

They are acid and medium in natural fertility.

•	 Lr—Limestone rocklands have steep slopes

mostly covered with limestone cobbles and stones. 

These lands are shallow and low in moistureholding capacity.

•	 PzB—Pozo blanco gravelly clay loams are nearly

level to strongly sloping, moderately deep, welldrained soils on alluvial and colluvial foot slopes. 

They are highly calcareous and medium in 

natural fertility.

•	 PzC—Pozo blanco gravelly clay loams are a

steeper variant of the above.

•	 SoA—Sosa loamy sands are moderately deep,

excessively drained soils on nearly level to sloping 

Coastal Plains. They are underlain by stratified, 

dense, sandy clay and are low in available moisture 

and natural fertility.

•	 Tf—Tidal flats are nearly level lands along the

coast. They are poorly drained and frequently 

flooded by salt water during high tides. They have 

a high water table, are saline, and when dry, are 

thinly crusted with salt.

•	 Ts—Tidal swamps are areas that are permanently

flooded by shallow water along the coast.

At the Laguna Cartagena NWR, the soils situated 

in the Lajas Valley are distinct from those in the 

Sierra Bermeja. The fertile, alluvial soils of the Lajas 

Soils (area [ha])

AcD (11.34)

AcE (1.71)

AmC2 (17.86)

AsB (86.77 )

Ca (19.34)

Co (24.09)

FrB (89.89)

Gf (28.20)

JcB (0.58)

Lr (24.46)

PzB (1.47)

PzC (14.20)

SoA (105.91)

Tf (107.43 )

)

)

Ts (1.25

Water (223.41

0 0.250.5 1 1.5 2

km

0 0.25 0.5 1 1.5 2

Miles

Figure 5—Soils map of the Cabo Rojo National Wildlife Refuge (Carter 1965).
12

Valley are deep, heavy, difficult to drain, and often 

affected by salt, particularly in previously irrigated 

areas (Koenig 1953, Picó 1974, Willardson 1958). In

contrast, the steep slopes of the Sierra Bermeja are 

shallow, stony, well drained, and dry. 

Twelve soil types occupy the refuge, including 

three types of clay, two types of silty clay, two 

gravelly clay loams, two cherty clay loams, one silty 

clay loam, sand, and volcanic rock (table 3). Slightly 

more than one-quarter of the Lagoon tract is occupied 

by the Cartagena Lagoon, i.e., about 20 percent of 

the entire refuge (table 3). Soils not described above 

for Cabo Rojo follow with map designations (Carter 

1965) (table 3, fig. 6):

•	 Ag—Aguirre clays are deep, level or nearly level,

poorly drained soils on alluvial fans. They are 

sticky and plastic when wet, highly calcareous, 

and of medium natural fertility. 

•	 AmB—Amelia-Magüayo gravelly clay loams are

a series that occur together. They are characterized 

by deep, well-drained acid soils in alluvium and 

colluvium, occurring on the gentle foot slopes 

of hills. The mixture is about 60 to 70 percent 

Amelia, and 30 to 40 percent Magüayo.

•	 Cc—Cartagena silty clay loams, acid variant are

level, somewhat poorly drained, and slightly saline 

soils on alluvial fans. They have a slightly coarser 

textured surface layer than Cartagena clays. They 

shrink and crack when dry, swell when wet, and 

are of medium natural fertility.

•	 Gc—Guánica clays are deep, nearly level, slowly

permeable, calcareous, and with medium natural 

fertility. They have a high shrink-swell potential, 

and crack when dry. 

•	 GuD—Guayama cherty clay loams are shallow,

well-drained soils on the steep slopes of Sierra 

Bermeja. The soils are acid and medium in 

natural fertility.

Soils (area [ha])

Ag (19.73)

AmB (37.85)

AmC2 (55.93)

Ca (6.86)

Cc (23.50 )

Gc (84.43)

GuD (0.70)

GuF (65.21)

Sc (0.92)

Sn (20.54)

Va (7.77)

Vo (12.29)

Water (87.690)

0 0.1 0.2 0.4 0.6 0.8

km

0 0.1 0.2 0.4 0.6 0.8

Miles

Figure 6—Soils map of the Laguna Cartagena National Wildlife Refuge (Carter 1965). 
13

•	 GuF—Guayama cherty clay loams are a steeper

variant of the former. 

•	 Sc—San Anton silty clays (moderately deep) are

moderately deep, well drained, nearly level alluvial 

soils on flood plains adjacent to streams. They are 

slightly acid to mildly calcareous, and moderately 

permeable, with a firm heavy clay substratum. 

They are high in natural fertility. 

•	 Sn—Santa Isabel clays are deep, moderately welldrained soils on nearly level terrain in valleys. 

High in fertility, they shrink and swell with 

changes in moisture. 

•	 Va—Vayas silty clays are nearly level, poorly

drained soils on flood plains. They have a high 

water table, and are neutral to mildly alkaline 

above and saline at lower levels. 

•	 Vo—Volcanic rocklands are characterized by

acid volcanic rock outcrops that cover more than 

one-half of the mapping area. Areas not covered 

by outcrops are covered by stony or gravelly clay 

loams or clays. 

Physiography and Hydrology 

The western part of the Lajas Valley rises from sea 

level at Bahía Boquerón to about 13 m in elevation 

at PR Route 303 (fig. 1). The highway runs along the 

drainage divide that separates the eastern and western 

parts of the valley (Graves 1991). The principal 

aquifer of the Lajas Valley consists of alluvial 

deposits. Consolidated sedimentary rocks with 

different hydraulic characteristics underly the deposits 

and could be considered as a distinct aquifer (Graves 

1991). The principal aquifer is recharged by rainfall 

and streamflow, most of which occurs through coarse 

grain alluvial fans along the valley’s edges. Seasonal 

changes of 0.6 m are apparent in the altitude of the 

potentiometric surface, which averages about 15 m 

at the northern and southern boundaries of the valley, 

and 4 m in the middle. Discharge of ground water 

occurs through pumping, evapotranspiration, and 

subsurface seepage. 

The Headquarters tract, characterized by gentle 

slopes, ranges from <5 m in the southwest corner to 

about 35 m in elevation in the northwest corner (fig. 

3). The entire Salinas tract is virtually at sea level. 

The large, shallow Laguna Candelaria and Laguna 

Fraternidad salt water lagoons, which fluctuate in 

size according to rainfall, runoff from upland areas, 

and tidal levels are the most striking features of the 

tract (Tripp and Collazo 2003). Evaporation from the 

lagoons is high, and salinity tends to increase with 

distance from the water control structures. Water 

movement in the lagoons is influenced by wind and 

flow is generally from east to west, particularly in 

Laguna Fraternidad. 

The Lagoon tract ranges in elevation from 40 

m on the south side bordering the slopes of Sierra 

Bermeja to 11 m at the shoreline of the Cartagena 

Lagoon (Schaffner 1995b, see endnotes) (fig. 4). To 

the north, the property is flat, rising to 12 m at the 

border with private holdings. At the time of discovery, 

Cartagena Lagoon was once part of a system of 50 

freshwater lagoons throughout Puerto Rico (Colón 

1982a). Within the Lajas Valley, it was part of a series 

of wetlands that extended from Laguna Guánica 

westward for about 30 km through Ciénaga El 

Anegado and the Cartagena Lagoon to Laguna Rincón 

on the west coast (Ramírez Toro and Minnigh 1997, 

see endnotes).Cartagena Lagoon was a landlocked 

water body maintained by runoff from the surrounding 

mountains; heavy rainfall, however, caused ephemeral 

streams to raise the water level in the lagoon, which 

then overflowed, draining westward 7 km through 

the Río Boquerón, i.e., today the Drainage Canal, 

to a mangrove swamp along Bahía Boquerón 

(Danforth 1926). The water level of the lagoon varied 

seasonally, rising during the fall with heavy rainfalls 

and declining during the warm summer months. The 

size of the lagoon could fluctuate from 0 to 140 ha 

and its depth from 0 to 1.5 m. The lagoon’s dynamic 

nature provided a variety of habitats—open water 

for the foraging of resident and migratory waterfowl, 

emergent vegetation with nesting areas for resident 

aquatic birds, and foraging habitat for wading birds 

during the seasonal change in depth. 

Agricultural development subsequently wrought 

dramatic changes to the lagoon’s water regimen. 

Cartagena Lagoon remains today, but by the early 

1980s, it had lost 95 percent of its open water and 

some of its wildlife. This was due to a host of 

activities such as deforestation around the lagoon, 

pumping water for irrigation, eutrophication and 

pollution, canal construction, the growth of weedy 

plants, cattle grazing, and the concentration of hunters 

at the only remaining lagoon in the Lajas Valley. 

Today, water enters the Lajas Valley from the east 

via the Irrigation Canal and flows to the Cartagena 

Lagoon through the Margara Canal. This gravity-fed 
14

Tinaja. Rising to 300 m at Cerro Mariquita, the 110-ha Tinaja tract has 

a long history of fires and grazing. Today, with protection, secondary 

forest is rapidly replacing the grass cover. (Photo by Peter L. Weaver) 

Table 4—Mean annual rainfall at the U.S. Fish and Wildlife Service 

refuges, 1991–2005

Year Cabo Rojo Laguna Cartagena Comparison a

 mm/year percent b mm/year percent b percent

1991 575 62 707 67 81

1992 1032 111 1149 109 90

1993 870 94 1135 107 77

1994 715 77 870 82 82

1995 668 72 918 87 73

1996 877 94 978 92 90

1997 459 49 709 67 65

1998 1464 157 1687 159 87

1999 740 80 1140 108 65

2000 980 105 1094 103 90

2001 945 102 1199 113 79

2002 794 85 602 57 131

2003 1509 162 1119 106 135

2004 996 107 1110 105 90

2005 1327 143 1461 138 91

Mean 930 100 1058 100 88

a Comparison: annual rainfall Cabo Rojo/Laguna Cartagena.

b Percent = rainfall for particular year divided by 15-year mean for each site. 

aqueduct extends 37 km from Lago Loco, passing 

north of the town of Magüayo, through the Cartagena 

Lagoon, and west to Bahía Boquerón. The aqueduct 

supplies about 5 670000 m3

 (4,600 acre-feet) of water 

per month (Graves 1991). 

The Tinaja tract, contiguous with the lagoon 

property, lies about 0.5 km south of Cartagena 

Lagoon. The tract measures approximately 0.8 by 

1.6 km (upslope), and ranges from 20 to 290 m 

in elevation (fig. 4). The highpoint of the Sierra 

Bermeja is Cerro Mariquita at 301 m. It lies about 

50 m upslope to the west. Tinaja is dissected by two 

major arroyos that carry water towards the Cartagena 

Lagoon during heavy downpours. 

Climate

Puerto Rico, lying in the path of the easterly trade 

winds, has a climate tempered by proximity to the 

ocean (Calvesbert 1970). The island’s weather is 

influenced by major systems, including easterly waves 

and cold fronts, and occasionally by 

tropical storms and hurricanes. The 

southwestern part of the island, in the 

shadow of the Central Mountains, 

is drier than the north coast and 

mountain summits. 

Precipitation—During the 15-year 

period between 1991 and 2005, mean 

annual rainfall at the Cabo Rojo 

Headquarters tract, 13 km southwest 

of the town of Lajas, averaged about 

930 mm/year (table 4). During the 

same period, mean annual rainfall 

at the Cartagena Lagoon tract, 5 km 

southwest of Lajas, averaged about 

1058 mm/year. At Cabo Rojo, rainfall 

in particular years varied from 49 to 

162 percent of the 15-year mean (table 

4). At Cartagena Lagoon, the range was 

from 57 to 159 percent. The driest years 

during the period were 1997 at Cabo 

Rojo and 2002 at Cartagena Lagoon; the 

wettest years were 2003 at Cabo Rojo 

and 1998 at Cartagena Lagoon. These 

values agree with previous estimates 

(Calvesbert 1970, Ravalo and others 1986). 
15

Heavy rains. A heavy downpour converts the normally dry arroyo 

traversing the Tinaja tract into an instant stream with a small waterfall.

(Photo by Peter L. Weaver)

250

200

150

100

50

0

1 2 3 4 5 6 7 8 9 10 11 12

Climatic variables

Month

(A) Climate - Lajas

180

160

140

120

100

80

60

40

20

0

Climatic variables

1 2 3 4 5 6 7 8 9 10 11 12

Month

Temperature (C) Rainfall (mm)

(B) Climate - Magueyes

Figure 7—Rainfall and temperature diagram (1971–2000): 

(A) Lajas, (B) Magueyes.

When comparisons are made by refuge site, rainfall 

at Cabo Rojo averaged 88 percent of that at Cartagena 

Lagoon for the entire measurement period. During 

particular years, rainfall at Cabo Rojo varied from 65 

to 135 percent of that at Cartagena Lagoon. Rainfall at 

Cartagena Lagoon exceeded that at Cabo Rojo except 

for 2002 and 2003.

Rainfall in Puerto Rico for the period from 1990 

to 1997 was only 87 percent of the normal (30 years 

of record) for the 12 stations with the longest records 

(Larsen 2000). Between 1991 and 1997, mean annual 

rainfalls were 742 mm at Cabo Rojo and 924 mm 

at Cartagena Lagoon (table 4). Comparable rainfall 

values for 1998 through 2005 were greater, averaging 

1094 mm at Cabo Rojo and 1176 mm at Cartagena 

Lagoon. In this instance, the earlier 7-year records 

represent only 68 and 78 percent, respectively, of 

the later 8-year records, and 80 and 87 percent, 

respectively, of the entire 15-year record (table 4). 

Rainfall averages between 1971 and 2000 are 

available for two sites nearby—the town of Lajas, 

and Isla Magueyes on the coast 8 km south of Lajas 

(figs. 7A and 7B). Mean annual rainfall at Lajas is 

1143 mm, ranging from 50 mm in January to 173 mm 

in October. At Magueyes, the mean annual rainfall 

is only 768 mm, ranging from 29 mm in February to 

131 mm in October. At both sites, most rainfall occurs 

from the beginning of August through the end of 

November, averaging 54 percent of the total at Lajas 

and 59 percent at Magueyes. A second, but lower 

rainfall peak, is evident during May.

Temperature—Temperature averages are also 

available for Lajas and Magueyes (figs. 7A and 7B). 

The mean annual temperature at Lajas is 25.3 °C, 

ranging from 23.2 °C in January to 26.8 °C in July. 

At Magueyes, it is 27.1 °C, ranging from 25.2 °C in 

January to 28.5 °C in June and August. These values 
16

0 5 10 20 30 40

Miles 0 5 10 20 30 40

km

Legend for major hurricanes in Puerto Rico since 17001, 2

1

Key: A = hurricane vortex passed over island with hurricane force winds; B = hurricane vortex passed over part of the island or offshore, with hurricane force winds on part of the

island; A/? and B/? = uncertain trajectory (i.e., San Ramón II in 1772 passed northeast of the island). Trajectories in 1899 and later are known; pre-1899 trajectories were

sketched from descriptions except those classed as A/? or B/?.

2

Sources: Monzón 1996, Neumann and others 1988, Quiñones 1992, Salivia 1972.

32B 22B

26A

27A 30A 5A, 25A

10B, 13A, 14B, 15B, 21B

33A

19A 17A 24A

1A, 2A, 3A

20A 11A, 18A

29A

31B

6A

4B

23A

28B

Key

10-B

11-A

12-B/?

13-A

14-B

15-B

16-A/?

17-A

Year

1805

1807

1809

1813

1814

1816

1819

1825

Name

San Vicente II

San Jacinto

San Esteban III

San Liborio II

San Liborio III

San José de Cupertino

San Mateo III

Santa Ana

Key

18-A

19-A

20-A

21-B

22-B

23-A

24-A

25-A

Year

1827

1835

1837

1851

1852

1867

1876

1893

Name

San Jacinto II

San Hipólito

N.S. de los Angeles

San Agapito II

San Llorenzo

San Narciso

San Felipe I

San Roque III

Key

26-A

27-A

28-B

29-A

30-A

31-B

32-B

33-A

Year

1899

1928

1931

1932

1956

1989

1996

1998

Name

San Ciriaco

San Felipe II

San Nicolás

San Cipriano II

Santa Clara

Hugo

Hortense

Georges

Key

1-A

2-A

3-A

4-B

5-A

6-A

7-B/?

8-B/?

9-B/?

Year

1738

1738

1751

1766

1766

1772

1772

1780

1785

Name

Santa Rosa II

San Leoncio III

San Agapito

San Jenaro

San Marcus

San Agustín

San Ramón II

San Antonio

San Lupo

agree closely with the previous mean estimate of 26 

°C, with monthly means varying by only 3 °C or 

so during the year (Calvesbert 1970, Daly and 

others 2003).

Relative humidity and evaporation—Two estimates 

of pan evaporation in the Lajas Valley are available: 

1930 mm/year (Anderson 1977) and 1525 mm/year 

(Graves 1991). Potential evapotranspiration at Cabo 

Rojo, estimated by the Hargreaves-Samani model, 

averaged 1660 mm/year (Goyal 1991). Another 

estimate of potential evapotranspiration for the Lajas 

Valley was 1250 mm/year (Anderson 1977). 

Hurricanes—Since 1700, Puerto Rico has apparently 

experienced hurricane force winds at least 33 times. 

Twenty hurricanes had trajectories over much of the 

island (type A hurricane) and 13 had trajectories over 

a portion of the island, or immediately offshore (type 

B hurricane) (Neumann and others 1988, Quiñones 

1992, Salivia 1972) (fig. 8). Local effects from more 

distant storms or hurricanes (type C storms) were 

also experienced more than 50 times. Since hurricane 

size, duration, and wind speed vary considerably, 

estimating trajectories and classifying storm types 

before the 20th century is a matter of conjecture. 

Several of the earlier hurricanes could have impacted 

the forests in southwestern Puerto Rico to some 

degree (fig. 8). 

Four hurricanes damaged Puerto Rico’s southwest 

since the end of the 19th century (fig. 8). San Ciríaco 

of 1899 and San Felipe II of 1928 passed to the 

northeast, and Georges of 1998 to the north. San 

Ciríaco with sustained winds around 180 km/hour, and 

San Felipe II with winds at 250 km/hour, were major 

storms, the latter perhaps being the most powerful on 

record for Puerto Rico. Hortense in 1996, a category 

1 hurricane on the Saffir-Simpson Hurricane Scale, 

passed directly over the refuges (Monzón 1996). With 

sustained winds of 135 km/hour and rainfall averaging 

between 75 and 125 mm in the southwest, Hortense 

caused flooding, uprooted trees, and damaged 

buildings and electrical lines. Its short residence time 

over Puerto Rico, poorly defined center, and generally 

low winds prevented more damage. Georges in 1998, 

Figure 8—Hurricane trajectories over Puerto Rico since 1700.
17

a category 3 hurricane with sustained winds of 180 

km/hour, flooded the Lajas Valley and damaged 

both refuges. 

Flora

“Puerto Rico was originally mantled by forests 

from the level of the sea to the summit of its 

mountains” (Hill 1899, p. 7), and the dominant 

vegetation in southwestern Puerto Rico at the time 

of Columbus’ arrival was forest (Murphy 1916). 

Mangroves probably covered about 12000 ha of 

Puerto Rico’s shorelines, declining to nearly onehalf by 1938 (Carrera and Lugo 1978). The dry, 

rocky slopes along the south coast were covered 

by woodland and cactus (Wadsworth 1950, 1962). 

Prominent tree species included Amyris elemifera, 

Bucida buceras, Bursera simaruba, Ceiba pentandra, 

Colubrina arborescens, Exostema caribaeum, 

Guaiacum officinale, Pictetia aculeata, and Tabebuia 

hetrophylla (Wadsworth 1962) (see appendix A for 

authorities, and common English and Spanish names 

of plant species mentioned in text). 

The flora of Puerto Rico, ranging from coastal 

areas through the mountainous interior, was studied 

in detail during the 1920s (Britton and Wilson 

1923–1930, Cook and Gleason 1928, Gleason and 

Cook 1927). Britton, the founder and first director 

of the New York Botanical Garden, helped carry out 

the first systematic inventory of natural history in 

the Caribbean (Sastre-D.J. and Santiago-Valentín 

1996). Many of his expeditions included sites in 

southwestern Puerto Rico. Major environmental 

concerns at that time included the island’s loss of 

forest cover, and the need for forest policy 

and reforestation. 

Mangroves grew along the coast. Bucida buceras, 

often draped with the epiphyte Tillandsia recurvata, 

and its associates were common in the stream deltas 

and alluvial valleys of the original forest (Gleason 

and Cook 1927). Capparis cyanophallophora, 

Citharexylum fruiticosum, Coccoloba diversifolia, 

Guaiacum officinale, Guapira fragrans, Pisonia 

albida, and Tabebuia hetreophylla were common 

along the coastal strip (Gleason and Cook 1927). 

Pterocarpus officinalis, associated with the landward 

side of mangroves and swamps, probably grew 

around Cartagena Lagoon at the time of the island’s 

discovery. Subsequently, during colonization and 

settlement, trees were cut for construction, fuel, and 

agriculture. At the beginning of the 20th century, 

the deciduous forests, ranging from Patillas in 

southeastern Puerto Rico along the south coast to 

Hormigueros near Mayagüez, were recurrently burned 

and grazed (Murphy 1916). 

In the early 1900s, aside from locally grown fruit 

trees, the most conspicuous trees of the southwestern 

coastal areas were the natives Cassine xylocarpa, 

Ceiba pentandra, Hippomane mancinella, Hymenaea 

courbaril, Manilkara bidentata, and the exotics 

Delonix regia and Tamarindus indica (Hill 1899). 

Other native trees observed during the early 1900s 

were Acacia farnesiana, Andira inermis, Bucida 

buceras, Guazuma ulmifolia, Inga laurina, Inga 

vera, Pictetia aculeata, Randia aculeata, Stahlia 

monosperma, and the exotic Haematoxylum 

campechianum (Gleason and Cook 1927, Murphy 

1916). Pastureland, if unattended, would soon revert 

to a thorn thicket characterized by Capparis flexuosa, 

Parkinsonia aculeata, Pilosocereus royenii, and 

Pithecellobium unguis-cati among other species 

(Gleason and Cook 1927). 

The Great Depression and World War II forced 

most islanders to utilize available land for pasture and 

crops. During the late 1940s, only 6 percent of Puerto 

Rico was in natural forest. After the mid-1980s, 

however, much of the agricultural land had been 

abandoned and secondary forests occupied about onethird of the island, including much of the southwest 

(Birdsey and Weaver 1982, Franco and others 1997). 

Occasional remnants of past forest cover, for example, 

the large Hymenaea courbaril along the jeep road in 

Tinaja, show that the largest trees in the original dry 

forest attained diameters of at least 1 m and heights 

approaching 20 to 25 m. 

All of Puerto Rico’s native tree species and 

many introduced exotics have been described and 

illustrated in local publications (Francis and Liogier 

1991, Francis and Lowe 2000, Little and Wadsworth 

1964, Little and others 1974). Moreover, taxonomic 

descriptions of the flora (grasses, trees, and vines) 

are available for Puerto Rico and other islands in the 

Caribbean (Acevedo-Rodríguez and Woodbury 1985; 

Hitchcock 1936; Howard 1979, 1988–1989; Liogier 

1985–1997; Liogier and Martorell 1982, 2000). Eight 

common species of grasses have also been illustrated 

in a field guide (Más and García 1990). Plant 

nomenclature in this report has followed Liogier.
18

Recently, all of Puerto Rico’s forest type and 

land cover types were mapped (Helmer and others 

2002). Mapping zones encompassing the refuges 

and protected areas of the southwest were designated 

as dry-alluvial and dry-volcanic/sedimentary/

limestone areas. The vegetation formations included 

lowland dry semideciduous forest or woodland/

shrubland, tidal and semipermanently flooded 

evergreen sclerophyllous forest, and lowland dry 

mixed evergreen drought-deciduous shrubland 

with succulents. The major land use of much of the 

southwest is pasture. 

Major Forest Types in the Southwest

Southwestern Puerto Rico, including all of the 

refuge properties, is situated in the Subtropical 

Dry Forest according to the ecological life zone 

system (Ewel and Whitmore 1973, Holdridge 1967). 

Within the life zones, associations are referred 

to as zonal (that is, with a typical climate and a 

typical soil type), or azonal if strongly influenced by 

edaphic, atmospheric, or hydric conditions. Another 

classification system used throughout the Caribbean 

also recognizes environmental effects on vegetation 

(Beard 1949, 1955). 

The major types of vegetation cover in the 

southwest are: mangroves; salt flats; littoral woodland 

(beach thickets); mesquite and semievergreen 

woodland; coastal shrub or thorn woodland; 

deciduous woodland; agricultural lands, including 

pastures; and residential areas and roadside trees 

(McKenzie and Noble 1990). Mangroves and salt

flats are the vegetation types that most closely 

approach the original cover. Moreover, the littoral 

woodland still contains numerous tree species that 

were part of the original vegetation. In contrast, 

pastures, agricultural lands, and residential areas 

are the most modified landscapes. The remaining 

vegetation types contain exotic and native species in 

compositions that differ from the original vegetation. 

Currently, the refuge sites are highly disturbed by 

human activity and occupied by secondary vegetation, 

including numerous exotics. The following vegetation 

descriptions, mainly overstory trees, are derived from 

previous work on forest cover in southwestern Puerto 

Rico (McKenzie and Noble 1990).

Mangrove forest—Mangroves are dominated by 

four tree species: (1) Avicennia germinans, (2) 

Conocarpus erecta, (3) Laguncularia racemosa, 

and (4) Rhizophora mangle. Rhizophora mangle

frequently forms a fringe along the coast and 

occasionally isolated islands of vegetation in shallow 

water. Avicennia germinans and Laguncularia 

racemosa are common in estuaries and around salt 

flats. Conocarpus erecta is commonly found at the 

landward edge of tidal mangrove swamps. 

Salt flats—Often adjacent to mangrove swamps, 

the salt flats contain a ground cover with Batis 

maritima, Heliotropium curassavicum, Sesuvium 

portulacastrum, and other salt-tolerant plants. Trees 

are usually limited to scattered patches of Avicennia 

germinans and Laguncularia racemosa. 

Littoral woodland (beach thickets)—The littoral 

woodland, stretching in a fringe along the coast 

behind beaches, contains several native canopy 

species like Canella winterana, Clerodendron 

aculeatum, Coccoloba diversifolia, Coccoloba 

uvifera, Colubrina arborescens, Erithalis fruticosa, 

Erythroxylum aerolatum, Jacquinia arborea, 

Krugiodendron ferreum, Piscidia carthagenesis, 

Pisonia albida, Randia aculeata, and Rochefortia 

acanthophora. The exotic Thespesia populnea

reproduces abundantly in some areas. 

Mesquite and semi-evergreen woodland—Land 

clearing for pasture and agriculture in the lowlands 

between the salt flats and nearby mountains has 

modified the original forest dominated by Bucida 

buceras (Cook and Gleason 1928), into an association 

with grass and scattered Prosopis juliflora, resembling 

a savanna (Garcia-Molinari 1952). Other common 

trees include the natives Bucida buceras, Guaicum 

officinale, and Pisonia albida along with numerous 

understory species. Common exotics are Leucaena 

leucocephala, Pithcellobium dulce, and Tamarindus 

indica. Trees common to arroyos in the mesquite 

and semievergreen woodland include natives such 

as Acacia farnesiana, Crecentia cujete, Piscidia 

carthagenesis, and Tabebuia heterophylla and 

exotics like Albizia lebbeck, Bauhinia monandra, and 

Swietenia mahagoni.

Coastal shrub or thorn woodland—Some areas 

are dominated by a coastal scrub previously called 

cactus scrub or thorn woodland (Beard 1949). The 

dominant canopy trees in this vegetation type are 

almacigo Bursera simaruba, Guaicum offincinale, 

Pisonia albida, Plumeria alba, Prosopis juliflora, 

and scattered Bucida buceras. Near Bahia Sucia, the 
19

Large native tree. One of the largest native trees in Puerto Rico’s dry 

southwest, Hymenaea courbaril L. (West Indian locust) grows on the 

refuge sites. (Photo by Peter L. Weaver) 

composition varies somewhat in small areas of 

limestone. Bucida buceras, Crescentia cujete, 

Guapira discolor, Pisonia albida, Prosopis 

juliflora, and Tabebuia heterophylla are the 

principal canopy species. 

Deciduous woodland—The major tree species 

in the deciduous woodland are Bourreria 

succulenta, Bucida buceras, Bursera simaruba, 

Clusia rosea, Coccolobia diversifolia, 

Colubrina arborescens, Colubrina elliptica, 

Erothroxylum aerolatum, Guazuma ulmifolia, 

Pisonia albida, Rauvolfia nitida, Thouinia 

striata var. portoricensis, Zanthoxylum 

martincense, Zanthoxylum monophyllum, and 

Ziziphus reticulata. Occasionally Guaicum 

offincinale is found.

Other Vegetation Types

Other vegetation types are solely the result of 

human activities and are maintained in a highly 

modified state by regular tending, at least for a 

period of time. 

Agricultural lands and pastures—Agricultural 

lands often contain Carica papaya, Persea 

americana, and ground crops such as melons, 

peppers, pineapples, pumpkin, and yucca. 

Pastures are occupied by about 15 species 

of native and introduced grasses. Secondary 

regeneration of native tree species and planted 

Prosopis juliflora and Swietenia mahagoni are 

seen in the vicinity of the refuges. 

Residential and roadside trees—Selected 

for shade, ornament, and fruit, or regenerated 

naturally, several species of trees survive around 

private homes, along fence lines, and scattered 

in fields. The most common trees planted for 

shade or as ornamental are Delonix regia, 

Swietenia mahagoni, Tabebuia heterophylla,

Tecoma stans, and Tectona grandis. Among the 

most common fruit trees are Tamarindus indica. 

Forest tree plantings (plantations)—Forest 

trees are planted for a variety of purposes, 

usually timber production. In Puerto Rico’s dry 

southwest, however, timber production was not 

a priority (Birdsey and Weaver 1982), although 

experimental work with timber species was 

attempted more than 60 years ago (Marrero 

1950; Wadsworth 1943, 1990). During the past 25 

years, at least 80 tree species have been planted on the 

refuge lands to stimulate native forest regeneration 

and to restore wildlife habitat. 

Vegetation Types at the Refuges

Frequent fires, heavy grazing, and continuous 

cropping, mainly in sugar cane, characterized the past 

use of the refuges. Soil erosion and sedimentation 

were rampant. During this period, native plants were 

severely reduced in numbers and several exotics 

increased in areal extent. Today, the refuges are 

covered with pasture interspersed with native and 

exotic trees, patches of secondary forest, and tree 

plantings of various species.

Cabo Rojo NWR—The Cabo Rojo NWR is divided 

into two tracts: Headquarters and Salinas. Intensive 

farming and grazing were the main uses of the 

Headquarters tract. In 1978, livestock were removed 

when the first refuge manager arrived. In 1980, the 

planting of Bucida buceras trees began in several 

areas. The planting continued irregularly through 1995 

and annually thereafter when several new tree species 

were introduced (Weaver and Schwagerl 2004). 

The existing forest cover in the wooded portion 

of the Headquarters tract contains Prosopis juliflora 

along with a few other exotic tree species such as 
20

Fires. The exotic Prosopis juliflora (Sw.) DC. (mesquite) was planted 

long ago on the Cabo Rojo National Wildlife Refuge (at left in photo). 

Many mesquite trees have survived recurrent fires which are a constant 

threat to wildlife and undermine efforts to restore the natural habitat. 

(Photo by Peter L. Weaver) 

Grass and cactus. Grass and cactus, a vegetation type maintained by 

fires, glistens in the early morning sun. (Photo by Peter L. Weaver)

Salt flats. The salt flats and ponds on the Salinas tract at Cabo Rojo 

provide wildlilfe habitat for numerous species of migratory and 

resident birds. (Photo by Peter L. Weaver)

Albizia lebbeck, Leucaena leucocephala, Melicoccus 

bujugatus, Parkinsonia aculeata, Pithecellobium 

dulce, and Tamarindus indica (Stacier 1992; Zuill 

1985, see endnotes). Maturing Bucida buceras 

trees also grow along with occasional Hymenaea 

courbaril and other native species scattered in 

secondary patches. Regeneration of native tree species 

is apparent under existing tree cover. In general, 

drainages contain the greatest concentration of 

large trees. 

In the grass-dominated areas, Panicum maximum 

and Cenchrus ciliaris grow along with widely 

scattered trees of Prosopis juliflora or clumps of 

cactus. The grasses cause serious problems. They are 

aggressive competitors for light and water and retard 

the regeneration of native tree species. Moreover, 

the grasses facilitate the spread of wildfires that 

sometimes enter from surrounding farmlands. 

The Salinas tract of the refuge contains three 

principal vegetation types: (1) scattered patches of 

mangrove, (2) stretches of littoral woodland, and (3) 

the salt flats with salt-tolerant plants. The mangroves 

and the littoral woodland have been affected by the 

harvest of fenceposts and fuelwood in the past. The 

tract also contains shallow lagoons, not exceeding 

0.5 m in depth, and occupying about 225 ha (Negrón 

González 1986). 

Laguna Cartagena NWR—The Laguna Cartagena 

NWR is partitioned into two major tracts: Lagoon 

and Tinaja. The Lagoon tract has three distinct areas: 

(1) the lagoon, centrally located, with its aquatic 

vegetation; (2) a recently abandoned sugar cane 

plantation, now largely planted with trees lying to 
21

the north; and (3) an abandoned pasture with naturally 

occurring trees and several patches of planted trees, 

lying to the south (Weaver and Schwagerl 2005). The 

lagoon itself consists of open water and areas with 

cattails (Díaz-Soltero 1990). 

The Tinaja tract also has three distinct areas. The 

lower, gentler slopes were grazed and recurrently 

burned through 1996 when the F&WS acquired 

the property. At that time, the lower slopes had the 

appearance of savanna-like grassland with scattered 

native and exotic trees. Today, in the absence of 

grazing and fire, numerous trees have regenerated, 

principally the exotic Leucaena leucocephala (Weaver 

and Schwagerl 2004). The midslopes are covered with 

secondary vegetation, some dating back at least to the 

mid-1930s when it was visible in aerial photographs 

(Weaver and Chinea 2003). The upper slopes around 

Cerro Mariquita are very steep, rocky, and heavily 

eroded by past grazing. Much of the vegetative cover 

is short, yet of considerable ecological value due to 

the presence of rare and endangered plant species. 

The partially forested Tinaja tract is important to 

Cartagena Lagoon as a source of unpolluted water. 

Vegetative surveys have been carried out at 

both the Cabo Rojo Refuge (McKenzie 1986, see 

endnotes) and the Laguna Cartagena Refuge (DíazSoltero 1990; Proctor 1996, see endnotes). The 

identified plants include: 10 ferns, 1 gymnosperm, 103 

monocotyledons, and 533 dicotyledons (appendix B). 

Among the monocots are 3 Alismataceae, 3 Araceae, 

5 Arecaceae, 2 Commelinaceae, 22 Cyperaceae, 55 

Graminae, 3 Liliaceae, 1 Najadaceae, 5 Orchidadceae, 

1 Pontederiaceae, 1 Smilaceae, and 1 Typhaceae. 

The dicots include 88 families. Twelve families 

had 53 percent of the species: (1) 20 Boraginaceae, 

(2) 23 Compositae, (3) 20 Convolvulaceae, (4) 

34 Euphorbiaceae, (5) 71 Leguminosae, (6) 31 

Malvaceae, (7) 14 Myrtaceae, (8) 11 Polygonaceae, 

(9) 19 Rubiaceae, (10) 11 Sapindaceae, (11) 12 

Solanaceae, and (12) 16 Verbenaceae. 

Fauna on the Refuges 

Several studies of fauna, some dating back to 

the early 20th century, have been carried out on the 

refuges. Although mainly interested in birds, Danforth 

(1926, 1931) also observed other animal groups, 

particularly those associated with the avifauna. 

Mammals

Eight species of mammals, other than domestic 

livestock and household pets, have been reported 

at Laguna Cartagena: Herpestes auropunctatus

(mongoose), two bats, two rats, one mouse, and 

two monkeys (table 5). The most easily seen is the 

mongoose, which is widespread. Originally introduced 

from Asia between 1877 and 1879 to control rats in 

sugar cane fields, the mongoose is now Puerto Rico’s 

most detrimental predator on ground-nesting birds and 

lizards (Raffaele 1983). At Cartagena Lagoon during 

the 1920s, mongooses were seen eating eggs from 

nests and attacking waterfowl (Danforth 1926). They 

also consume frogs, blind snakes, lizards, and several 

species of invertebrates, including ants (Nellis 1999, 

Wolcott 1953).

Mongoose populations are greatest in coastal 

grassy areas near small streams and low in forested 

areas even when water is available (Pimentel 1955). 

A study in Guánica Forest concluded that the 

mongoose and Caprimulgus noctitherus (Puerto Rican 

nightjar) were separated by habitat requirements—the 

mongoose was more prevalent in coastal scrub below 

75 m elevation and the nightjar at higher elevations 

(Vilella and Zwank 1993a) (Pine 1980).

In the past, rats fed mainly on sugar cane but also 

ate bird eggs in and around the refuges. Elimination of 

cane production from surrounding areas has probably 

reduced rat populations, and possibly mongoose 

numbers. Mice, in contrast, feed mainly on seeds and 

fruits. Of the 14 species of bats endemic to Puerto 

Rico (Pine 1980), only 2 species have been observed 

on the Cabo Rojo Headquarters tract (table 5). 

In 1960, the small islands of Isla Cueva and Isla 

Guayacán near Parguera were acquired from the 

government of Puerto Rico for research on monkeys 
22Table 5—Animal species other than birds at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in 

southwestern Puerto Rico a b

Refuges c

Common name Cabo Rojo

 Laguna

Cartagena Other Reserves d

Group and species English Spanish H S L T A B C D E

Mammals

Native—bats

Artibeus jamaicensis Jamaican fruit-eating bat Murciélago frutero de Jamaica D

Brachyhylla cavernarum Antillian fruit-eating bat Hocico de cerdo D

Erophylla sezekoini Brown flower bat Murciélago de las flores D

Molossus molossus fortis Velvety free-tailed bat Murciélago de techos X

Monophyllus redmani Greatre Antillian long-tailed bat Murciélago lenguilargo D

Mormoops blainvilli Antillian ghost-faced bat Barbicacho D

Noctilio leporinus Greater bulldog bat Murciélego pescador A ENyctinomus murinus e Brazilian free-tailed bat Murciélago de viejo X

Introduced—various

Erythrocebus patas Patas monkey Mono patas X X

Macaca mulatta Rhesus monkey Mono macaca X E

Herpestes auropunctatus Mongoose Mangosta X X X X A C D

Mus musculus House mouse Ratón X

Rattus norvegicus House rat Rata común XR. rattus Roof rat Rata negra X A

Reptiles

Lizards

Ameiva exsul Puerto Rican ground lizard Siguana común X A B D EA. wetmorei Blue tailed ground lizard Siguana de rabo azul X B D E

Anolis cooki Dry land anole Legartijo del seco X A B D EA. cristatellus Puerto Rican crested anole Lagartijo común X A B D EA. ponsensis Ponce’s garden lizard Lagartijo jardinero X B D

A. pulchellus Grass anole Lagartijo rayon X A B D

A. stratulus Spotted lizard Lagartijo manchado X B D

Geckos

Phyllodactylus wirshingi NA f D

Sphaerodactylus macrolepsis Common gecko Salamanguita común X

S. roosevelti Roosevelt’s dwarf gecko D

S. nicholsi nicholsi Nichol’s dwarf gecko Salamanguita pigmea X D E

continued
23

Table 5—Animal species other than birds at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in southwestern Puerto Rico a b (continued)

Refugesc

Common name Cabo Rojo

 Laguna

Cartagena Other Reserves d

Group and species English Spanish H S L T A B C D E

Reptiles (continued)

Turtles

Dermochelys coriacea Leatherback turtle El tinglar XEretmochelys imbricata Hawksbill turtle El carey XTrachemys stejnegeri West Indian slider La jicotea X A

Snakes

Epicrates inoratus Puerto Rican boa Culebron D

Typhlops richardi Blind snake Vibora toro X

Amphibians

Bufo marinus Marine toad Churi X X A

Eleutherodactylus antillensis Puerto Rican red-eyed frog Coqui churi X A D

E. coqui Coqui Coqui común X A

Leptodactylus albilabris White-lipped frog Ranita de labio blanco X X APelthophryne lemur Puerto Rican crested toad Sapo concho D

Rana catesbiana Bull frog Sapo toro X

Fish

Anguilla rostrata Eel Anguilla X

Awaous taiasica Striated river goby Saga XBathygobius soporator Frillfin goby Gobio mapo XCaranx latus Horse eye jack Jurel, Ojon X ACentropomidae Snook family Familia de Robalos X

Centropomus parallelus Fat snook Robalo blanco X

Centropomus undecimales Common snook Robalo ADormitator maculatus Fat sleeper Mapiro XEleotris pisonis Spiny cheek sleeper Moron XGerres cinereus Yellow fin mojarra Mojarra C

Gobiomorus dormirtor Bigmouth sleeper Guavina X

Hypostomus plecostamus Plecostomus Pez gato XIctalurus nebulosus Brown bullhead Barbudo XLepomis gibbosus Pumpkin seed Chopa XL. macrochirus Bluegill Chopa XLiposarcus multiradiatus g Armored sailfin catfish Pleco? X

continued
24

Table 5—Animal species other than birds at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in southwestern Puerto Rico a b (continued)

Refuges c

Common name Cabo Rojo

 Laguna

Cartagena Other Reserves d

Group and species English Spanish H S L T A B C D E

FIsh (continued)

Megalops atlanticus Tarpon Sabalo X A

Mugilidae Mullet family Familia de Jareas o Lizas X A

Mugil curema Mullet Jarea A C

Poecilia reticulata Guppy Gupi X

P. vivipara Top minnow Cisi XTilapia aurea Tilapia Tilapia XT. mossambica Tilapia Tilapia mozambica X A C

a Sources: Danforth (1926), Diaz-Soltero (1990), Joglar (1998), Negrón González (1986), Rivero (1978). Domestic animals are omitted.bSources (fish names): Erdman (1972), Erdman and others (1985).c Cabo Rojo Refuge, with tracts; H = Cabo Rojo Headquarters; S = Salinas. Laguna Cartagena Refuge, with tracts; L = Lagoon; T = Tinaja. d Other reserves and information sources: A = Boqueron Wildlife Refuge (Chabert and others 1982, Negrón González 1986). Fifteen species of fish were mentioned for the lagoon but only 

six were identified to species. B = Los Morrillos de Cabo Rojo (Municipio Autonomo de Cabo Rojo 1998); C = Punta Guaniquilla Natural Reserve (Fuentes Santiago and Quevedo Bonilla 2002); D = Guánica Forest (Canals Mora 1990, Conde Costas y González 1990, Genet 2002, Rivero 1978); E = Parguera Natural Reserve (Departamento de Recursos Naturales 1981; Departamento de Recursos Naturales y Ambientales 2000a: Ventosa-Febles and others 2005).

e Now called Tadarida brasiliensis.f NA = not known.g Cartagena Lagoon (identified by Carlos Santos, University of Puerto Rico).
25

Troupial. Icterus icterus (troupial) is a permanent resident introduced 

from South America, commonly seen in southwestern Puerto Rico 

between Ponce and Boquerón. (U.S. Fish and Wildlife Service photo) 

(Vandenbergh 1989). Before its closure in 1982, 

some Erythrocebus patas and Macaca mulatta

(patas and rhesus monkeys) kept at the La Paguera 

Primate Facility escaped into the surrounding area 

(González-Martínez 1990). The rhesus monkeys, 

excellent swimmers, migrated into the Sierra Bermeja 

range where during the 1980s observers Phoebus and 

others (1989, page 157) noted “the escapees and their 

offspring are now doing quite well, living on several 

large adjoining cattle ranches” (Phoebus and others 

(1989, p. 157). Rhesus monkeys have been recorded at 

Laguna Cartegena and favor the fruits of Clusia rosea

and Hymenacea coubaril. In 1966, rhesus monkeys 

were also released on the Isla Desecheo NWR (Evans 

1989). By 1970, they were recognized as a serious 

threat to native fauna and flora, feeding on sea bird 

eggs and native vegetation. The largest known colony 

of Sula leucogaster (brown booby) was decimated 

and the Sula sula (red-footed booby) colony also 

declined in numbers, as a result of monkey predation. 

In addition, the monkeys consumed Bursera simaruba

wood pulp for water and used local cactus plants as a 

secondary source of moisture and nutrition. Not easily 

seen by visitors, they disperse into small groups to 

optimize foraging. 

In the late 1990s, about 120 patas monkeys ranged 

freely throughout 125 km2 of southwestern Puerto 

Rico where they were recorded on both refuges 

(González-Martínez 1998). At that time, the patas 

roamed in four heterosexual groups and several 

all-male bands, traveling a minimum daily distance 

of 0.8 to 2.0 km. They depend heavily on five 

introduced tree species: (1) Leucaena leucocephala, 

(2) Mangifera indica, (3) Melicoccus bijugatus, (4) 

Prosopis juliflora, and (5) Tamarindus indica along 

with the native Bourreria succulenta. They also eat 

the fruits of Stenocereus hystrix and Bromelia penguin

as well as grasshoppers and dung beetles (GonzálezMartínez 1990). Fruits of Bursera simaruba and 

Spondias mombin are eaten by both rhesus and patas 

monkeys. All monkeys are a threat to public health, 

damage crops, prey on bird eggs, and have spread in 

the southwest. If they reach other parts of the island 

they could become virtually impossible to control. 

Birds

Birds include resident, migrant, vagrant, and 

introduced species. West Indian bird communities 

have higher proportions of frugivores than mainland 

communities; the arrival of migrant parulids during 

winter months, however, reverses the pattern (Faaborg 

and Terborgh 1980). Moreover, small gleaning 

insectivores that spend the winter on the islands are 

overrepresented whereas hawks and large, gleaning 

insectivores, frugivores, and granivores are 

either absent or underrepresented (Faaborg and 

Terborgh 1980). 

Of the 212 species sighted on the refuges, 12 are 

endemic, 25 are listed as game or protected game 

species, and 14 have been introduced as game or for 

other purposes. Moreover, seven species are listed 

as threatened or endangered or as candidates for 

listing by the F&WS, five are listed as threatened by 

the Commonwealth of Puerto Rico, and two species 

are listed by both. Finally, 13 are target species for 

management at Cartagena Lagoon (table 6, top). 

Raffaele’s (1983, 1989) compendia of Puerto Rican 

bird species, based on nearly 7 years of observations, 

disclosed that about one-half of the island’s 269 

recorded species, including exotics, had been recorded 

at one site—Cartagena Lagoon. Another 22 bird 
26Table 6—Bird species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in southwestern Puerto Rico a

Refuges b

Common name Cabo Rojo

Laguna 

Cartagena Status c

Other

Reserves d

Genus and species English Spanish H S L T E N M/R GIP C/F C/T A B C D E

Species reported for Cabo Rojo and Laguna Cartegena Wildlife Refuges, as well as other reserves in southwestern Puerto RicoActitus macularius Spotted sandpiper Playero coleador X X X C M A B C D E

Agelaius xanthomas Yellow-shouldered blackbird Mariquita X X X E U R F T A B D E

Amandava amandava Strawberry finch Chamorro fresa X X C R A

Ammodramus savannarum Grasshopper sparrow Gorrión chicharra X X C R CT A

Anas acuta Northern pintail Pato pescuecilargo X U M G A D

Anas americana American wigeon Pato cabeciblanco X C M G A CAnas bahamensis White-cheeked pintail Pato quijada colorada X X C R P F CT A C D

Anas clypeata Northern shoveler Pato cuchareta X U M G A

Anas crecca Green-winged teal Pato aliverde X R M G A

Anas discors Blue-winged teal Pato zarcel X X C M G T A C D

Anas penelope Eurasian wigeon Silbón Europeo X R M

Anas platyrhynchos Mallard Pato cabeciverde X R M G

Anas rubripes American black duck Pato oscuro X A M G

Anous stolidus Brown noddy Cervera parda X C R

Anthracothorax dominicus Antillean mango Zumbador dorado X X X C R C A C D

Anthracothorax viridis Green mango Zumbador verde X E U R A B D

Ardea alba Great egret Garza real X C R A B C D E

Ardea herodias Great blue heron Garzón cenizo X X X C MR A B C D E

Arenaria interpres Ruddy turnstone Playero turco X C M A C D E

Asio flammeus Short-eared owl Múcaro real X X U R C A D

Aythya affinis Lesser scaup Pato pechiblanco X U M G A

Aythya collaris Ring-necked duck Pato acollarado X U M G A

Bartramia longicauda Upland sandpiper Playero pradero X R M

Botaurus lentiginosus American bittern Yaboa americana X R M C A

Bubulcus ibis Cattle egret Garza ganadera X X X X C R A B C D E

Buteo jamaicensis Red-tailed hawk Guaragua colirrojo X X X X C R A C D E

Butorides virescens Green heron Martinete X X X C R A B C D E

Calidris alba Sanderling Playero arenero X C M D

Calidris alpina Dunlin Playero espaldicolorado X R M

Calidris canutus Red knot Playero gordo X X U M C

Calidris fuscicollis White-rumped sandpiper Playero de rabadilla blanca X X X U MCalidris himantopus Stilt sandpiper Playero patilargo X X C M C A C D E

Calidris mauri Western sandpiper Playero occidental X X X C M C A D E

Calidris melanotos Pectoral sandpiper Playero pectoral X X X C M A D

Calidris minutilla Least sandpiper Playero menudillo X X X C M A C E

continued
27

Table 6—Bird species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in southwestern Puerto Rico a (continued)

Refuges b

Common name Cabo Rojo

Laguna 

Cartagena Status c

Other

Reserves d

Genus and species English Spanish H S L T E N M/R GIP C/F C/T A B C D E

Calidris pusilla Semipalmated sandpiper Playero gracioso X X X C M C A D ECaprimulgus carolinensis Chuck-wills-widow Guabairo de las Carolinas X U M D

Caprimulgus noctitherus Puerto Rican nightjar Guabairo de Puerto Rico X E R R F D ECathartes aura Turkey vulture Aura tiñosa X X X C R I A B C D E

Catharus fuscesens Veery Zorzalito rojizo X A T

Catharus ustulatus Swainson’s thrush Zorzal de Swainson X A T D

Charadrius alexandrinus Snowy plover Chorlito blanco X U R C B

Charadrius melodus Piping plover Chorlito melódico X R M FCharadrius semipalmatus Semipalmated plover Chorlito acollarado X X C M A C D ECharadrius vociferus Killdeer Chorlito sabanero X X X C R A B D

Charadrius wilsonia Wilson’s plover Chorlito marítimo X X C R C A B C D E

Chlidonias niger Black tern Fumarel común X X C M

Chlorostilbon maugaeus Puerto Rican emerald Zumbadorcito de Puerto Rico X X X E C R C D

Chordeiles gundlachii West Indian nighthawk Querequequé antillano X X X U MR A D

Chordeiles minor Common nighthawk Querequequé migratorio X R M B D

Circus aeruginosus Western marsh harrier Aguilucho lagunero occidental X A M

Circus cyaneus Northern harrier Gavilán de ciénaga X X X U M A

Coccyzus americanus Yellow-billed cuckoo Pájaro bobo piquiamarillo X X X U MR A D

Coccyzus minor Mangrove cuckoo Pájaro bobo menor X X X C R A B C D E

Coccyzus vieilloti Puerto Rican lizard cuckoo Pájaro bobo mayor X E C R C DCoereba flaveola Bananaquit Reinita común X X X C R A B C D E

Columba livia Rock pigeon Paloma domestíca X X C R C D

Columbina passerina Common ground dove Rolita X X X C R P A B C D E

Contopus latirostris Lesser Antillean pewee Bobito antillano menor X X X C R C A B D E

Contopus virens Eastern wood peewee Pibi oriental X R MT

Crotophaga ani Smooth-billed ani Judio X X X C R A B C D E

Cypseloides niger Black swift Vencejo negro X U R C D

Dendrocygna arborea West Indian whistling-duck Chiriría caribeña X R R P C CT A C

Dendrocygna autumnalis Black-bellied whistling-duck Chiriría pinta X A MDendrocygna bicolor Fulvous whistling-duck Pato bicolor X U MR P A

Dendroica adelaidae Adelaide’s warbler Reinita mariposera X X X E C R C B C D E

Dendroica caerulescens Black-throated blue warbler Reinita azul X C M C C D

Dendroica coronata Yellow-rumped (myrtle)warbler Reinita coronada X X X C M A D

Dendroica discolor Prairie warbler Reinita galana X X X C M C A C D

Dendroica dominica Yellow-throated warbler Reinita gargantiamarilla X U M A D

Dendroica magnolia Magnolia warbler Reinita manchada X X R M A C D

Dendroica palmarum Palm warbler Reinita palmera X U M A C D

continued
28Table 6—Bird species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in southwestern Puerto 

Rico a (continued)

Refuges b

Common name Cabo Rojo

Laguna 

Cartagena Status c

Other

Reserves d

Genus and species English Spanish H S L T E N M/R GIP C/F C/T A B C D E

Dendroica pensylvania Chestnut-sided warbler Reinita flaquicastaña X X R M D

Dendroica petechia Yellow warbler Canario de mangle X X X C R C A B C D E

Dendroica striata Blackpoll warbler Reinita rayada X X CU MT A DDendroica tigrina Cape May warbler Reinita tigre X X X C M A D

Dendroica virens Black-throated green warbler Reinita verdosa X R M DDolichonyx oryzivorus Bobolink Chambergo X U M

Egretta caerulea Little blue heron Garza azul X X X C R A C D E

Egretta rufescens Reddish egret Garza rojiza X X R M

Egretta thula Snowy egret Garza blanca X X X C R A C D E

Egretta tricolor Tricolored heron Garza pechiblanca X X C R A B C D E

Elaenia martinica Caribbean elaenia Juí blanco X X X C R C D

Empidonax virescens Acadian flycatcher Mosquiro verdoso X A T

Estrilda melpoda Orange-cheeked waxbill Veterano X X X C R I A B C D E

Estrilda troglodytes Black-rumped waxbill Veterano orejicolorado X R R I D

Eudocimus albus White ibis Ibis blanco X R T

Euphonia musica Antillian euphonia Jilguero X X R R D

Euplectes afer Yellow crowned bishop Napoleón tejedor X U R I

Euplectes francicanus Orange bishop Osbispo anaranjado X U R I D

Falco columbarius Merlin Falcón migratorio X X X U M I A C D E

Falco peregrinus Peregrine falcon Halcón peregrino X X X U M F C C D

Falco sparverius American kestrel Fálcon común X X X C R A B C D E

Fregata magnificens Magnificent frigatebird Tijereta X X C R C A B C D E

Fulica caribaea Caribbean coot Gallinazo Caribeño X U R P CF CT A

Fulica americana American coot Gallinazo americano X C MR P A DGallinago delicata Wilson’s snipe Becasina X X X C M G T A

Gallinula chloropus Common gallinule Gallareta común X X X C R G T A B C D

Gallus gallus Red junglefowl Gallo/Gallina silvestre X X C R I D

Geothlypis trichas Common yellowthroat Reinita picatierra X X X C M AGeotrygon montana Ruddy quail-dove Paloma perdiz rojiza X U R P C D

Haematopus palliatus American oystercatcher Ostrero caracolero X U R C DHimantopus mexicanus Black-necked stilt Viuda X X X C R A B C D E

Hirundo rustica Barn swallow Golondrina horquillada X X X C M A D

Icterus dominicensis Greater Antillean oriole Calandria X X C R C A C

Icterus galbula Baltimore oriole Bolsero de Baltimore X X U M DIcterus icterus Troupial Turpial X X X C R I A B C D E

Iridoprocne bicolor Tree swallow Golondrina biclor X R M

Ixobrychus exilis Least bittern Martinetito X X C R C A C D E

continued
29

Table 6—Bird species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in southwestern Puerto Rico a (continued)

Refuges b

Common name Cabo Rojo

Laguna 

Cartagena Status c

Other

Reserves d

Genus and species English Spanish H S L T E N M/R GIP C/F C/T A B C D E

Jacana spinosa Northern jacana Jacana Centroamericana X A TLarus argentatus Herring gull Gaviota argéntea X X R MLarus atricilla Laughing gull Gaviota gallega X X X C R B D

Laterallus jamaicensis Black rail Gallito negro X R M CLimnodromus griseus Short-billed dowitcher Agujeta piquicorta X X X C M A C D E

Limosa fedoa Marbled godwit Barga canela X A MLimosa haemastica Hudsonian godwit Barga albiblanca X X R M

Lonchura cucullata Bronze mannikin Diablito X X X X C R I A C D

Lonchura malabarica Indian silverbill Gorrión picoplata X X X C R I A B D

Lonchura malacca Troicolored munia Monjita tricolor X X X C R I D

Lonchura punctulata Nutmeg mannikin Gorrión canela X X X X C R I B C D

Lophodytes cucullatus Hooded merganser Mergansa encapuchada X A MLoxigilla portoricensis Puerto Rican bullfinch Come ñame X X X X E C R A B C D E

Margarops fuscatus Pearly-eyed thrasher Zorzal pardo X X X X C R A C D E

Megaceryle alcyon Belted kingfisher Martín pescador norteño X X X C M A C D

Megascops nupides Puerto Rican screech-owl Múcaro común X X U R C A D

Melanerpes portoricensis Puerto Rican woodpecker Carpintero de Puerto Rico X X X X E C R A B C D

Mergus serrator Red-breasted merganser Mergansa piquilarga X A MMimus gundlachii Bahama mockingbird Ruiseñor de las Bahamas X A T

Mimus polyglottos Northern mockingbird Ruiseñor X X X X C R A B C D E

Mniotilta varia Black-and-white warbler Reinita trepadora X X X C M A C D

Molothrus bonariensis Shiny cowbird Tordo lustroso X X X X C R A B C D E

Myiarchus antillarum Puerto Rican flycatcher Juí de Puerto Rico X X X X C R C A B C D E

Nesospingus speculiferus Puerto Rican tanager Llorosa X E C R D

Numida meleagris Helmeted guineafowl Guinea torcaz X X U R G D

Nyctanassa violacea Yellow-crowned night-heron Yaboa común X X X C MR C A B C D E

Nycticorax nycticorax Black-crowned night-heron Yaboa real X X X U MR A D

Olor columbianus Whistling swan Cisne silbador X A MOporonis agilis Connecticut warbler Reinita de Connecticut X X R MOporonis formosus Kentucky warbler Reinita de Kentucky X R M D

Oporonis philadelphia Mourning warbler Reinita enlutada X X R M

Oxyura dominica Masked duck Pato enmascarado X R R P C CT A

Oxyura jamaicensis Ruddy duck Pato chorizo X C R P CF CT A C D

Pandion haliaetus Osprey Águila pescadora X X X C M A B C D E

Parula americana Northern parula Reinita pechidorada X X X X C M A C D

Passerina caerulea Blue grosbeak Azulejo X R M D

Passerina cyanea Indigo bunting Gorrión azul X X U M A D

continued
30Table 6—Bird species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in southwestern Puerto Rico a (continued)

Refuges b

Common name Cabo Rojo

Laguna 

Cartagena Status c

Other

Reserves d

Genus and species English Spanish H S L T E N M/R GIP C/F C/T A B C D E

Patagioenas leucocephala White-crowned pigeon Paloma cabeciblanca X X R R C A D

Patagioenas squamosa Scaly-naped pigeon Paloma turca X R R G D

Pelicanus occidentalis Brown pelican Pelícano pardo X X C R F A B C D E

Petrochelidon fulva Cave swallow Golondrina de cueva X X X C R A C D

Petrochelidon pyrrhonata Cliff swallow Golondrina de penasco X X R M

Phaethon lepturus White-tailed tropicbird Rabijunco X C R C D

Phalacrocorax olivaceus Olivaceous cormorant Cormorán oliváceo X A M

Pheucticus ludovicianus Rose-breasted grosbeak Piquigrueso X R M D

Phoenicopterus ruber American flamingo Flamenco X R T C EPiranga olivacea Scarlet tanager Piranga escarleta X R MT

Piranga rubra Summer tanager Piranga roja X R TPlegadis falcinellus Glossy ibis Ibis lustroso X R RT A D

Pluvialis dominica American golden plover Playero dorado X X R M

Pluvialis squatarola Black-bellied plover Playero cabezón X X C M A C D E

Podiceps dominicus Least grebe Tigua X X R R C CT A C

Podilymbus podiceps Pied-billed grebe Zaramago X C R P C A C

Porphyrula martinica Purple gallinule Gallareta azul X R R CT APorzana carolina Sora Gallito X R M APorzana flaviventer Yellow-breasted crake Gallito amarillo X U R C CTProgne cryptoleuca Cuban martin Golodrina Cubana X R T

Progne subis Purple martin Golondrina púrpura X R M

Progne dominicensis Caribbean martin Golondrina de iglesias X X X C R A B C D

Protonotaria citrea Prothonotary warbler Reinita anaranjada X X U M C A D E

Quiscalus niger Greater Antillean grackle Chango X X X C R A B C D E

Rallus longirostris Clapper rail Pollo de mangle X X C R A B D E

Riparia riparia Bank swallow Golondrina parda X X X C R A D

Seiurus aurocapilla Ovenbird Pizpita dorada X X X C M D

Seiurus motacilla Lousiana waterthrush Pizpita del río X X C M C A D

Seiurus noveboracensis Northern waterthrush Pizpita de mangle X X X C M C A B C D E

Setophaga ruticilla American redstart Candelita X X X C M A B D E

Sphyrapicus varius Yellow-bellied sapsucker Carpintero pechiamarillo X R M D

Spindalis portoricensis Puerto Rican spindalis Reina mora X X X X E C R A D E

Sterna anaethetus Bridled tern Gaviota monja X C R CSterna antillarum Least tern Gaviota pequeña X X U R C C A D E

Sterna dougalli Roseate tern Palometa X C R F D

Sterna fuscata Sooty tern Gaviota oscura X X C R D

Sterna hirundo Common tern Gaviota común X C MR A B D

continued
31

Table 6—Bird species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in southwestern Puerto Rico a (continued)

Refuges b

Common name Cabo Rojo

Laguna 

Cartagena Status c

Other

Reserves d

Genus and species English Spanish H S L T E N M/R GIP C/F C/T A B C D E

Sterna maxima Royal tern Gaviota real X C R A B C D

Sterna nilotica Gull-billed tern Gaviota piquigorda X X U M A

Sterna sandvicensis Sandwich tern Gaviota piquiaguda X X U R A B C

Streptopelia roseogrisea African collared-dove Tórtola collarina X X U R I D

Sula leucogaster Brown booby Boba prieta X C R D

Sula sula Red-footed booby Boba patiroja X C R C

Tiaris bicolor Black-faced grassquit Gorrión negro X X X X C R A C D E

Tiaris olivaceus Yellow-faced grassquit Gorrión barba amarilla X X X C R A B C D

Todus mexicanus Puerto Rican tody San Pedrito X X X X E C R A B C D E

Tringa flavipes Lesser yellowlegs Playero guineilla menor X X X C M A B C D E

Tringa melanoleuca Greater yellowlegs Playero guineilla mayor X X X C M A B C D E

Tringa semipalmatus Willet Playero albiblanco X X CU M A C D E

Tringa solitaria Solitary sandpiper Playero solitario X X C M C A D

Turdus plumbeus Red-legged thrush Zorzal de patas coloradas X X C R A C D E

Tyrannus caudifasciatus Loggerhead kingbird Clérigo X X X C R C D

Tyrannus dominicensis Gray kingbird Pitirre X X X C R A B C D E

Vermivora chrysoptera Golden-winged warbler Reinita alidorada X X R M DVermivora pinus Blue-winged warbler Reinita aliazul X R T

Vidua macroura Pin-tailed whydah Viuda colicinta X X U R I

Vireo altiloquus Black-whiskered vireo Julián chiví X X X C R C D

Vireo griseus White-eyed vireo Julián chiví ojiblanco X X X U M

Vireo latimeri Puerto Rican vireo Bien-te-veo X X X E C R C D EVireo olivaceus Red-eyed vireo Julián chiví ojirrojo X X R T D

Wilsonia canadensis Canada warbler Reinita de Canadá X A M

Wilsonia citrina Hooded warbler Reinata de capucha X X U M A D

Zenaida asiatica White-winged dove Tórtola aliblanca X X U R G A B C D E

Zenaida aurita Zenaida dove Tórtola cardosantera X X X C R G A B C D E

Zenaida macroura Mourning dove Tórtola rabilarga X X U R G D E

Zonotrichia albicollis White-throated sparrow Chingolo X A M

continued
32Table 6—Bird species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges and Commonwealth protected areas in southwestern Puerto Rico a (continued)

Refuges b

Common name Cabo Rojo

Laguna 

Cartagena Status c

Other

Reserves d

Genus and species English Spanish H S L T E N M/R GIP C/F C/T A B C D E

Species reported for other reserves in southwestern Puerto Rico but not recorded at Cabo Rojo or Laguna Cartegena Wildlife RefugesArchilochus colubris Ruby-throated hummingbird Zumbadorcito de garganta roja R M D

Brotogeris versicolurus White-winged parakeet Perquito aliblanco C R I D

Catharus bicknelli Bicknell’s thrush Zorzal de Bicknell R M C D

Catharus minimus Gray-cheeked thrush Zordal de mejilla gris R M D

Colinus virginianus Northern bobwhite Codorniz R R I A

Dendroica castanea Bay-breasted warbler Reinita castana R M D

Dendroica fusca Blackburnian warbler Reinita de fuego R T D

Eulampis holosericeus Green-throated carib Zumbador pechiazul R R D

Geotrogon chrysia Key West quail-dove Paloma perdiz áurea U R C D

Geotrygon mystacea Bridled quail-dove Paloma perdiz de Martinica R R C D

Helmitheros vermivorum Worm-eating warbler Reinita gusanera U M C D

Larus delawarensis Ring-billed gull Gaviota piquianillada C M ALomnothlypis swainsonii Swainson’s warbler Reinita de Swainson R M D

Myopsitta monachus Monk parakeet Perico monje U R I A B

Numenius phaeopus Hudsonian curlew Playero pico corvo R M A EOrthohyncus cristatus Antillean crested hummingbird Zumbadorcito crestado C R D

Passer domesticus House sparrow Gorrion ingles U R I D

Phalaropus lobatus Red-necked phalarope Faloropo picofino A M C

Stercorarius pomarinus Pomarine jaeger Pagalo pomarino R M E

Streptopelia decaocto Eurasian collared-dove Tórtola turca I D

Tachornis phoenicobia Antillean palm swift Vencejito A D

Wilsonia pusilla Wilson’s warbler Reinita de Wilson A M D

a Sources: Species list and information derived from several sources (Chabert Llompart and others 1986; Danforth 1926, 1929, 1931, 1936; Diaz-Soltero 1990; Hernández Prieto 1990; Kepler 

1971, 1972; McCandless 1961, 1962; Moreno 1997; Ortiz Rosas 1981; Raffaele 1983; Toro McCown and Chabert 1986; U.S. Fish and Wildlife Service 2002, 2004; Wetmore 1927; Wiley 1974, see endnotes). Some species listings are from historical records. 

b Cabo Rojo Refuge, with tracts; H = Cabo Rojo headquarters; S = Salinas. Laguna Cartagena Refuge, with tracts; L = Lagoon; T = Tinaja. Species present are indicated with an X. c Status: E = endemic; N = numbers in Puerto Rico (C = very common or common; U = uncommon; R = rare to extremely rare; A = accidental) (Raffaele 1983); M/R: M = migrant, transient 

(T), or visitor and R = resident or resident breeder; GIP: G = game species, I = introduced as game or other purpose, P = protected game species; C/F (listed species): (C = threatened or endangered by Puerto Rico; F = listed or candidate for listing by U.S. Fish and Wildlife Service); C/T (species of concern): C= conservation concern, local or regional; T = target species for managment at the Cartagena Lagoon; blank space = no detail.

d Reserves and information sources: A = Boqueron Wildlife Refuge (Chabert and others 1982); B = Los Morrillos de Cabo Rojo (Municipio Autonomo de Cabo Rojo (1998); C = Reserva 

Natural Punta Guaniquilla (Fuentes Santiago and Quevedo Bonilla 2002); D = Guánica Forest (Arendt and others 2007; Hernández Prieto 1990); E = Parguera Natural Reserve (Departamento de Recursos Naturales 1981, Departamento de Recursos Naturales y Ambientales 2000a; Ventosa-Febles and others 2005).
33

species not yet recorded at the refuges have been 

reported on five nearby reserves in southwestern 

Puerto Rico (table 6, bottom; fig. 1). 

The salt flats at Cabo Rojo are among the best 

sites to attract birds migrating from North and South 

America through the eastern Caribbean. The fall 

bird migration may be divided into three events with 

some overlap: (1) shorebirds dominate from July to 

mid-September, (2) warblers from September to midOctober, and (3) migrant waterfowl in late October 

and November (Collazo and others 1995, McCandless 

1961). Data recorded at the salt flats showed: 

•	 More than 25 species of shorebirds

•	 As many as 40,000 shorebirds migrating through

the salt flats during the fall months

•	 Daily counts in the fall sometimes exceeding

7,000 birds, with historic daily counts as high 

as 10,000 birds 

•	 A small breeding population of Charadrius 

alexandrinus (snowy plover) not found elsewhere 

in Puerto Rico 

In 1900, “Christmas counts” began in the United 

States as a means to protect bird species from 

traditional holiday hunts. In 1921, Stuart Danforth 

began Christmas bird counts in Mayagüez, PR, when 

Aeglaius xanthomus (yellow-shouldered blackbird) 

comprised 62 percent of the 720 birds sampled. In 

1925 and 1927, the count was done at Cartagena 

Lagoon, and the 1926 and 1928 counts were between 

Guánica and Boquerón (McCandless 1962). Results 

were published in the local Journal of Agriculture. 

Counts between 1955 and 1960 were carried out 

by McCandless and colleagues (Burden 1961) and 

included the Cartagena Lagoon and Cabo Rojo 

Lighthouse, as have all subsequent counts from 1972 

Ducks. Anas discors (blue-winged teal) is a common wintering duck in 

Puerto Rico that is found on several types of water bodies, including 

freshwater lagoons and hypersaline salt ponds. The species may 

be observed throughout the year but is most common from October 

through April. (U.S. Fish and Wildlife Service photo) 
34

to the present (figs. 9A, 9B, and 9C). In 1955 and 

1956, about one-quarter of the birds tallied were 

Bubulcus ibis (cattle egrets). 

The earliest counts were undertaken by Danforth 

alone, or in 1928, with the help of the well-known 

ornithologist James Bond (McCandless 1962). The 

1955 to 1959 counts were carried out by McCandless 

with fewer than 10 assistants. The Danforth counts are 

not comparable with later work because of the lack of 

field assistance and travel difficulties. 

The 31-year record between 1972 and 2002 showed 

a mean of 9,203.5 ± 795.0 (range = 996 to 20,475) 

individuals of 97.8 ± 2.2 species (range = 65 to 112) 

tallied by an average of 21.3 ± 2.0 (range = 3 to 44) 

observers (Puerto Rican Ornithological Society 2006, 

see endnotes).The greatest number of species tallied 

was in 1972 and 1974 and the fewest in 1978. It is 

noteworthy that the lowest number of species tallied 

was between 1976 and 1979, a period when drought 

may have negatively impacted bird species at nearby 

Guánica Forest (Faaborg and Arendt 1989a, 1989b, 

1990). This same period also coincides with the 

fewest observers participating in the count. 

Early in the 20th century, sugar cane fields 

on the south coast were largely devoid of tree 

cover (Wetmore 1916). At that time, Cartagena 

Lagoon was considered to be Puerto Rico’s most 

important breeding ground for resident and migrant 

waterfowl (Danforth 1926). During nearly 2 years of 

observations, Danforth identified 105 resident and 

migratory bird species and described 8 vegetation 

types. He feared that the lagoon would be drained and 

that some species would be lost or decline in numbers. 

He suggested protective measures to increase bird 

numbers and diversity and pointed out the need for 

public education. His interest in government control 

Falcon. Falco sparverius (American kestrel) is a locally common 

permanent resident in Puerto Rico that prefers coastal areas with large 

trees for nesting. It is often seen on exposed perches in open areas from 

which it hunts. (U.S. Fish and Wildlife Service photo) 
35

120

100

80

60

40

20

0

Species (number)

Individual birds

(thousand) Participants (number)

1920 1940 1960 1980 2000 2020

1920 1940 1960 1980 2000 2020

1920 1940 1960 1980 2000 2020

25

20

15

10

5

0

50

45

40

35

30

25

20

15

10

5

0

Year

Year

Year

(A)

(B)

(C)

Figure 9—Christmas bird counts: (A) number of species, (B) number of individual birds, 

and (C) number of participants.

of the lagoon was realized many years later. Only 

recently, sugar cane production discontinued north 

of the Cartagena Lagoon. Today, the land is used as 

pasture for livestock. 

Reptiles and Amphibians

Eleven reptiles and five amphibians have been 

reported as terrestrial residents on the refuges: seven 

lizards, one land turtle, two geckos, one blind snake, 

one toad, and four frogs. In addition, two sea turtles, 

Eretmochelys imbricata and Dermochelys coriacea, 

briefly use the beaches at Cabo Rojo for laying 

eggs (table 5). 

Ameivas, commonly called iguanas (or preferably 

siguanas), actively forage on the ground, rarely 

ascending trees. Ameiva exsul, the most common 

and widely distributed siguana in Puerto Rico, is 

found in coastal areas to about 350 m in elevation, 

and on offshore islands (Rivero 1978). Ameiva 

wetmorei, a very attractive siguana that is rarely 
36

Ground lizard. Ameiva wetmorei (blue-tailed ground lizard) is uncommon 

at higher elevations on the Tinaja tract of the Laguna Cartagena National 

Wildlife Refuge. (U.S. Fish and Wildlife Service photo)

seen, is confined to the open, deciduous forest and 

scrub of southwestern Puerto Rico. During 1920, 

Ameiva exsul, Anolis cristatellus, Anolis pulchellus, 

and Anolis stratulus were all reported for the Lagoon 

tract (Danforth 1926). On the Tinaja tract, Ameiva 

wetmorei is found at higher elevations and Anolis 

poncensis on the lower slopes. The only amphibian 

reported then was the frog, Leptodactylus albilabris,

from the Cartagena Lagoon. Leptodactylus albilabris

is seldom found distant from wetlands, including 

streams. Tadpoles and small frogs are eaten by 

the lagoon waterfowl, particularly the herons 

(Danforth 1926). 

Of the eight species of anoles found in Puerto 

Rico, five have been recorded in grass and bushes 

on the refuges (Rand 1964, Rivero 1978). Anolis 

cooki occurs in disjunct populations between Cabo 

Rojo and Guayanilla on the south coast. Currently, 

it is endangered due to habitat changes and possibly 

competition from other lizards. Anolis cristatellus

occurs all over Puerto Rico up to 550 m in elevation. 

It commonly enters houses. Anolis poncensis is found 

along the dry south coast, mainly from Cabo Rojo 

to the town of Salinas east of Ponce. It is often seen 

in low bushes and cactus. Anolis pulchellus, also 

common, prefers grasses in open areas. Sometimes it 

climbs low bushes but is never found in trees. Anolis 

stratulus is widely distributed in the lowlands, and 

common at intermediate elevations in the mountains. 

Two geckos of the genus Sphareodactylus are 

found in the refuges. Sphareodactylus macrolepsis is 

the most common and widely distributed species of 

the genus, with subspecies occurring from the coast 

to 850 m in elevation (Rivero 1978). Sphareodactylus 

nicholsi prefers dry habitat, extending along the north 

coast from the town of Manati west to Mayagüez and 

south and east to Ponce and Juana Díaz (Rivero 1978). 

It frequents open deciduous forest where it forages 

among the leaves. 

Trachemys stejnergeri (previously Pseudemys 

terrapen) is Puerto Rico’s only native turtle. It 

occupies lagoons and streams in the lowlands where 

it feeds on plants and animals. Typhlopids are 

blind snakes, also called worm snakes, which live 

underground. Typhlops richardi feed on invertebrates 

including ants and termites. 

To most islanders, the amphibian fauna of Puerto 

Rico includes the group commonly known as “coquis” 

in the genus Eleutherodactylus, and Bufo marinus, the 

common toad (Rivero 1978). The toad was imported 

from Jamaica and Barbados during the 1920s to 

control a grub on sugar cane. A few years later, it 

was abundant. Rana catesbiana, the bullfrog, was 

introduced for similar purposes in 1935 and since has 

spread in coastal areas to 150 m in elevation. It prefers 

wetlands including lagoons and streams. 

Two species of Eleutherodactylus, both widely 

distributed on the island, have been identified on the 

refuges (table 5). Eleutherodactylus coqui, the best 

known, climbs into tree branches at night whereas 

Eleutherodactylus antillensis calls from low branches 

and bushes. Insectivorous birds are common predators 

of frogs; moreover, ameivas, anoles, and several 

invertebrates may also eat young amphibians 

(Rivero 1978). 

Fish

Four species of fish were reported for Cartagena 

Lagoon during the 1920s (Danforth 1926) (table 5). 

At that time, when the lagoon waters evaporated 

during June and July, local fishermen speared 

Anguilla rostrata. Some of the eels were in excess 

of 1 m in length. Smaller eels served as food for the 

Ardea herodias (great blue heron) (Danforth 1926) 

and probably other wading bird species as well. 

Among the fish species, Centropomus parallelus were 

reported by local farmers. Individuals of Dormitator 
37

Butterflies. Sunning themselves in the early morning hours, these 

Heliconius charitonia (zebra butterflies) are occasionally seen along 

the dirt road and in the arroyos of the Tinaja tract. 

(Photo by Peter L. Weaver)

maculates, rather scarce, were usually observed 

among the cattails where they would bury themselves 

in the mud when water levels were low. Poecilia 

vivipara were most abundant, often seen in great 

numbers swarming in shallow water. The latter fish 

species was the only one used to any extent as food by 

the birds.

Additional fish surveys were carried out in 1957, 

1980, and 1985 (Díaz-Soltero 1990). At least 17 

species were recorded one or more times (table 5). 

Moreover, unknown individuals called “pez grande” 

(big fish) by local folks were reported to have lived 

in the mud and attained a weight of 9 kg (Danforth 

1926). The population was apparently exterminated 

when the lagoon dried up in 1923. 

Other Flora and Fauna

A compendium of plant, animal, and fungi species 

was listed in the plan for the development and 

management of Cartagena Lagoon (Díaz-Soltero 

1990). The tables derived from several unpublished 

reports included 26 species of algae, 8 species 

of fungi, 23 species of protozoans, 11 species of 

metazoans, and 96 species grouped as “microbiota.” 

Also tallied were 2 species of Platyhelminthes, 

6 species of Mollusca, 5 species of Annelida, 5 

species of Arthropoda (Crustacea), and 27 species 

of Arthropoda (Insecta). In a study of the lagoon’s 

water quality, 67 insect species belonging to five 

orders—Hemiptera, Diptera, Coleoptera, Odontata, 

and Ephemeroptera—were collected (Deliz Quiñones 

2005). Six families and twenty-four species of 

butterflies were tentatively identified on both refuges 

(Asencio López and Rentas Soto 2006, see endnotes; 

Haselmayer 2000, see endnotes; Hernández 2004; 

Ramos 1982; Smith and others 1994) (table 7). 

At the Salinas tract, the water boatmen and 

brine shrimp found in Candeleria and Fraternidad 

lagoons serve as prey for numerous shorebirds 

(Bonilla Soto 1984, Tripp and Collazo 2003). The 

lagoons are also the habitat for two new, extremely 

halophilic species of bacteria—Halogeometricum 

borinquense (Montalvo-Rodríguez and others 

1998); and Haloterrigena thermotolerans, a highly 

thermo-tolerant Archaeon belonging to the order 

Halobacterias (Montalvo-Rodríguez and others 2000). 

Also, three strains of halophilic Archaebacteria were 

reported (Montalvo-Rodríguez and others 1997). 

Finally, about 100 species of parasites found on 63 

species of freshwater sport fishes were identified 

(Bunkley-Williams and Williams 1994). The fish are 

commonly caught in the island’s reservoirs, streams, 

and lagoons, including those in and around the refuges 

and other protected areas. 

Environmental Issues

The need to increase, improve, and protect nesting 

and foraging habitat for native forest birds, shorebirds, 

and wetland species, including threatened and 

endangered species, was indicated in the mid-1970s 

(Raffaele and others 1977, Woodbury and others 

1975). In 1979, after 6 years of investigation, several 

of Puerto Rico’s critical wildlife areas were identified 

including the Boquerón Commonwealth Forest, the 

Boquerón Wildlife Refuge, the Cabo Rojo Salt Flats 

and cliffs, the Cartagena Lagoon, Punta Guaniquilla 

Natural Reserve, and the Guánica Forest (Department 

of Natural Resources 1979). At the beginning of the 

new millennium, the importance of species and habitat 

protection remain as critical issues (Núñez-García and 

Hunter 2000, U.S. Fish and Wildlife Service 2002, 

Ventosa-Febles 2005). A number of factors indicate 

the need for additional habitat in the Caribbean 

Islands NWR Complex, including the refuges in 

southwestern Puerto Rico: 
38

Table 7—Butterfly species found on the Cabo Rojo and Laguna Cartagena National Wildlife Refuges a

Names

Scientific—family species English Occurrence b

Danaidae

Danaus plexippus Monarch Common in open areas

Heliconidae

Agraulis vanillae Gulf fritillary Fairly common in open areas

Heliconius charitonia Zebra Uncommon in arroyos

Hesperidae

Cymaenes tripunctus Three-spotted skipper Abundant in open areas

Pyrgus oileus oileus Tropical checkered skipper Fairly common in open areas

Ubanus proteus Long-tailed skipper Very common everywhere in September

uncommon after mid-October

Lycaenidae

Hemiargus hanno watsoni Hanno blue

Strymon columella Hairstreak Common in open areas

Nymphalidae

Anartia jatrophae Peacock Fairly common everywhere

Biblis hyperia Red rim Fairly common near arroyos

Ephyriades arcas Skipper Uncommon in wooded area

Euptoieta hegesia watsoni Mexican fritillary Seen once in open area

Hamadryas amphichloe Pale cracker Fairly common in wooded area

Junonia evarete Mangrove buckeye Common closest to salinas

Junonia genoveva Tropical buckeye Very common everywhere

Pieridae

Anaea borinquenalis Leafwing Seen once in wooded area

Aphrissa sp. Sulphurs

Appias drusilla monomorpha Amarilla white

Ascia monuste eubotea Great southern white Very common in open areas

Eurema elathea False-barred sulphur Common in open areas

Eurema lisa euterpe Little yellow

Kricoginia sp. Suphurs

Phoebis agarithe/sennae Large orange sulphur/ Uncommon in open areas

cloudless sulphur

Phoebis philea Orange-barred sulphur

a Sources: Haselmayer (2000), see endnotes; Asencio Lopez and Rentas Soto (2006), see endnotes. Lists compiled during 

visits to Cabo Rojo Refuge.

b Occurrence: Uncommon, fairly common, common, very common; abundant; sightings are a function of site and season. 

•	 The past cutting of Puerto Rico’s forests for

agriculture and selective harvest of commercial 

timber, i.e., habitat destruction, which occurred on 

99 percent of the island (Wadsworth 1950) 

•	 The devastation of dry forest in Puerto Rico,

the Neotropics, and worldwide, so much so that 

some consider it as the most endangered tropical 

ecosystem (Janzen 1988a)

•	 Habitat destruction resulted in 22 percent of Puerto

Rico’s native bird species being considered as 

either endangered or threatened (Raffaele 1983)

•	 The considerable number of threatened and

endangered species that use the current refuges 

in southwestern Puerto Rico; including resident, 

migratory, and endemic birds; and those with 

special habitat requirements such as mangroves 

and salt flats (U.S. Fish and Wildlife Service 2002)

•	 The relatively limited size of the refuges in

comparison to the island, and the need to secure 

sufficient habitat to assure recovery of listed 

species in recovery plans (fig. 1)
39

Sea birds. Sterna sandvicensis (sandwich terns), a species widely 

distributed throughout the world including southwestern Puerto Rico, 

breeds in dense colonies along the seacoast and on islands. 

(Photo by Jorge Saliva) 

•	 The actual and projected sharp increase in human

population and economic activity in the southwest, 

notably the municipality of Cabo Rojo (NúñezGarcía and Hunter 2000) (fig. 2) 

•	 The devastating impacts of recent hurricanes

which may portend a gloomy future given current 

concerns over global warming and more active 

and destructive hurricane periods (Van Bloom and 

others 2005) (fig. 8) 

•	 The threat of recurrent droughts and associated

wildfires that could destroy portions of the existing 

habitat and their wildlife, or negatively influence 

the breeding of some bird species 

•	 The increase of Erythrocebus patas and Macaca

mulatta monkeys and their potential impact on 

native wildlife along with the persistent, negative 

impact of the mongoose 

•	 The undetermined impact of exotic species on

native flora both in dry forest and wetland habitats 

•	 The relatively small size of the Lagoon tract, the

condition of farmlands immediately surrounding it, 

and the need for good land management to secure 

quality water for wildlife protection 

In addition, 14 of the 21 species considered as 

Caribbean sea bird fauna (Schreiber and Lee 2000) 

have been recorded on the refuges and protected areas 

of southwestern Puerto Rico. The species include: 

Anous stolidus (brown noddy), Fregata magnificans

(magnificent frigatebird), Larus atricilla (laughing 

gull), Phaethon lepturus (white-tailed tropicbird), and 

Pelecanus occidentalis (brown pelican), two boobies, 

Sula leucogaster (brown booby) and Sula sula (redfooted booby), and seven terns, Sterna anaethetus

(bridled tern), Sterna antillarum (least tern), Sterna 

dougallii (roseate tern), Sterna fuscata (sooty tern), 

Sterna hirundo (common tern), Sterna maxima

(royal tern), and Sterna sandvicensis (sandwich 

tern) (table 6). Most of these species exist in low 

densities and normally feed at considerable distances 

from breeding sites. Typically they produce a single, 

slow-growing offspring and populations are slow to 

recover after disturbances (Schreiber and Lee 2000). 

Primary threats to these sea birds are loss of nesting 

habitat due to development, human disturbance of 

colonies, introduced predators, and limited biological 

information for management purposes (Halewyn 

and Norton 1984, Schreiber 2000). Other problems 

include egg predation, pollution of oceanic waters 

due to various sources, excessive fishing, and 

recurrent hurricanes that can negatively impact 

small populations. 

Endangered Species

The Endangered Species Act of 1973, as amended 

through 1988, directs Federal Agencies to promote the 

recovery of all listed species (U.S. Fish and Wildlife 

Service 1988). Species’ names are published in the 

Federal Register and recovery plans are developed 

to enhance their survival. Recovery plans include 

a description of the current status of endangered 

species, their ecology (life history, distribution, 

and population trends), habitat description and 

requirements, factors that influence their survival, 

(e.g., fires, hurricanes), and reasons for listing the 

species. Also provided are recovery goals, recovery 

criteria, and the required actions to assure 

species recovery. 

Recovery criteria deal with sustainability of 

the species. Among the concerns are adequate 

reproduction, genetic robustness to avoid inbreeding 

and to allow for adaptation, sufficient habitat, and 

control of threats to survival. Between 1970 and 1996, 

the F&WS developed recovery plans for 123 species 

that are found on either the Cabo Rojo orand Laguna 

Cartagena NWR, as well as for one sea turtle and one 
40

plant species still not recorded but known from nearby 

sites (table 8).

The endangered plant species include two endemic 

grasses in the genus Aristida (McKenzie and others

1989). Aristida chaseae is found on exposed rocky 

outcrops at 150 to 200 m in elevation near Cerro 

Mariquita on the Tinaja tract. Aristida portoricensis

grows in exposed rock crevices in association with 

the former. Both have also been recorded at the 

western end of Sierra Bermeja. Three other listed 

plant species are endemic to the Sierra Bermeja and 

grow on the Tinaja tract. Eugenia woodburyana is 

a small, evergreen tree that grows on the slopes of 

Cerro Mariquita. Lyonia truncata var. proctorii, an 

evergreen shrub, grows on steep slopes in the open. 

Veronia proctorii, a small, erect shrub, grows on upper 

slopes near Cerro Mariquita. Ottoschulzia rhodoxylon, 

a small evergreen tree found in western Puerto Rico 

and Hispaniola (Little and others 1974), grows on 

the Tinaja tract. Stahlia monosperma, is a mediumsized tree that survives as scattered populations in dry 

Table 8—U.S. Fish and Wildlife Service species recovery plans applicable to species at the Cabo Rojo and Laguna 

Cartagena National Wildlife Refuges a

Dates

Group and species Common name Federal Register

Federal 

Register

Recovery 

plan

- - - - mo/day/yr - - - -

Flora

Aristida chaseae A.S. Hitchc. NA b (Poaceae) 58 FR 25755-25758 04/27/1993 07/31/1995

A. portoricensis Pilger c NA (Poaceae) 55 FR 32257 08/08/1990 05/16/1994

Catesbaea melanocarpa

Krug & Urban d Tropical lilythorn 64 FR 13116-13120 03/17/1999 07/15/2005

Eugenia woodburyana Alain e NA (Myrtaceae) 59 FR 46715 09/09/1994 10/06/1998

Lyonia truncata Urban var. proctorii f NA (Ericaceae) 58 FR 25755-25758 04/27/1993 05/16/1994

Ottoschulzia rhodoxylon 

(Urban) Urban g NA (Icacinaceae) 55 FR 13488-13491 04/10/1990 09/20/1994

Stahlia monosperma (Tul.) Urban h Cobana negra 55 FR 12790-12792 04/05/1990 11/01/1996

Vernonia proctorii L.E. Urbatsch i

 NA (Asteraceae) 58 FR 25755-25758 04/27/1993 05/16/1994

Fauna

Agelaius xanthomus j Yellow-shouldered blackbird 41 FR 51019-510122 11/19/1976 11/12/1996

Caprilumgas noctitherus k Puerto Rican whip-poor-will 38 FR 14678 06/04/1973 04/19/1984

Chelonias mydas l Green turtle 43 FR 32808 07/28/1978 10/29/1991

Dermochelys coriacea l Leatherback turtle 35 FR 8495 06/02/1970 04/06/1992

Eretmochelys imbricata l Hawksbill turtle 35 FR 8495 06/02/1970 12/15/1993

Pelecanus occidentalis occidentalis Brown pelican 35 FR 16047 10/13/1970 08/??/1985

a Source: Code of Federal Regulations 1970–1999.

b NA = not applicable (i.e., no common names or no existing recovery plans).

c Source: Silander (1994).

d Source: Rivera and Foote (2004). Reported on private property near the refuges.

e Sources: Breckon and Kolterman (1994); U.S. Fish and Wildlife Service (1993b, 1998). 

f Source: Breckon and Kolterman (1994).

g Sources: Breckon and Kolterman (1994); Silander (1994), see endnotes; U.S. Fish and Wildlife Service (1990). Recorded for Sierra 

Bermeja and possibly present on the Tinaja tract. 

h Source: Saliva (1996).

i Source: Breckon and Kolterman (1994).

j Sources: U.S. Fish and Wildlife Service (1976), see endnotes; Rivera (1996). 

k Source: Diaz Diaz (1984), see endnotes.

l Of the sea turtles, Chelonias mydas has not yet been recorded on the Salinas tract.
41

Rare and endangered. A recently discovered rare and 

endangered plant without a common name, Vernonia proctorii

L.E. Urbatsch, grows on the slopes of the Tinaja tract near 

Cerro Mariquita. (U.S. Fish and Wildlife Service photo)

coastal areas near mangroves (Little and Wadsworth 

1964). It has also been planted on both refuges. The 

remaining plant species, Catesbaena melanocarpa, 

has not yet been found on the refuges but is reported 

on private property nearby. It is also reported on St. 

Croix and in the Lesser Antilles. Six of the refuge 

plants, (i.e., all except Lyonia truncata var. proctorii 

and Ottoschulzia rhodoxylon,), are also listed 

as priority species that purportedly could be 

extirpated within 5 to 10 years (Missouri Botanical 

Garden 1992). 

Endangered animal species (table 8) include 

Agelaius xanthomus, formerly reported on the 

Headquarters tract (Wiley 1974, see endnotes) 

and still occasionally seen; and Falco peregrinus

(peregrine falcon), a recent winter migrant that has 

been delisted.The Puerto Rican whip-poor-will, 

Caprimulgus noctitherus, was recently reported on the 

Tinaja tract and sea turtles, Dermochelys coriacea and 

Eretmochelys imbricata, nest on the beaches at 

Cabo Rojo. 

Management Activities 

Management of the refuges involves routine 

surveillance as well as activities aimed at the 

protection, maintenance, and improvement of the 

habitat for wildlife, and education of the public. The 

major activities have included: fire prevention and 

control on both refuges, restoration of native habitat 

on both refuges, management of the Cartagena 

Lagoon, development of structures and improvements, 

and collaborative activities (table 9). 

Fire and Habitat Restoration 

Fires at both refuges like those during February 

of 2002 have caused considerable damage in the 

past. Fire continues to be used to clear pastures by 

farmers surrounding the refuges, particularly during 

the January through April dry season. Occasionally, 

these fires escape onto refuge lands. Fire prevention 

involves public education, training for fire fighting, 

and the construction and maintenance of fire breaks, 

especially during the dry season. In 2006, all fire 

breaks were widened. 

During the 1990s, a grassland management plan 

and a reforestation plan were developed for both 

refuges (Schaffner 1995a, 1995b, see endnotes). The 

long-term objectives of the grassland plan were to 

alter the composition favoring native grasses and 

change the overstory to benefit the bird species. 

Mowing grass for forage, prescribed burning, and 

the provision of firebreaks were incorporated into the 

plan. The reforestation plan was aimed at increasing 

the variety of native tree species. The planting list 

contained about 50 tree and shrub species, among 

them Roystonea borinquena, which is used for nesting 

by at least 15 resident bird species (Perez Rivera 

1977), 12 of which are recorded on the refuges. 

Both management plans were implemented to some 

extent although the vegetation conversion goals and 

proposed annual planting goals were not met. 

From 1980 to 2001, more than 9,000 trees 

representing 78 species were planted on the refuges 

for habitat restoration (Weaver and Schwagerl 2004) 

(table 10). By 2006, the number had increased to 
42

Table 9—Chronology of activities that influenced the Cabo Rojo National Wildlife Refuge (Headquarters and Las 

Salinas National Wildlife tracts) a

Date Events and activities 

700 BC Indians harvest salt; Indians throughout Puerto Rico engage in the commerce of salt from Cabo Rojo 

1500s + Las Salinas and Cerro Mariquita in Sierra Bermeja become reference points for travelers around southwestern

Puerto Rico

1508 Juan Ponce de Leon is the first Spanish explorer to set foot on the Salinas de Cabo Rojo; in 1511, he begins 

procedures for salt extraction

1528 Spain’s first confrontation with another European power (France) takes place in Bahía Sucia at Las Salinas

1585 British construct a bunker at Las Salinas as part of a military strategy to control Puerto Rico 

Early 1600s Las Salinas become communal property when the Spanish crown announces the collapse of the salt extraction 

1799 Last attack of the British in Puerto Rico takes place at Puerto Real, north of Punta Guanaquilla, in Cabo Rojo

municipality

1815 Spain proclaims La Real Cédula (Royal Document) allowing greater freedom in economic development and trade 

including the importation of equipment to facilitate salt extraction at Las Salinas

Mid-1800s Ramón Baldorioty de Castro works as administrator of Las Salinas

1883 Town of Lajas founded with 6,238 persons b

Early 1900 Forests in Lajas Valley are cut to increase land for sugar cane production b

1972 Department of Natural Resources is created with jurisdiction over the island’s natural resources b

1973 Passage of the Endangered Species Act b

1974 U.S. Fish and Wildlife Service acquires the Headquarters tract from the Foreign Broadcast Information Service; 

first avifauna survey on the Headquarters tract records 24 species

1976 Water Law (Ley de Aguas) is approved and Department of Natural Resources has authority to regulate the 

development and use of the island’s water resources b

1978 Full time manager arrives at the Headquarters site

1980 Reforestation begins by planting Bucida buceras at Headquarters tract

1986 McKenzie prepares the first critical plant list for the Headquarters site

1997 Major restoration begins with planting trials of numerous tree species

1999 Las Salinas is the second tract acquired as part of the Cabo Rojo Refuge 

2003 Visitor information and interpretative center is completed on Salinas tract 

2004 Observation tower of salt flats is completed; visitor’s house at Headquarters tract is refurbished 

2008 New Headquarters building is constructed

a Sources: Toro McCown and Chabert (1986); McKenzie (1986), see endnotes; Ramírez Padilla (1985). 

b These five events are also important with regard to the Cartagena Lagoon Refuge but are not repeated in table 11.
43

Table 10—Tree species planted at the Cabo Rojo and Laguna Cartagena National Wildlife 

Refuges, 1980–2003

Trees planted by Refuge a Assessment b

Species CR LC Total Survival Growth

- - - - - - - - no. - - - - - - - -

Acrocomia media O.F. Cook 10 5 15 P MR

Amyris elemifera L. c 14 0 14 MR MR

Andira inermis (W. Wright) DC. 15 146 161 G G

Annona glabra L. 2 30 32 G F

Annona muricata L. 2 2 4 MR MR

Annona squamulosa L. 35 0 35 G F

Avicennia germinans (L.) L. 40 0 40 P P

Bourreria succulenta Jacq. 57 31 88 G G

Bucida buceras L. 1,429 1,204 2,633 G G

Bunchosia glandulosa (Cav.) L.C. Rich. 32 0 32 MR MR

Bursera simaruba (L.) Sarg. 710 488 1,198 G G

Calophyllum calaba Jacq. 4 0 4 P MR

Capparis cyanophallophora L. c 6 0 6 MR MR

Capparis hastata Jacq. 4 0 4 MR MR

Capparis indica (L.) Fawe. & Rendle 20 2 22 F P

Cedrela odorata L. 32 6 38 F F

Ceiba pentandra (L.) Gaertn. 20 164 184 G G

Chamaesyce articulata (Aubl.) Britton 1 0 1 MR MR

Citharexylum fruticosum L. 72 107 179 G F

Clusia rosea Urban 2 1 3 MR MR

Cnidoscolus aconitifolius (Mill.) I.M. Johnst. 6 0 6 MR MR

Coccoloba diversifolia Jacq. c 12 0 12 MR MR

Coccoloba uvifera (L.) L. 190 5 195 P F

Cocos nucifera L. 13 69 82 P MR

Colubrina arborescens (Mill.) Sarg. 57 18 75 G G

Comocladia dodonaea (L.) Urban 2 0 2 MR MR

Conocarpus erectus L. 190 0 190 F F

Cordia alliodora (Ruiz & Pav.) Oken 12 0 12 MR MR

Cordia collococca L. 9 67 76 G F

Cordia sulcata DC. 62 92 154 G G

Cresentia cujete L. 25 39 64 G F

Cresentia portoricensis Britton 5 15 20 F F

Crossopetallum rhacoma Crantz c 12 0 12 MR MR

Erythoxylum areolatum L. 91 7 98 G G

Erythroxylum rotundifolium Lunan c 10 0 10 MR MR

Eugenia ligustrina (Sw.) Willd. 3 0 3 MR MR

Eugenia woodburyana Alain 2 0 2 P P

Ficus benjamina L. 4 0 4 MR MR

Ficus citrifolia Mill. 57 107 164 P P

Guaiacum officinale L. 312 159 471 G P

Guaiacum sanctum L. c 14 0 14 MR MR

Guazuma ulmifolia Lam. 25 63 88 G G

Gymnanthes lucida Sw. c 14 0 14 MR MR

Helicteres jamaicensis Jacq. 8 0 8 G F

Hymenea courbaril L. 10 8 18 P P

Inga fagifolia (L.) Willd. 2 0 2 MR MR

Jaquina arborea Vahl 41 20 61 P P

Mangifera indica L. 2 0 2 MR MR

continued
44

Table 10—Tree species planted at the Cabo Rojo and Laguna Cartagena National Wildlife 

Refuges, 1980–2003 (continued)

Trees planted by Refuge a Assessment b

Species CR LC Total Survival Growth

- - - - - - - - no. - - - - - - - -

Manilkara bidentata (A. DC.) Chev. 0 13 13 MR MR

Masticodendron foetidissimum (Jacq. Crong. 6 39 45 F F

Melococcus bijugatus Jacq. 11 5 16 MR MR

Metopium toxiferum (L.) Krug & Urban 3 0 3 MR MR

Petitia domingensis Jacq. 5 0 5 MR MR

Pimenta racemosa (Mill.) J.W. Moore 2 0 2 MR MR

Piscidia carthagenesis Jacq. 4 0 4 MR MR

Pisonia albida (Heimer) Britton 24 1 25 MR MR

Pisonia subcordata Sw. 5 36 41 MR MR

Samanea saman (Willd.) Merrill 21 23 44 G G

Polygala cowellii (Britton) Blake 78 14 92 P F

Pterocarpus officinalis Jacq. 0 13 13 MR MR

Randia aculeata L. 3 1 4 MR MR

Reynosia uncinata Urban c 4 0 4 MR MR

Roystonea borinquena O.F. Cook 2 9 11 P P

Sabal casuarium (O.F. Cook) Beccari 4 0 4 MR MR

Sapindus saponaria L. 0 21 21 F F

Senna polyphylla (Jacq.) Irwin & Barneby 24 0 24 G F

Spondias mombin L. 28 48 76 G F

Stahlia monosperma (Tul.) Urban 330 368 698 G F

Sterculia apetala (Jacq.) Karst. 11 0 11 MR MR

Swietenia mahogani Jacq. 716 0 716 G G

Tabebuia heterophylla (DC.) Britton 128 147 275 G F

Tamarindus indica L. 10 9 19 G F

Terminalia catappa L. 4 0 4 MR MR

Thespesia populea (L.) Soland 150 25 175 P MR

Trichilia hirta L. 47 37 84 G G

Trichilia tricantha Urban 4 0 4 P MR

Zanthoxylum flavum Vahl 73 36 109 P MR

Ziziphus reticulata (Vahl.) DC. 3 0 3 MR MR

 Total 3,700 5,397 9,097

a Refuges: CR = Cabo Rojo; LC = Laguna Cartagena.

b Tentative assessment—Survival and growth: G = good; F = fair; P = poor; MR = more research is needed.

c Tree species planted in research plot near airstrip by Sandra Molina.
45

Habitat restoration. Several tree species were tested in the continuing 

effort to reforest abandoned agricultural land surrounding the 

Cartagena Lagoon. (U.S. Fish & Wildlife Service photo) 

Research on survival and growth. A sapling of Andira inermis

(W. Wright) HBK (moca) is measured for growth at the Laguna 

Cartagena National Wildlife Refuge. The early survival and growth of 

most planted species were assessed recently. (Photo by Peter L. Weaver) 

about 17,500 trees, 88 percent of which were on 

the Headquarters and Lagoon tracts (Weaver and 

Schwagerl 2008) (fig. 10). At an average spacing of 

5 by 5 m, i.e., 400 trees/ha, the original plantings were 

designed to cover about 44 ha. 

Five species—Bucida buceras, Bursera simaruba, 

Guaiacum officinale, Stahlia monosperma, and 

Swietenia mahagoni—represented 63 percent of the 

plantings. Survival and growth rates for all of the 

species were assessed recently (table 10). Growth 

in diameter and height showed that Bucida buceras, 

Ceiba pentandra, and Guazuma ulmifolia performed 

best among native species, and Swietenia mahagoni

among the exotics. Survival was rated as good 

for 25 species, 12 of which also had good growth 

rates. Another six species had fair survival rates. 

The plantings showed that a concerted effort using 

available resources can be successful if fire protection 

is provided. The plantings also proved that several 

tree species perform satisfactorily and may be used in 

habitat restoration. 

Puerto Rico’s dry forest refuges are critical nesting 

and foraging habitat for native and migratory birds. 

In the past, dry forests were widespread, not only 

on the island but throughout the neotropics. Today, 

major efforts are being made in some countries like 

20

18

16

14

12

10

8

6

4

2

0

Cumulative number of trees (thousand)

3 5 7 9 11 13 15 17 19 21 23 25 27

Year—1980 to 2006

Headquarters

Salinas

Lagoon

Tinaja

Total

1

Figure 10—Tree plantings by refuge tract and year since 1980.
46

Costa Rica to restore wildlife habitat (Janzen 1988b). 

Research has shown that some species of woody 

plants regenerate satisfactorily under tree plantings. 

By comparison, recruitment of woody plants into 

pasture lands is slow because grasses compete for 

water and nutrients, or because recurrent fires inhibit 

woody growth (Guariguata and others 1995). Patterns 

of understory regeneration vary with overstory tree 

species, their densities, and their management. 

Complete recovery of dry forest from past 

disturbance takes considerable time and involves 

biomass accumulation, the development of a mature 

forest structure, and regrowth of a typical species 

composition. In southwestern Puerto Rico, the 

recovery of plant biomass, regardless of species, may 

take 50 years or more. Recovery of mature structure, 

i.e., a mixture of small and large diameters of native 

and exotic species, could take 200 years or more. 

Finally, recovery of biomass and structure along 

with a species composition similar to that at the 

time of the island’s discovery, i.e., typical speciessite relationships, could easily take one-half of a 

millennium or more (Weaver and Schwagerl 2005). 

At that time, shade-tolerant exotic trees are likely to 

persist. Moreover, unknowns such as climatic change 

could impose other ecosystem changes. A sustained 

reforestation effort using native tree species should 

accelerate natural succession through its effects 

on vegetative structure, microclimate, and soils 

(Parrotta 1993). 

Cartagena Lagoon 

The Cartagena Lagoon was originally an open, 

freshwater wetland with a short hydroperiod, and 

without a permanent inlet or outlet (Schaffner 1994, 

see endnotes). The bottom of the lagoon, consisting of 

dense alluvial clay, trapped local rainfall, sheet flow, 

and ephemeral streamflow to accumulate water during 

the wet season. Shallow, open water, occupying most 

of the lagoon, provided foraging habitat for resident 

and migrant waterfowl and the surrounding emergent 

vegetation was used for nesting by resident aquatic 

birds. The gradual decline in water levels associated 

with the dry season extended foraging opportunities 

for wading species. 

Agricultural development during the late 19th

century and throughout most of the 20th century 

negatively impacted the Cartagena Lagoon (table 11). 

In the late 1800s, farming intensified in and around 

the Lajas Valley when forest cover was gradually 

replaced by sugar cane plantations. Both infiltration 

of rainwater into the soil and runoff were influenced. 

About 1920, the farmers began pumping the lagoon to 

irrigate surrounding lands, reducing the lagoon’s dry 

season water levels. Early ornithologists expressed 

concern over the potential impacts of water removal 

on the lagoon’s avifauna (Danforth 1926). In the 

1950s, the completion of the Lajas Valley irrigation 

project (Koenig 1953) left Cartagena Lagoon as

the only body of fresh water in the valley. In 1955, 

a proposal for draining the lagoon was opposed by 

government officials in Puerto Rico and the United 

States. In the late 1950s, widespread recognition of 

wildlife values associated with the lagoon resulted in a 

recommendation that it be expropriated. At that time, 

landowners and renters had constructed dikes and 

drainage canals to further lower water levels. 

In 1961, the Margara Canal was completed to 

conduct unused irrigation water from the north and 

to maintain lagoon water levels. Drainage water from 

sugar cane fields, however, aggravated eutrophication; 

and the relatively static water levels promoted the 

encroachment of water lettuce, water hyacinths, and 

cattails into open water. Moreover, the stable water 

levels and fertilized runoff from cane fields caused 

the gradual formation of a thick, floating peat mat 

(Ramírez Toro and Minnigh 1997, see endnotes). In 

the early 1980s, avifauna abundance and diversity 

were notably less than in earlier years. In 1982, the 

lagoon dried out completely. 

In 1989, the lagoon was finally established as a 

wildlife refuge; however, maintenance of the Margara 

Canal and drainage canals was discontinued by 

the local government, causing aquatic vegetation 

to proliferate. In the early 1990s, the water control 

structure was repaired and the floating peat mat was 

discovered. The peat mat, along with the lack of 

canal maintenance, contributed to greater flooding 

in Magüayo. The mat, by virtue of its volume, 

increased the margin of the lagoon during rainfall 
47

Table 11—Chronology of activities that influenced the Laguna Cartagena National Wildlife Refuge (Lagoon and 

Tinaja tracts) a

Date Events and activities

1493 Columbus lands in southwestern Puerto Rico; lagoon functions naturally, receiving water by overland flow and

from small, intermittent streams (arroyos) during heavy rains; avifuna abounds in diversity and numbers

Late 1800s – Farming begins in Lajas Valley; trees are cut for housing, agriculture, and fuel; water regimen changes,

early 1900s possibly at first by accentuating wet season runoff and gradually diminishing water levels during the dry 

season; the Lajas Valley becomes privately owned farms

1920s Local farmers begin pumping Cartagena Lagoon to irrigate sugar cane; dry season water levels diminish; 

Danforth investigates bird species at Cartagena Lagoon and initiates a Christmas bird count; Gleason and

Cook describe the vegetation in the Lajas Valley and around the lagoons at Cartagena and Guánica

1945 Quebrada Los Llanos diverts water toward Cartagena Lagoon

1950s Lajas Valley irrigation system is built (Lago Loco at Río Loco dam is constructed to supply water to the Lajas 

Valley; the Laguna Anegada and Laguna Guánica are drained to reclaim land for cultivation); a drainage canal

runs through the Cartagena Lagoon to the Bahía de Boquerón; Cartagena remains as only fresh water lagoon

in the Lajas Valley, resulting in additional hunting pressures on its wildlife

Mid–1955 Proposed drainage of Cartagena Lagoon is opposed by the governor of Puerto Rico and the U.S. Secretary of

the Department of the Interior

1956 McCandless revives the Audubon Christmas count between Cartagena Lagoon and Los Morrillos de Cabo

Rojo; a new water control structure is built at Cartagena Lagoon and continuous pumping drastically reduces

lagoon area; water levels remain relatively constant except for major rainfall events

1958-59 Importance of Cartagena Lagoon’s wildlife, scientific, economic, sport (hunting), tourism, and recreation values

is recognized and expropriation is recommended; land owners and renters construct dikes and drainage canals

lowering the water levels in the lagoon

Late 1950s Federal aid project establishes aquatic vegetation (water lettuce, water hyacinth, and cattails); water extraction 

continues to reduce the lagoon size creating more land to irrigate for sugar cane production 

1960s Drainage water from sugar cane fields aggravates eutrophication (i.e. fertilizers, pesticides, and sediment); 

untreated sewage enters from town of Magüayo along the refuge’s eastern border; plant associations change; 

relatively static water levels promote the growth of water lettuce, water hyacinths, and cattails into open water

areas; bird species decline in the lagoon

1961 Magara Canal is completed to conduct unused irrigation water from the north to maintain lagoon water levels; 

vandalism of the water control structure prevents filling the lagoon to the proposed target level of 11 m

1979 Department of Natural Resources recognizes the Cartagena Lagoon as a critical wildlife habitat

Early 1980s Avifauna diversity and abundance is significantly less than in earlier years; Magüayo expands northward to the 

Magara Canal

1982+ Lagoon dries up and the pumping of irrigation water stops; growth and annual burning of cattails results in

accumulation of cattail biomass into a floating mat

1983 The Department of Natural Resources requests the U.S. Congress to acquire Cartagena Lagoon; the Puerto

Rico Land Administration receives Cartagena Lagoon and surrounding properties 

1989 Cartagena Lagoon is established as a wildlife refuge through lease agreement with the Land Administration; 

maintenance of Margara and drainage canals terminates, inhibiting the drainage of canal waters; aquatic 

vegetation develops in the canals; heavy rainfalls cause the overflow of canals, saturating adjacent soils and

increasing flood events in Magüayo

continued
48

Table 11—Chronology of activities that influenced the Laguna Cartagena National Wildlife Refuge (Lagoon and 

Tinaja tracts) a (continued)

Date Events and activities

1990 The Puerto Rico Planning Board designates the Cartagena Lagoon National Wildlife Refuge as a Natural

Reserve

Early 1990s Water control structure is repaired and the floating peat mat is discovered 

1993 Water quality studies are initiated by Interamerican University, San German; observations show that nutrients

leaving the lagoon are greater than the amounts entering; sugar cane harvest terminates around Cartagena

Lagoon

1994 Worst drought in several decades causes the lagoon to dry up

1996 The Tinaja tract is acquired as part of the Laguna Cartagena Refuge with fee title from the U.S. Department of

Agriculture Farm Service; Proctor prepares the first plant list for the Tinaja tract

1997+ Grazing and fire are eliminated from the Tinaja tract; reforestation begins on the Lagoon and Tinaja tracts

2005 The observation tower is built at Cartagena Lagoon

a Sources: Danforth (1926); Díaz-Soltero (1990); Gleason and Cook (1927); Proctor (1996), see endnotes; Ramírez Padilla (1985); Schaffner 

(1994), see endnotes; Toro McCown and Chabert (1986).

Cartagena Lagoon. The lagoon, with ample water, supports an 

uncommon burst of flowering Eichornea crassipes (Mart.) Solms.-Laub. 

(water hyacinths). (Photo by Peter L. Weaver)

events, and hampered water flow through the lagoon. 

The intent of the water control structure was to gain 

control over water levels and reestablish the lagoon’s 

historic hydroperiod. Fluctuations in water level 

would purportedly result in the characteristic species 

changes along the margin of the lagoon and benefit 

the wildlife. 

In 1990, a comprehensive management plan was 

prepared for the Cartagena Lagoon with information 

on its flora and fauna as well as suggested monitoring 

and research activities (Díaz-Soltero 1990). In 1994, 

the lagoon dried out again during the island’s worst 

drought in decades. 

In 1996, the Tinaja tract was added to the Laguna 

Cartagena NWR. For several years, the lower part 

of the tract had been managed for livestock grazing, 

which included recurrent burning. Fencing around the 

tract was repaired and grazing eliminated. In 2000, 

the lagoon entered the new millennium in a degraded 

condition, suffering from a century of problems, 

and replete with a legacy of efforts to successfully 

mitigate them. However, surveys conducted in 1998 

and 2003 showed that woody vegetation had increased 

in the absence of fire and grazing (Weaver and Chinea 

2003; Weaver and Schwagerl 2004, 2005). Finally, an 

unpolluted source of fresh water was again available 

to the lagoon. 

Structures and Improvements

The major human improvements at the Cabo 

Rojo and Laguna Cartagena NWRs consist of the 

development of various types of buildings, roads, 

trails, towers, water control structures, security 

measures, and commercial salt ponds (table 12), 
49

Table 12—Structures and improvements at the Cabo Rojo and Laguna 

Cartagena National Wildlife Refuges a

Cabo Rojo Laguna Cartagena

Structures or improvements Headquarters Salinas Lagoon Tinaja

Buidlings and related facilities

Office and visitors center (m2

) 1935

Shop building (m2) 595

Guard house (m2

) 9

Microwave building and pole (m2) 21

Oil building (m2) 6

Refuge workshop (m2

) 186

Tree shade houses–nursery (m2) 160

Museum complex (m2

) 209

Museum parking (m2) 17

Other b NA

Improvements—visitors

Observation towers (no.) 1 1

Information kiosk (no.) 1

Signs (information) (no.) 5 1

Interpretative trail (km) 2.3

Cactus garden (ha) 0.1

Farmhouse (no.) 1

Silo (no.) 1

Ruins (no.) 5

Structures—water

Canals or drainages (km) 5.6 3.5

Water level controls (no.) 2 3

Wells (no.) c 4 5

Watering troughs (no.) 2 1

Impoundments (no.) 4 1

Access roads 

Paved roads—entrance (km) 1.2

Dirt roads (km) d 24.6 3.2 3.1 2.4

Security

Boundary fencing (km) 17.7 11.5 7.1 4.5

Gates (no.) 2 2 2 1

Firebreaks (km) 6.5 1.9 14.8

Commercial activities

Salt ponds (no.) 2

a Cabo Rojo (Headquarters and Salinas tracts); Laguna Cartagena (Lagoon and Tinaja tracts).

b Septic tank, water lines, sewer lines, transformers, electrical lines, and telephone lines at 

Cabo Rojo headquarters site.

c At Cabo Rojo, one for irrigation and three out of service; at Laguna Cartagena, one for 

observation, one for irrigation, and three out of service.

d Only 1 km of dirt road is being maintained at Tinaja.
50

Nursery production. The small headquarters nursery regularly 

produces seedlings for planting. Between 1980 and 2001, more than 

9,000 trees representing 78 species were planted on the Cabo Rojo 

and Laguna Cartagena National Wildlife Refuges. (Photo by 

Peter L. Weaver)

along with the planting of trees. The Headquarters 

tract includes an office building with a visitor’s 

reception area and interpretative displays, a workshop, 

a tree nursery, four wells, an old farm house and silo 

used today for volunteers, and cattle watering troughs 

scattered throughout the property. Improvements 

include a 0.1-ha cactus garden, 1.2 km of paved roads, 

25 km of dirt roads, and 6.5 km of firebreaks. The 

property also has a security gate and is surrounded by 

17.7 km of fencing. 

The major structures at the Lagoon tract are the 

lagoon water control structures, 3.5 km of canals, 

five wells, scattered cattle watering troughs, and a 

7-m observation tower overlooking the lagoon. In 

addition, there are 3.1 km of dirt roads and 14.8 km 

of firebreaks. About 7.1 km of fencing surround the 

property. The only structure on the Tinaja tract is a 

watering trough. In addition, there are 2.4 km of dirt 

roads and 4.5 km of fencing surrounding the property. 

Descriptive information regarding all wells in the 

Lajas Valley, including those found on the refuges, 

was compiled and published by the U.S. Geological 

Survey (Graves 1991). 

Collaborative Activities 

The Coastal Zone Management Program 

(PMZCPR in Spanish) was established in Puerto 

Rico by virtue of the 1978 Federal Coastal Zone 

Management Law (Departamento de Recursos 

Naturales y Ambientales 1997, 2000b). Special 

planning areas were designated by the Puerto Rican 

Government to protect the island’s natural, historic, 

cultural, and archaeological resources. These special 

areas include Boquerón, Parguera, and Guánica along 

Puerto Rico’s southwest coast. The Boquerón sector 

of the Southwest Special Planning Area (SBAPES 

in Spanish) covers the area from Punta Pitahaya 

to PR Route 101, north of Punta Guaniquilla, and 

encompasses the Cabo Rojo NWR and borders the 

Laguna Cartagena NWR. In carrying out its mission, 

the F&WS cooperates with the Puerto Rico Planning 

Board, the Puerto Rico Department of Natural and 

Environmental Resources, and the Puerto Rico 

Department of Public Works. 

In 1993, an interagency committee was formed 

among Federal and Puerto Rican Agencies with 

responsibility or authority for the Cartagena 

Lagoon. Since the F&WS was responsible for land 

management and leasing the lagoon from the Puerto 

Rico Land Authority, both were represented. Other 

entities included were:

•	 Federal—the Army Corps of Engineers, the U.S.

Department of Agriculture, the Natural Resources 

Conservation Service, and the U.S. Environmental 

Protection Agency

•	 Puerto Rico—the Department of Natural

and Environmental Resources and the 

Energy Authority

The Salinas tract has a small visitor information 

center, salt ponds, an 8-m high observation tower 

overlooking the salt operations, the foundation of an 

old windmill, a short birdwatchers trail, culverts, and 

an unimproved road on top of a dike leading to the 

fishermen’s facility. This facility is a 2.5 -ha enclosure 

with a building that contains equipment for the 

storage of fish and fish products by the municipality’s 

fishermen (Departamento de Agricultura 1995, 

see endnotes). Since 1981, the Commonwealth 

Department of Agriculture has rented the property 

and structure to the Municipality of Cabo Rojo for a 

nominal sum. In addition, there are at least 3.2 km of 

dirt roads and 1.9 km of firebreaks. A portion of the 

property along PR Route 301 is fenced. 
51

Visitors to refuges. The refuges receive nearly 150,000 visitors each 

year, about 90 percent for a day on the beach. The remainder, like 

these Boy Scouts, visit for a hike and birdwatching at either the 

Headquarters tract or Laguna Cartagena. (Photo by Ken Foote)

•	 Center for Education, Conservation and

Environmental Interpretation (Centro para 

Educación y Interpretación Ambiental, or CECIA 

in Spanish) of Interamerican University at 

San German 

A work plan was devised with short- and long-term 

goals including habitat restoration, the installation of a 

water control structure, and the mitigation of floods in 

the town of Magüayo. The Puerto Rico Conservation 

Foundation (Fundación Puertorriqueña de 

Conservación in Spanish), the Department of Natural 

Resources and the Environment, and the Autoridad de 

Carreteras assisted in generating public support for the 

acquisition of the Salinas tract. 

In addition to the aforementioned groups, the 

F&WS cooperates with several other Commonwealth 

and Federal Government Agencies, and with 

numerous universities in Puerto Rico and elsewhere, 

including:

•	 U.S. Department of Agriculture Forest Service,

International Institute of Tropical Forestry

•	 U.S. Department of Agriculture Natural Resources

Conservation Service

•	 U.S. Geological Service, Biological Resources

Division

•	 Conservation Trust of Puerto Rico

•	 Puerto Rico Department of Agriculture

•	 Puerto Rican Rural Development Corporation

•	 Puerto Rican Ornithological Society

•	 Pontifical Catholic University of Puerto Rico

in Ponce

•	 Metropolitan University

•	 University of Puerto Rico

•	 McMaster University in Hamilton, Ontario,

Canada

•	 Plymouth State University in New Hampshire

•	 University of Missouri in Columbia

The F&WS also works closely with the local 

nongovernmental organization called the Cabo 

Rojo Residents for Environmental Health (Cabo 

Rojeños Pro Salud Ambiente in Spanish), which is 

a refuge friend’s group. Another important aspect 

of collaborative activities concerns the residents 

of the nearby community of Magüayo, since fires 

originating in that vicinity have burned refuge lands 

on several occasions. A recent survey showed that 

residents had little appreciation of the environmental 

values associated with the refuge and that the majority 

had not participated in environmental programs 

or restoration efforts (Ruiz and Huertas 2005, see 

endnotes). Linking benefits to the community with 

conservation efforts might stimulate local interest 

and participation. 

Public Use of Refuges

Visitors

During 2005, the Cabo Rojo NWR received 

135,400 visitors, mainly at beach sites, and 11,200 

visitors at the Headquarters Visitors Center. Another 

8,000 persons in 90 groups visited the Cabo Rojo 

Interpretative Center on the Salinas tract. About 1,000 

people visited Cartagena Lagoon. (U.S. Fish and 

Wildlife Service 2005, see endnotes). Fishing, but not 

hunting, is allowed on the refuges. However, fishing 

is considered poor and no records are kept. Other 

activities at both refuges included: 

•	 1,050 visitors for wildlife observations, including

920 that used foot trails

•	 200 visitors for photography, including 45 that

used photo blinds
52

•	 2,370 participants in environmental education

programs, including 38 teachers in programs held 

onsite and offsite, and 2,264 students in programs 

held onsite and offsite

•	 4,890 participants in interpretative programs,

including 2,330 held onsite and 1,560 held offsite

Residents and tourists are encouraged to visit the 

refuges through available brochures and books 

(Nellis 1999, Raffaele 1983, U.S. Fish and Wildlife 

Service 2004). 

Special Use Permits

A total of 174 special use permits have been 

granted for activities at the Cabo Rojo and Laguna 

Cartagena NWRs since 1979–97 at the Cabo Rojo 

Headquarters tract, 11 at the Salinas tract, 64 at the 

Lagoon tract, and 2 at the Tinaja tract (table 13). Two 

permits were issued during the 1970s, 15 in the 1980s, 

116 in the 1990s, and 41 since 2000. Of the total, 2 

involved structures, i.e., removal of antennae cables 

and anchors, and building culverts and water diversion 

devices; and 70 were administrative in nature dealing 

with various topics, i.e., release of floodwaters on 

refuge property, tick eradication, testing broadcast 

signals from South American stations, cutting forage 

for animals (silage), salt extraction operations, grazing 

to reduce grass cover or sugar cane, and maintenance 

of vegetation along the roadsides and in irrigation 

canals or drainages. The remaining 100 use permits 

dealt with 42 studies on various topics, some renewed 

at later dates, i.e., climate, botany and botanical 

collections, vegetation monitoring, native and exotic 

vertebrate wildlife species, invertebrates, wildlife 

monitoring, photography, and water sampling. 

The F&WS has sponsored studies, research 

projects, and educational programs with universities 

and government agencies through grants and 

cooperative agreements for several years. Most of 

the cooperating students and professors are from 

universities in Puerto Rico, Canada, and the United 

States. The topics of many studies are summarized in 

the list of special use permits (table 13). Study plans 

or proposals and final reports or publications are 

required for F&WS records. 

Among the formal activities of the F&WS are 

those related to planning for bird conservation 

(Núñez-García and Hunter 2000) and biological 

reviews (U.S. Fish and Wildlife Service 2002) 

where various research and monitoring needs are 

indicated. Christmas counts have also been carried 

out for several years. Among the less formal research 

activities are recurrent bird surveys, which include 

tallies of bird species and estimates of their numbers. 

Other Major Protected Areas 

in the Southwest

Six other major protected areas are situated in 

southwestern Puerto Rico, in addition to the F&WS 

refuges of Cabo Rojo and Laguna Cartagena. They 

are (Department of Natural Resources 1979, VentosasFebles and others 2005) (fig. 1):

•	 Boquerón Commonwealth Forest

•	 Boquerón Wildlife Refuge

•	 Guánica Commonwealth Forest

•	 Los Morrillos de Cabo Rojo

•	 Parguera Natural Reserve

•	 Punta Guaniquilla Natural Reserve

Boquerón Commonwealth Forest 

The Boquerón Commonwealth Forest is partitioned 

into three major areas: (1) an area along the west 

coast near Punta Guaniquilla extending southeast to 

Laguna Rincón, (2) a patch along the eastern edge 

of Bahía Sucia, and (3) an extensive area ranging 

from Punta Pitahaya east to Parguera (Department of 

Natural Resources 1976). The forest occupies 1876 

ha between sea level and 5 m in elevation (table 1). 

Most of the area is classified as tidal swamp and 

tidal flats (Carter 1965). The major types of wildlife 

habitat in the Boquerón Forest are islets, mangroves 

and mudflats, salinas, and coastal lowlands. The main 

cover types are basin and fringe mangroves, beach 

thicket, salt flat vegetation, and scrub woodland 

(Cintrón and others 1978). 

Mangrove roots provide habitat for invertebrates 

and algae, and offer protection for numerous species 

of juvenile fish (Austin 1971). Also, regardless 

of salinity or temperature, several species of fish, 

including Centropomus spp., Mugil spp., Tylosurus 

crocodilus, Gerres ciriereus, and Diapterus plumieri

frequent mangrove systems. During the 1970s, the 

social and foraging behavior of 18 species of warblers 

(Parulidae) was studied on 2 offshore islands—
53

Table 13—Special use permits for the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Decades a

Permit topics 1970s 1980s 1990s 2000+

number of permits

Cabo Rojo—Headquarters tract

Flood waters: release on refuge lands 1

Solar radiation: measurement 1 2

Climate: study before installation of areostat radar system 1

Botanical specimens: collect and examine 1

Broadcast antenna: remove cables and anchors 1

Ticks: administer U.S. Department of Agriculture eradiction program 1

Cattle ticks: study basic ecology and longevity under field conditions 1

Broadcast signals: investigate levels from South American stations 3

Herpestes auropunctatus (mongoose): field test oral vaccine for rabies; determine age and sex 1

Herpestes auropunctatus (mongoose): live trap and implant microchips for tracking 1 2

Hay: cut and remove for livestock feed and silage 2 22 3

Panicum maximum (Guinea grass): determine the impact of cattle grazing 1

Vegetation: conduct surveys and collect specimens except endangered species 1

Feral primates: collect data on ecology, distribution, and behavior 3

Grass: cut by hand and remove as feed for domestic animals 3

Dendroica discolor (prairie warbler) and Parula americana (northern parula): 

investigate the use of several habitats 4

Bird nesting: study reproductive success of nesting birds, notably exotic finches 2

Vegetation: collect samples for analysis of habitat and nest site selection by birds 2

Dry forest: study floristic composition of forest fragments 1

Bird species: survey distribution, abundance, and habitats using point counts 2

Molothrus bonariensis (shiny cowbirds): use radio transmitter to determine longevity 2

Shaerodactylus nicholsi (Nichol’s dwarf gecko): study 3

Bats: use mist nets to determine species composition and abundance 1

Polygala cowellii (violet-tree): study pollination and population ecology 2

Myiarchus antillarum (Puerto Rican flycatcher): study breeding biology and foraging ecology 1

Myiarchus antillarum (Puerto Rican flycatcher) and Icterus dominicensis (Greater Antillian 

oriole): nesting and breeding season observations 1

Monitor: Neotropical migrant and resident landbirds 1

Prosopis juliflora (mesquite) and Bucida buceras (oxhorn bucida) trees: 

cut mesquite to release planted oxhorn bucida 2

Seiurus motacilla (Louisiana water thrush) and S. noveboracensis (northern water 

thrush): determine if species are limited by available habitat in Puerto Rico 4

Bursera simaruba (gumbo-limbo) and Hymanea courbaril (West-Indian-locust) trees: 

collect leaf and seed samples for study of gene flow among forest fragments 1 2

Crotophaga ani (smooth-billed ani): tracking, banding, blood samples 3 1

Aristida chaseae (endemic grass): life history and competition with other grasses 2 1

Native tree species: develop protocol for seed germination for use at refuges 2 1

Falco sparverius (American kestrel): capture for genetic analyses 1

Refuge: site sampling to determine bacterial mats, diatoms, and cilates 1

Aristida chaseae (endemic grass): collect samples 1

Seeds: collect in traps under crowns of native and exotic trees and identify 2

Subtotal 2 14 68 13

continued
54

Table 13—Special use permits for the Cabo Rojo and Laguna Cartagena National Wildlife Refuges (continued)

Decades a

Permit topics 1970s 1980s 1990s 2000+

number of permits

Cabo Rojo—Salinas tract

Structures: install culverts and water diversion devices 1

Artemia franciscana (brine shrimp): ultraviolet B radiation as factor in reproduction 1

Birds: blood samples and banding 1

Photography: series of photos of habitat and species 1

Migratory shorebirds: identify factors that affect their distribution and density 1

Salt extraction: operations 3

Vegetation: maintain roadside plants 3

Subtotal 1 10

Laguna Cartagena—Lagoon tract

Water: sample and determine basidiomycete and ascomycete fungi 1

Panicum maximum (Guinea grass): use grazing to help control growth 7

Vegetation: survey and collect specimens except endangered species 2

Sugar cane: eliminate by cattle grazing 1

Grass: cut and remove for livestock 10

Canals: provide access to existing irrigation and drainage structures 2

Feral primates: collect data on ecology, distribution, and behavior 3

Dendroica discolor (prairie warbler) and Parula americana (northern parula): 

investigate the use of several habitats 2

Bird nesting: study reproductive success of nesting birds, notably exotic finches 2

Vegetation: collect samples for analysis of habitat and nest site selection by birds 2

Monitor: Neotropical migrant and resident landbirds 1

Bursera simaruba (gumbo-limbo) and Hymanea courbaril (West-Indian-locust) trees: 

collect leaf and seed samples for study of gene flow among forest fragments 1

Myiarchus antillarum (Puerto Rican flycatcher) and Icterus dominicensis (Greater Antillian 

oriole): nesting and breeding season observations 1

Crotophaga ani (smooth-billed ani): tracking, blood samples and banding 3 1

Hay: cut and remove for silage 8 1

Gastropods and bivalves: collection and identification 1 1

Water: collect samples for determination of ionic composition and salinity 1

Grazing: livestock forage 13

Subtotal 47 17

Laguna Cartagena—Tinaja tract

Vegetation: survey and collect specimens except endangered species 1 1

Total 2 15 116 41

a Permits first granted in 1979. 
55

Boquerón Commonwealth Forest. The Boquerón Forest has a long 

history of human activity. As early as 1875, the local forestry agency 

(Inspección de Montes) attempted to control the annual removal of 

bark and fuelwood from these mangrove woodlands. (Photo by 

Peter L. Weaver) 

Guayacán of 35 ha and Cueva of 20 ha—within the 

Boquerón Forest (Post 1978). Population densities 

were high and the birds usually forged alone or in 

stationary (passive) flocks. The stationary flocks 

were usually composed of birds that had gathered 

to consume insects foraging in different patches of 

vegetation. Aggressive behavior occurred frequently 

and was common among the same species or those 

with similar foraging behavior 

Despite their known importance as critical 

habitat for fish and bird species, pressures 

continue for the development of mangrove areas 

as recreation and tourism sites. The 1976 master 

plan for Commonwealth forests outlines issues and 

recommendations concerning wildlife protection 

and potential uses (Department of Natural Resources 

1976). The master plan also states that the Boquerón 

Forest never had a management plan; however, as 

long ago as 1875, the “Inspección de Montes” 

(a forestry agency under Spanish authority) attempted 

to manage the annual harvest of bark and fuelwood 

in the Boquerón mangrove (Dominguez Cristóbal 

2000, 2001). 

Boquerón Wildlife Refuge 

During the 1950s, both Ciénaga El Anegado (about 

780 ha) and Laguna Guánica (about 450 ha) were 

drained as part of the Lajas Valley irrigation project; 

earlier observations noted at least 24 bird species 

in the vicinity (Beatty 1931, Bonnett and Tirado 

1950). In addition, the project resulted in serious 

deterioration of the Cartagena Lagoon. In 1964, the 

Puerto Rico Department of Agriculture designated 

182 ha of the Boquerón Forest as the Boquerón 

Wildlife Refuge to protect the flora and fauna, 

including mangroves, sea-grass beds, and pelicans 

(Cruz Alonso 1997, Departamento de Recursos 

Naturales 1982). Laguna Rincón is in the western 

part of the refuge (Burger and others 1992) (fig. 1). 

The lagoon occupies 74 ha and connects with Bahía 

Boquerón via a canal that is 0.2 km long and 0.75 m 

deep (Negrón González 1986). About 40 percent of 

the refuge is comprised of three mangrove species: 

(1) Avicennia germinans, (2) Laguncularia racemosa, 

and (3) Rhizophora mangle. Much of the remainder 

is occupied by water with submerged, emergent, and 

floating vegetation (Chabert and others 1982). 

The Boquerón Wildlife Refuge includes a water 

impoundment that was built to help mitigate the 

loss of wetlands in the Lajas Valley a decade earlier 

(Chabert and others 1982). The impoundment 

measures 4 km in length and has six flood gates 

constructed along dikes to control water levels. The 

impounded wetland system is composed of a series of 

canals and small water bodies which are influenced by 

five sources of water, the first three being primary: 

•	 The brackish Laguna Rincón (to the west), which

varies in salinity according to rainfall 

•	 The Lajas Valley irrigation canal (to the east),

which drains the western part of Lajas Valley and 

provides the greatest portion of fresh water

•	 Direct rainfall captured during the wet season

•	 Quebrada Boquerón (to the north), which dilutes

brackish water coming from Laguna Rincón

•	 Runoff from areas south of the refuge

The impoundment structures retain water in the 

lagoon, controlling salinity, and creating open areas 

for the avifauna. The refuge also has a boardwalk for 

recreational use. 
56

Boquerón Wildlife Refuge. A boardwalk conducts visitors 

through mangrove forest. The refuge has 124 bird species in 30 

families, 60 of which are seen regularly. (Photo by Peter L. Weaver)

Several surveys of the refuge disclosed 124 bird 

species in 30 families, about 60 of which are seen 

regularly (Chabert and others 1982) (table 6). Bird 

populations are greatest during the winter months 

when the refuge provides a rest area for migratory 

species from North America. The refuge also has a 

rookery of Nyctcorax nyctcorax (black-crowned 

night herons). Other surveys showed the presence of 

(table 5): 

•	 Mammals—native Noctilio leporinus, and the 

introduced species Herpestes auropunctatus and 

Ratus ratus

•	 Amphibians—Bufo marinus, Eleuterodactylus 

antillensis, Eleuterodactylus coqui, and 

Leptodactylus albilabris

•	 Reptiles—Ameiva exsul; Anolis cooki, 

Anolis cristatelis, and Anolis puchellus; and 

Trachemys stejnegeri

Frog. Eleutherodactylus antillensis (Puerto Rican red-eyed frog) 

is found at the Boquerón Wildlife Refuge. (U.S. Fish and Wildlife 

Service photo)
57

Guánica Forest. Continuous monitoring of resident and migratory birds 

has been carried out at Guánica since the 1970s. Guánica, the largest 

forest in Puerto Rico’s reserve system, also has a visitor’s center. 

(Photo by Jorge Saliva)

Both of the exotic mammals negatively impact bird 

populations, often feeding on eggs. In contrast, the 

amphibians and reptiles constitute an important part of 

the diet of many birds. 

The lagoon contains at least 35 different genera of 

phytoplankton; moreover, several species of fish occur 

there, the most abundant being Tilapia mossambicus, 

Centropomus undecimalis, Mugil curema, and Caranx 

latus (Negrón González 1986) (table 5). Tilapia 

mossambicus, an introduced species, is more common 

than native species which also include Megalops 

altanticus (Burger and others 1992). Currently, 

the refuge is managed for hunting and fishing. It 

also provides opportunities for outdoor education, 

recreation, tourism, and crab catching. Unfortunately, 

dumping of garbage in the vicinity has been 

a problem. 

Guánica Commonwealth Forest

The Guánica Forest, lying 25 km east of Cabo 

Rojo along the south coast, occupies about 4016 

ha (table 1, fig. 1). The forest was created in 1919 

and was designated as a biosphere reserve in the 

United Nations Educational, Scientific and Cultural 

Organization—Man and the Biosphere Program in 

1981 (Departamento de Recursos Naturales 1985). 

Guánica is a semideciduous coastal limestone forest 

that was cut in the past for agriculture, wood, and 

fuel. Today, much of the woody vegetation is dense 

and of small dimensions (Murphy and Lugo 1986b). 

Tree species with multiple stems are common, a 

condition that appears due to environmental stresses 

such as drought, wind-transported salt, and periodic 

hurricanes (Dunphy and others 2000). 

Guánica, the best studied Subtropical Dry Forest 

on the island, supports nine vegetation types. Dry 

deciduous forest and dry forest with evergreen 

species account for 80 percent of the area (Dugger 

1979, see endnotes). The remaining types include 

mangroves, natural salinas, beaches, rocky areas with 

cactus, thorn forest, plantations, and disturbed areas. 

Guánica receives about 800 to 900 mm/year of rain 

and has a mean annual temperature near 26 °C (Ewel 

and Whitmore 1973, Lugo and others 1978). Actual 

and pan evapotranspiration average about 700 and 

2000 mm/year, respectively. Ravines and sinkholes 

have lower air temperatures and higher humidity, 

and may support plant assemblages different from 

the surrounding scrub vegetation (Farnsworth 1993). 

Extremely dry years rather than average years may 

be the most important factor controlling the forest 

structure and composition (Murphy and Lugo 1986a). 

Coppicing is common during succession and helps to 

hasten recovery. 

The flora of Guánica contains at least 500 species 

of vascular plants, 19 endemic to Puerto Rico, in 85 

families (Quevedo and others 1990). Bird species 

number more than 150 with 10 endemic to the island 

(Canals Mora 1990). Numerous caves provide habitat 

for five species of bats, one-third of the island’s 

native mammals (Conde Costas and González 1990). 

From 1922 to 1970, about 150,000 trees 

representing 9 species including 2 natives were 

planted on 150 ha in Guánica Forest (Wadsworth 

1990). Guanica’s low rainfall and recurrent droughts 

resulted in poor survival and slow growth for most 

species and indicated the importance of the dry forest 

for uses other than timber production. However, 

the introduced Haematoxylon campechianum, 

planted in 1924, showed 75 percent survival after 
58

Los Morrillos de Cabo Rojo (Cabo Rojo Lighthouse area). A lone hiker ascends the trail to the Cabo Rojo Lighthouse. Built in 

1882, the lighthouse casts a beam that is visible for nearly 25 km. (Photo by Peter L. Weaver)

18 years although growth was slow (Wadsworth 

1943). Moreover, other exotics such as Leucaena 

leucocephala and Swietenia mahagoni appear to be 

permanent additions to the flora of the dry forest. Of 

the 167 tree species reported for the Guánica Forest, 

23 were introduced (Chinea 1990). Of the introduced 

species, only eight showed some degree of success, 

three in forested conditions and five in the open.

Past measurements of tree diameters showed slow 

growth in comparison to wetter areas on the island 

(Anon. 1951, Wadsworth 1990, Weaver 1979). Forest 

studies were initiated in the 1970s and permanent 

plots were established during the 1980s; subsequently, 

information has been published on forest structure, 

biomass, and nutrient cycling (Lugo and Murphy 

1986; Lugo and others 1978; Murphy and Lugo 

1986b, 1990; Murphy and others 1995). It is probable 

that most of the plant studies and virtually all the 

avifaunal studies carried out at Guánica are applicable 

to much of the Subtropical Dry Forest region in the 

southwest, including the F&WS refuge sites. 

Los Morrillos de Cabo Rojo Lighthouse

Cabo Rojo, the southwest tip of Puerto Rico, is a 

tombolo that is comprised of two separate limestone 

islands of Miocene and Oligocene age connected to 

each other and the mainland by sand bars (Kaye 1959,

Monroe 1976, Weaver 1976). The 105-ha property is 

controlled by the Department of Natural Resources. 

The area around the lighthouse, previously under the 

U.S. Coast Guard, is currently managed by the Cabo 

Rojo municipality as part of SBAPES.

Wave-cut cliffs characterize the island units of the 

tombolo where the Cabo Rojo Lighthouse is located. 

The lighthouse, constructed in 1882 and recently 

restored as a museum and educational center, is 

recorded on the National Register of Historic Sites 

of the U.S. Department of the Interior (Departmento 

de Recursos Naturales y Ambientales 2000b). The 

lighthouse beam, which emanates from 37 m above 

sea level, is visible for nearly 25 km (Cerame 

Vivas 1988). 
59

Parguera Natural Reserve. The 408-ha reserve, managed by the Puerto Rican Conservation Trust, is situated near the fishing 

village of Parguera. Dry scrub forest, mangroves, and salt flats dominate the reserve. A bioluminiscent bay, numerous coral and 

fish species, scenic views, and opportunities for boating are available to tourists. (Photo by Joseph Schwagerl)

The environmental impact statement developed 

for the lighthouse and surrounding areas contained 

the following details on flora and fauna (Municipio 

Autónomo de Cabo Rojo 1998):

•	 The highly impacted vegetation is secondary and

contains at least 52 plant families with at least 

147 species, 3 of which—Bulbostylis pauciflora, 

Polygala hecatantha, and Waltheria calcicola—are 

listed as critical.

•	 The avifauna is represented by 27 bird families

with 52 species. Two species—Agelaius 

xanthomus and Pelecanus occidentalis—are 

listed as critical, and two others—Charadius 

alexandrinus and Sterna antillarum—are listed as 

critical and vulnerable (table 6). 

•	 The herpetofauna is represented by seven lizards

(two ameiva and five anoles), with Anolis cooki

listed as critical and vulnerable (table 5).

•	 During the past several years two sea turtles—

Eretmochelys imbricata and Dermochelys 

coriacea—have been recorded as using nearby 

beaches for nesting.

Parguera Natural Reserve

The Parguera Natural Reserve occupies 408 ha 

along the coast between the Boquerón and Guánica 

Forests (table 1, fig. 1). The reserve, managed by the 

Conservation Trust of Puerto Rico, contains numerous 

beaches, cays, salt flats, and islands dominated by 

mangroves, beach thicket, salt tolerant vegetation, and 

scrub woodland (Anon. 1982). 

The fishing village of Parguera and surrounding 

areas were first brought to national attention through 

an article in National Geographic describing the 

“Bahía Fosforescente,” which is situated a short 

distance to the east (Zahl 1960). Subsequently, the 

bioluminescent organisms in the bay were studied 
60

and plans devised for the protection of the area, 

including the watershed above the bay (National Park 

Service 1968, Seliger and others 1971). Floral and 

faunal surveys in and adjacent to the reserve reported 

(tables 1, 5, and 6):

•	 184 species of plants, including the endemics

Cordia rickseckeri, Machaonia portoricensis, 

Rondeletia inermis, Thounia striata, and Wedelia 

lanceolata; species listed as critical such as 

Rochefortia acanthophora and Bulbostylis 

pauciflora; and species listed as endemic and 

critical such as Leptocerus quadricostatus and 

Psychilis krugii

•	 64 species of birds, including the endemics—

Caprimulgus noctitherus, Loxigilla portoricensis

(Puerto Rican bullfinch), Todus mexicanus

(Puerto Rican tody), and Vireo latimeri (Puerto 

Rican vireo); the federally endangered Pelicanus 

occidentalis; the endangered endemic Agelaius 

xanthomas; and the rarely seen Stercorarius 

pomarinus (pomarine jaeger) five species of 

herpetofauna (one gecko, two ameivas, and two 

anoles); including the endemics Sphaerodactylus 

nicholsi and Ameiva wetmorei

Surveys for native mammals were unavailable in 

1981 (Departamento de Recursos Naturales 1981). 

At that time, however, the bat Noctilio leporinus was 

considered a probable resident. Also, feral Macaca 

mulatto, long ago having escaped from Isla Magueyes, 

still occasionally roam the reserve. 

Punta Guaniquilla Natural Reserve

Since 1976, the 170-ha Punta Guaniquilla Natural 

Reserve has been administered by the Conservation 

Trust of Puerto Rico. The property contains the 

remnants of Hacienda La Romana, a sugar producing 

estate built during the 1870s (Fuentes Santiago and 

Quevedo Bonilla 2002; Ramos y Ramírez de Arellano 

1986, see endnotes). The ruins of the Hacienda 

including walls, a well and a cistern, are still evident 

today. Several archeological sites, probably Taino, 

have also been identified through historical records 

and verbal accounts. 

The reserve extends from sea level to 46 m in 

elevation at Cerro Guaniquilla. There are two cave 

systems in limestone of Cretaceous age, one reputed 

to have contained petroglyphs in the past (Monroe 

1976; Ramos y Ramírez de Arellano 1986, see 

endnotes). The other, erroneously associated by the 

public with the death of the famous Cabo Rojo pirate, 

Roberto Cofresí (1791–1825), has dozens of tunnels. 

Actually, it was Cofresí’s nephew, Antonio Cofresí, 

an interpreter in the Port of Mayagüez, who died in 

the cave. 

The land contains upland and wetland habitats with 

seven major vegetation types (Quevedo Bonilla 2000, 

Vázquez 1997, Vázquez and Kolterman 1998):

•	 Pasture—some areas heavily impacted by grazing

•	 Shrubland—areas in transition to woodland

•	 Woodland—canopy from 6 to 9 m tall and

dominated by a few tree species

•	 Evergreen forest—canopy to 12 m with a mixture

of several tree species

•	 Marsh—flooded seasonally for extended periods

•	 Beach—sand, rocky areas, and few tree species

including red mangrove

•	 Beach thickets—to 5 m tall with several tree

species including white and black mangroves

The vascular flora consists of 253 species, 70 

percent of them shared with Guánica Forest, and 

35 percent of them trees. Endemic trees include 

the Gaussia attenuata, Machaonia portoricensis, 

Rondeleteia inermis, and Thouinia striata. The other 

endemic species are Hohenbergia antillana and 

Zamia portoricensis. The presence of Cynometra 

portiricensis, Oxandra lanceolata, and Stahlia 

monosperma suggest that the reserve could be 

considered as critical habitat for these species. Eightyone bird species have been identified at the reserve 

(Fuentes Santiago and Quevedo Bonilla 2002). Five 

are endemic and three are classed as rare or extremely 

rare (table 6). 
61

Punta Guaniquilla Natural Reserve. Managed by the Puerto 

Rican Conservation Trust, the 152-ha reserve contains the 

remains of Hacienda La Romana. The reserve is also well 

known for its limestone rocks which have eroded into attractive 

formations. (Photo by Peter L. Weaver)

Laguna Guaniquilla occupies 9 ha of the reserve 

(Negrón González 1986). The lagoon has a mean 

depth of 0.4 m and surrounds rocks that have been 

eroded into attractive formations. The lagoon was 

previously connected to the sea via a canal that was 

closed around 1955 to retain fresh water runoff 

(Ramos y Ramírez de Arellano 1986, see endnotes). 

Before then, the lagoon was used to trap marine fish 

where they were grown for domestic use. The only 

fish species recorded in the lagoon today are Tilapa 

mossambicus, Mugil curema, and Gerres cinereus. 

Tilapa mossambicus accounted for two-thirds of the 

catch. In the past, the area was used by Dendrocygna 

arborea (West Indian whistling duck) as a nesting site. 

General Observations and Research 

in Southwestern Puerto Rico 

Major wildlife research for the refuge sites is 

summarized below along with information from other 

protected areas in southwestern Puerto Rico. Most of 

the formal research has been carried out at four sites: 

the Cabo Rojo and Laguna Cartagena NWRs and 

the Commonwealth forests at Boquerón and Guánica. 

In a few instances, important observations not 

available from the southwest have been reported from 

other areas. 

All research activities carried out partially or 

completely on the refuges—either at Cabo Rojo 

(Headquarters or Salinas tracts) or at Laguna 

Cartagena (Lagoon or Tinaja tracts)—are included as 

refuge studies. In some instances, both refuges were 

involved along with other sites, usually the Boquerón 

or the Guánica Commonwealth Forests. 
62

Flora—Mangroves and Phenology

Between 1974 and 1989, mangroves throughout 

Puerto Rico showed a 39.5-percent increase in 

area (Torres Rodríguez 1993). Along the entire 

southwestern coast from Guánica to Punta 

Guaniquilla, sampled mangrove areas increased 

from about 800 to 970 ha (21.2 percent). Within 

this region, the areas around Guánica and Cabo 

Rojo combined increased from 356 to 537 ha (50.8 

percent) whereas the area around Parguera decreased 

slightly from 445 to 433 ha (2.7 percent). Several 

reasons were suggested for the increase in mangrove 

areas including a decline of agriculture, reduction in 

conversion to other uses, isolation in some instances, 

legal protection, and the natural recovery of mangrove 

areas when left undisturbed. 

Information on phenology (flowering, fruiting, 

and leaf loss) is important for determining potential 

periods of limited food supply and possible habitat 

preferences for animal populations, particularly 

during droughts. Likewise, data on the origin of tree 

species are useful for the restoration of native habitat. 

Although detailed studies are not available for all 

of the island’s tree species, general phenological 

observations have been recorded over the years 

(Francis and Lowe 2000, Little and Wadsworth 1964, 

Little and others 1974). This information has been 

summarized for 162 tree species growing on or near 

the refuges (appendix A). Of the total, 135 trees are 

native, i.e., about 30 percent of the island’s total 

native species, and the remainder are introduced. 

Of the natives, 13 are endemic to Puerto Rico. 

Slightly more than 100 species are evergreen, about 

50 are deciduous, and most of the remainder may 

be evergreen or deciduous depending upon weather 

conditions (appendix A). 

Patterns of flowering and fruiting for 48 woody 

species in deciduous forest at Guánica were described 

as complex, with considerable variability within and 

among species (Murphy and others 1995). A 3-year 

study showed that flowering is rarest during the 

February to -March dry season and more common 

from April through October, after which it begins 

to decline. Fruits, in turn, are relatively abundant 

throughout the year when all 48 woody species are 

considered together; most fruits, however, seem to 

fall during the dry season. About one-half of the 

48 species bear fruit twice annually, and nearly 

one-quarter have some fruiting specimens at least 

every other month (Murphy and others 1995). In 

addition, another 3-year study at Guánica showed that 

the amount of leaves, as determined by measures of 

leaf area and forest cover, was least during dry season 

and greater during the rest of the year (Murphy and 

Lugo 1986b).

Fauna—Birds

Early observations in western Puerto Rico and 

offshore islands revealed more than 160 bona fide 

species and subspecies, nearly one-half of which were 

migratory (Danforth 1926, 1931, 1936; Struthers 

1923). In the early 1960s, a checklist updated known 

species and clarified Spanish nomenclature for the 

island’s birds (Leopold 1963). Later, as part of Puerto 

Rico’s encyclopedia of natural resources, historical 

bird records from 1535 to present, including extinct 

species, were compiled by the Department of Natural 

Resources (Chabert Llompart and others 1986). 

Notes on bird feeding habits, reproductive ecology, 

behavior, and vocals were included. Several other 

publications highlight species’ identification, some 

with photographs or sketches of the island’s species 

(Hernández-Prieto 1994, Oberle 1999, Ortiz Rosas 

1981, Toro McCown and Chabert 1986) (see also 

table 6, footnotes). In addition, bird species’ guides 

(Raffaele 1983, 1989) include recent arrivals and 

summarize identifying characters, English and local 

names, favored habitats, flight and feeding behavior, 

voice, nest data, and regional distributions for all 

species among other observations. 

Cabo Rojo National Wildlife 

Refuge Studies—Headquarters 

Tract and Municipality

Flora—Tree Genetics, Mistletoe, 

and Restoration

Landscape fragmentation can lower visitation 

by pollinators and lead to inbreeding and increased 

genetic drift (Dunphy 2003, Dunphy and others 

2004). Accurate estimates of gene flow, however, 

may provide information on how certain species will 

endure habitat losses. Gene exchange in fragmented 

forest was explored for three species: (1) Bursera 

simaruba and (2) Hymenaea courbaril, both native 
63

tree species; and (3) Albizia lebbeck, an exotic tree 

introduced during historic times. For all species, a 

substantial proportion of seed production was the 

result of gene-flow pollen. Hymenaea courbaril, 

a bat pollinated species, had the lowest geneflow rate. Genetic diversity values were higher in 

Bursera simaruba and Albizia lebbeck than for other 

species with similar life history characteristics, 

and conversely, lower in Hymenaea courbaril. No 

relationship was found between gene-flow rates and 

distance to nearest neighbor or stand size. In another 

study, an analysis of 16 populations of Rhizophra 

mangle from the Dominican Republic; Florida; 

Panama; and Cabo Rojo, Puerto Rico, i.e., Bahía 

Sucia and the Boquerón Wildlife Reserve, was carried 

out to determine genetic variation within the species 

(Pérez Laguillo 1998). 

Dendrophthora bermejae, a recently discovered 

endemic mistletoe, is confined to a few localities in 

Cabo Rojo, including Sierra Bermeja (Kuijt and others

2005). The mistletoe’s only known host is Guaiacum 

officinale. The mistletoe is currently threatened by 

residential, commercial, and tourist development 

and should be established in protected areas in 

southwestern Puerto Rico.

An attempt was made to assess the potential of 16 

native tree species for restoration of subtropical dry 

areas, including the refuge sites (Caracal Velez 2001). 

Light and soil requirements, dry storage longevity of 

seeds, and early seedling survival were investigated in 

greenhouse conditions. All of the species tested grew 

naturally within the refuges and were suitable 

for reforestation.

Fauna—Birds 

Agelaius xanthomus (yellow-shouldered 

blackbird)— In 1974, an early morning survey of 

the Cabo Rojo Headquarters tract disclosed 26 bird 

species, 7 percent of which were Agelaius xanthomus

(Wiley 1974, see endnotes). The tract was suggested 

as a buffer zone to protect the mudflats on the Salinas 

tract to the south. At that time, the blackbirds used the 

Headquarters tract for feeding and daytime roosting. 

Cathartes aura (turkey vulture)—Cathartes aura, 

the only falconiformes established in Puerto Rico, 

is frequently sighted between Cabo Rojo and Ponce 

(Perez Rivera and Cotte Santana 1977, Santana and 

others 1986a). Nests have been located on the refuges 

and in nearby Lajas caves and in the Cerro de Cotui 

in San Germán. Observations made near Lajas show 

that peak soaring occurred in the early afternoon. The 

minimum home range during an 18-month period 

was 460 km2

. Daily range, however, varied from 1.2 

to 30.5 km2. A census suggested that Puerto Rico was 

not an important wintering area for migrating vultures 

(Santana and others 1986b). Other surveys report on 

the geographical distribution, and feeding and nesting 

habits of Cathartes aura, Buteo jamaicensis (redtailed hawk), and Falco sparverius (American kestrel) 

(Pérez Rivera and Cotte Santana 1977).

Columbids (pigeons and doves)—Predation accounts 

for 81 percent of columbid nest losses (RiveraMilán 1996). Moreover, in the Guánica and Cabo 

Rojo Forests, the total amount of rainfall during the 

first 6 months of the year accounted for about 70 

percent of the variability associated with nest density 

estimates during the reproductive peak. The presence 

of food plants and watering areas at the Headquarters 

tract, along with protection, could benefit columbid 

populations (Wiley 1974, see endnotes). 

Dendroica adelaidae (Adelaide’s warbler)—

The vocal behavior of the Dendroica adelaidae

was studied in relation to its social behavior and 

ecology (Staicer 1991). Breeding was seasonal with 

individuals maintaining monogamous pair bonds 

and exclusive pair territories throughout the year; 

however, when unpaired, only males maintained 

territories. Paired birds used duets of male songs and 

female calls. 

Myiarchus antillarum (Puerto Rican flycatcher)—

The foraging modes and resource use by Myiarchus 

antillarum are diverse (Cruz and others 1986). 

The flycatchers capture prey nearly one-half of the 

time when both the bird and prey are in the air. The 

remaining captures are when both are on a substrate 

or when the prey is on a substrate and the bird on 

the wing. Flycatchers also have a varied diet which 

includes mollusks, arthropods, and vertebrates, e.g., 

Anolis spp.; and occasionally, fruits. During the 

late 1980s, breeding biology and growth patterns of 

flycatchers were studied (Torres Báez and Collazo 

1992). Flycatchers breed from February through June 

with a peak in May and June. About three-quarters 

of the nests fledged at least one young. Flycatchers 
64

accept artificial cavities and will use wooden nest 

structures. Nest cavity dwellers fledge better than 

those that nest on the soil or on platforms. 

Dendroica discolor, Dendroica tigrina, Parula 

americana (prairie, Cape May, and northern 

parula warblers)—The behavior of Dendroica 

discolor, Dendroica tigrina, and Parula americana

was studied to identify factors of importance for 

maintaining wintering populations (Staicer 1992). 

Most individuals exhibited strong site fidelity within 

and among wintering seasons. Usually they roamed 

independently, but sometimes were found in small, 

cohesive flocks. Dendroica tigrina maintained 

intraspecific territories and chased away other species. 

Parula americana were more plastic in behavior, 

ranging from territorial to gregarious to wandering. 

Dendroica discolor, in turn, was more prone to avoid 

other warblers, moving around widely. Current longterm studies demonstrate population declines for 

all three species in the wintering habitat at Guánica 

Forest (Arendt and collaborators 1992). 

Falco sparverius (American kestrel)—Falco 

sparverius feeds on insects, lizards, and mice (Perez 

Rivera and Cotte Santana 1977). The species has 

purportedly declined because of the lack of nesting 

sites. In response, 20 nest boxes were installed on the 

Cabo Rojo NWR, mainly at the Headquarters tract, 

where 2 currently have active nests.

Other Fauna

Numerous Antiacarsia gemmatalis (velvet bean 

caterpillar) larvae were observed feeding on the 

weedy legume Tephrosia cinerea at Cabo Rojo 

(Gregory and others 1991). Three bird species: (1) 

Molothrus bonariensis, (2) Quiscalus niger (Greater 

Antillian grackle), and (3) Agelaius xanthomus feed 

on the larva. Apparently, this is the first report of the 

caterpillar’s native host plant in the West Indies. 

Gecko populations of Sphaerodactylus 

nicholsi at the Cabo Rojo NWR vary with habitat, 

preferring evergreen vegetation that offers cover 

from insolation (López-Ortiz 1999). Evergreens help 

stabilize temperatures and provide leaf litter and 

humus during the year, supporting populations of 

arthropods. Arboreal lizards are sympatric and are 

potential predators. 

Cabo Rojo National Wildlife Refuge 

Studies—Salinas Tract

Lagoon System and Its Wildlife 

During autumn migration and winter months, the 

hypersaline lagoon system of the salt flats is the most 

important converging point for migrating shorebirds in 

Puerto Rico (Grear 1992a, 1992b; Grear and Collazo 

1999). Surveys recorded 32 species of shorebirds with 

Calidris pusilla and Calidris mauri (small calidrids) 

accounting for 60 to 70 percent of the species. The 

maximum populations were recorded in mid-October 

when changes in invertebrate abundance coincided 

with fluctuations in lagoon salinity. Shorebird diets 

were associated with food abundance and changes in 

water levels. Every site within the salt flats serves as 

an important habitat for at least one shorebird species. 

Additional studies should focus on the hydrology of 

the lagoons and the effects of water level fluctuations 

on salinity; the availability of primary producers, 

i.e., algae; and prey dynamics. The abundance of 

calidrids and their prey was correlated with the algal 

substrate. Fishermen report that two families of fish—

Centropomidae (snook) and Mugilidae (mullets)—

regularly enter the lagoons (Negrón González 1986) 

(table 5). 

Artemia spp. (brine shrimp) are suitable food for 

a wide variety of organisms, including crustaceans 

and fishes, and are also used in the development of 

aquacultural systems. Artemia populations in Laguna 

Fraternidad were influenced mainly by fluctuations 

in temperature and salinity and not other factors such 

as pH, dissolved oxygen, or turbidity (Bonilla Soto 

1984). The highest shrimp populations were recorded 

in May; however, shrimp were absent in August 

because of critical temperature and salinity levels. The 

lethal upper temperature and salinity limits for brine 

shrimp are 36 °C and 200 parts per thousand (p/t). 

The density and distribution of Trichocorixa spp. 

(water-boatmen) and Artemia spp. were monitored 

from early September to early November, the fall 

migratory season (Tripp 1996, Tripp and Collazo 

2003). The former were recorded in areas of lowest 

salinity (<65 p/t) and the latter in areas with the 

highest salinity (≥106 p/t). The foraging value 

of the salt flats for shorebirds may be enhanced 

by maintaining salinity at <65 p/t in selected 
65

management units, and by controlling water depth. 

In a second study carried out from January through 

December, populations of Artemia salina and 

Gomphosphaeria aponina (phytoplankton) increased 

during the period of heaviest rainfall, coincident with 

lowest salinity (Mercado Alvarez 2003). Alterations 

of populations were noted, suggesting that the shrimp 

feed on the phytoplankton. Samples of nitrates, 

nitrites, dissolved oxygen, and pH were also collected. 

Fauna—Shorebirds and Wading Birds 

Charadrius alexandrinus and Charadrius wilsonia

(snowy and Wilson’s plovers)—A snowy plover 

was first reported on the salt flats near the Cabo Rojo 

Lighthouse in 1928 (Danforth (1929). In 1988, a 

study of the snowy and Wilson’s plovers reported on 

their abundance, nesting chronology, nesting success, 

nesting habitat, and habitat use (Lee 1989). Peaks in 

the abundance of snowy plovers apparently coincide 

with the influx of migrants. Nest sites differed by 

species. Wilson’s plovers preferred live vegetation 

nearby whereas the snowy plovers nested on open 

sand. Chick mortality for both species appeared high. 

Causes of nest failure were due to abandonment for 

the snowy plover, and human disturbance and flooding 

for the Wilson’s plover. Sandflats appeared to be 

important roosting sites for both species. 

Calidris fuscicolis, Calidris mauri, Calidris 

minutilla, and Calidris pusilla (white-rumped, 

western, least, and semipalmated sandpipers)—

Banding of Calidris pusilla and Calidris mauri

from 1989 to 1993 provided information on their 

residency rates, i.e., length of stay in the 1992 season; 

annual return rates between 1989 and 1991; and their 

populations in 1992 for the Cabo Rojo Salt Flats 

and nearby Boquerón Forest at Punta Pitahaya (Rice 

1995). The residency rates suggested that the salt flats 

serve as both stopover and wintering area. Moreover, 

the return rates suggested that the salt flats are 

traditionally used by both species. About 7,300 birds 

of both species were estimated to use both areas in the 

fall of 1992, with Cabo Rojo accounting for about 90 

percent of the total. 

Nonbreeding territorial behavior for Calidris 

pusilla, Calidris minutilla, and Calidris mauri

was studied at the Cabo Rojo Salt Flats (Tripp and 

Collazo 1997). Territorial birds defended areas with 

greater prey densities than nonterritorial birds, using 

territories as large as competition permitted. The 

cost of territorial behavior may be offset by foraging 

in areas where prey density is greater, or through 

increased foraging efficiency. Nonterritorial birds, 

however, also foraged in areas where prey was more 

abundant than for average sites within the lagoon.

Three short-term student studies examined Calidrid

species at Fratrenidad Lagoon, a water body where 

foraging areas remain relatively constant diurnally. 

The first showed that Calidrid species preferred 

the shallow and warmer north shore of the lagoon, 

possibly because of differences in depth, or the lack 

of littoral vegetation that could hide avian predators 

(Santos 1988, see endnotes). The second study looked 

at habitat selection and foraging behavior in mixed 

flocks of Calidris fuscicolis, Calidris mauri, Calidris 

minutilla, and Calidris pusilla (Hopping 1989, see 

endnotes). The third investigated bill morphology 

of Calidris mauri, Calidris minutella, and Calidris 

pusilla in regard to prey size and substrate preferences 

(Bailey 1990, see endnotes). 

A recurrent census of migratory and resident 

shorebirds identified 28 species between 1985 and 

1992 (Collazo and others 1995). The highest number 

of individuals was recorded between August and 

December and in March. Migrants accounted for 87 

percent of the annual counts. Calidrid sandpipers,

Tringa flavipes (lesser yellowlegs), and Calidris 

himantopus (stilt sandpipers) were the most abundant 

migrants and Himantopus mexicanus (black-necked 

stilts), the most common resident bird. Peak daily 

counts ranged from about 3,500 to over 8,000 

individuals. Distribution of the birds was affected by 

food availability and hydrological conditions, both a 

function of the salt extraction operations. 

The stomach contents of 35 wading birds of 4 

species were examined in 1994 and 1995 to determine 

their feeding habits (Miranda 1995b, Miranda and 

Collazo 1997a). Eleven prey groups were identified 

including 7 families of fish, 2 genera of crustacea, 

a lizard, and insects. Egretta tricolor (tricolored 

herons) fed on guppies and snook. Ardea alba (great 

egrets) and Egreta thula (snowy egrets) fed mainly on 

shrimp and guppies, and Egreta caerulea (little blue 

herons) fed exclusively on fiddler crabs. A modified 

throw-trap was used to sample prey in shallow water 

mangrove swamps (Miranda and Collazo 1997b). 

The trap was light, easy to use, and suitable for 

sampling prey in areas with an uneven bottom in a 

standardized fashion. 
66

Fauna—Sea Turtles 

Dermochelys coriacea were reported nesting in 

1991, 1994, and 1997 and Eretmochelys imbricata

in 1999 on the beaches of southwestern Puerto Rico 

(Jiménez Marrerro 2000, see endnotes). A survey 

of the beaches at Bahía Salinas, Punta Águila, 

and Playa Combate in the winter of 1999–2000 

showed that motor vehicles cause serious problems. 

The beaches were judged satisfactory for turtle 

nesting, and their actual use could be documented by 

increased monitoring.

Laguna Cartagena National Wildlife 

Refuge Studies—Lagoon Tract

Water Quality, Insects, and Earthworms 

Physical and chemical conditions of the lagoon 

were assessed, including pH, temperature, dissolved 

oxygen, nutrients, and heavy metals (Deliz Quiñones 

2005). Lagoon conditions, vegetation, and insect 

populations all varied during the year. Ammonia and 

phosphate were at high levels, and dissolved oxygen 

and lagoon depth showed significant fluctuations. 

Sampling yielded 67 species of insects in 33 families. 

Insect abundance and diversity were related mainly to 

the amount of dissolved oxygen.

Only four earthworm species were encountered at 

Cartagena Lagoon (Alfaro and Borges 1996). The low 

diversity and density of earthworms was attributed 

to several factors, including high organic matter 

accumulation, the high resistance to penetration in the 

topsoil layer, high salinity, sand content in deeper soil 

layers, and the effects of rainfall on shallow soils. 

Native Bird Species 

Populations of four native aquatic birds: (1) Anas 

bahamensis (white-cheeked pintail), (2) Dendrocygna 

arborea (West Indian tree duck), (3) Fulica caribaea

(Caribbean coot), and (4) Oxyura jamaicensis (ruddy 

duck) were abundant at the beginning of the 20th

century (Bonilla and others 1992). All species were 

negatively affected by habitat loss, i.e., drainage 

of the Ciénaga El Anegado and Laguna Guánica, 

and deterioration of the Cartagena Lagoon during 

the 1950s and 1960s; excessive hunting and egg 

removal; occasional flooding; and predation by rats 

and mongooses. Islandwide waterfowl investigations 

indicated that all species are less common today and 

their hunting has been prohibited by the Department 

of Natural Resources (Belitsky 1978, Bonilla 1992, 

Chabert 1984, see endnotes for each). Interestingly, 

all four species have been reported recently at the 

Cartagena Lagoon, including nesting individuals of 

Dendrocygna arborea. Only one Fulica caribaea was 

reported in 1991–92 at Cartagena Lagoon. In addition, 

an Anas rubripes (American black duck) recorded at 

the lagoon January 1980 was later shot by a hunter 

(Colón 1982b). The previous recording for this 

species was at Laguna Guánica in December 1935. 

Communal breeding of Crotophaga ani (smoothbilled ani) was studied at Cabo Rojo and Laguna 

Cartagena where groups ranged from 2 to 9 adults 

with communal clutches between 11 and 25 eggs 

(Startek 1997). DNA loci were used to examine 

the mating system and extent of reproductive skew, 

i.e., the distribution of reproduction among group 

members. Within four focal groups, suggestions of 

monogamy, polygamy, and incidents of intraspecific 

brood parasitism were found. Alternative strategies 

to monogamy may be more prevalent than previously 

believed, and the reproductive skew may be affected 

by the availability of limiting resources during the 

breeding season. A second study addressed specific 

DNA markers, parentage of buried eggs, and adult 

relatedness (Blanchard 2000). Evidence emerged 

for an egalitarian system with low reproductive skew. 

The most significant factors affecting bias were 

the order of egg laying, timing, and number of 

breeding females. 

Members of a group tended to stay together during 

regular activities such as roosting, foraging, and 

resting, where they are better able to defend their 

nests and territories than would a single pair (Lentz 

2005). Shared incubation and sentinel behavior 

during foraging are group behavioral activities 

that may lower mortality. Territory quality, i.e., as 

determined by the time of first breeding and chances 

for renesting or second breeding, correlated negatively 

with group size. Moreover, ani habitats were not 

saturated, i.e., suitable breeding habitat was available 

but unoccupied. Buteo jamaicensis, Felis domesticus

(domestic cat), Rattus rattus, and Solenopsis invicta

(fire ants) were observed preying on ani nests 

(Lentz 2005). 
67

Laguna Cartagena National Wildlife 

Refuge Studies—Tinaja Tract

Secondary Succession and Restoration 

The natural recovery of trees after the elimination 

of fire and grazing was assessed on the Tinaja tract, 

including species-site relationships and changes in 

species composition over time (Weaver and Chinea 

2003, Weaver and Schwagerl 2005). Past land use 

history, slope, and distance to drainages accounted 

for variability in species occurrence. The survey 

disclosed at least 161 tree species on the Tinaja tract. 

The 10 most common species accounted for nearly 60 

percent of the stems and the 43 least common species, 

each with 5 or fewer individuals, accounted for only 

2.2 percent of the stems. In addition, the survival, 

growth, and future potential of tree species for site 

restoration were evaluated for both refuges (Weaver 

and Schwagerl 2004). 

Boquerón Commonwealth Forest

Fauna—Bird Species

Agelaius xanthomas (yellow-shouldered 

blackbird)—Agelaius xanthomas, endemic to 

Puerto Rico, including Isla Mona, declined from 

very abundant in 1864, to common in 1927 through 

the 1940s, to uncommon and restricted in range in 

the early 1970s (Post and Wiley 1976). The range in 

the 1970s was from Ensenada to Punta Guaniquilla 

in southwestern Puerto Rico. Among the factors 

suggested for its decline were loss of feeding habitat, 

reduction in nesting habitat, introduced rats and 

mongooses, fowl pox, and two competitive bird 

species—Margarops fuscatus (pearly-eyed thrasher)

and Molothrus bonariensis. Higher blackbird nest 

success on cays as compared to mainland sites was 

attributed to the lack of brood parasitism offshore. 

The roosting behavior of the blackbird was 

studied along with observations on the Margarops 

fuscatus and Quiscalus niger (Post and Post 1987). 

All of these species use primary (nocturnal) and 

secondary (diurnal) roosts. Primary roosts are 

located in predator-free sites, i.e., offshore islands, 

and secondary roosts in subcanopy trees near food 

sources. Blackbirds do not stay in nesting colonies at 

night unless they are incubating or brooding. Predator 

mobbing was common, and arrival and departure 

patterns from roosting sites varied among species.

The blackbird was listed as endangered in 

1976 because of brood parasitism by Molothrus 

bonariensis, an activity that reduced its nesting 

success (López-Ortiz and others 2002, Post 1981a, 

Wiley and others 1991). Between 1974 and 1982, 

blackbird populations declined by about 80 percent. 

Radio telemetry was used to learn more about 

blackbird patterns of movement and habitat use during 

the nonbreeding season (Cruz-Burgos 1999). The 

mean home range was about 230 ha during both the 

pre- and postbreeding season. Prosopis juliflora stands 

and residential areas with household feeders were 

used more frequently during the prebreeding season; 

caterpillars and vegetable matter were favored in scrub 

and mangrove woodlands during the postbreeding 

season. Foraging was mainly during the morning and 

afternoon, with the midday spent resting and preening.

An artificial nest structure program, initiated 

in the Boquerón Forest in 1984 and coupled with 

removal of Molothrus bonariensis, improved the 

blackbird’s reproductive success (Wiley and others 

1991). Blackbirds accepted the nest boxes which 

were concentrated in mangrove habitats. Avicennia 

germinans trees were later surveyed in areas east and 

west of Punta Pitahaya (Reitsma 1998, see endnotes). 

The dramatic increase in the number of natural 

blackbird nests was probably due to the success of 

the artificial nest structures installed during the past 

several years. From 1996 to 2000, the percentage 

of blackbird nests in natural substrates as opposed 

to artificial structures increased from about 1 to 35 

percent (López-Ortiz and others 2002).

In 1987, a flock of about 300 blackbirds were 

sighted foraging on caterpillars along with Quiscalus 

niger, and 1 day later, about 100 blackbirds with 

Quiscalus niger and Molothrus bonariensis

(McKenzie and Noble 1989). Both were in a

woodland of Prosopis juliflora near the Boquerón 

Forest north of Punta Pitahaya. A study of blackbird 

ectoparasites near Parguera showed 3 species of avian 

biting lice on 69 percent of the 265 birds sampled, 

with greater occurrence on males than females (Post 

1981b). Also, three species of mites were reported 

in blackbird nests. The frequency of avian pox 

lesions was also recorded and it was suggested that 
68

deformities resulting from avian pox could reduce 

blackbird life expectancy. One incident of the capture 

of a blackbird in flight by Falco sparverius was 

thought to be related to pox lesions (Miranda 1995a). 

Since 1980, the Commonwealth Department of 

Natural Resources and the F&WS have undertaken 

measures for the recovery of the blackbird including 

trapping and removing Molothrus bonariensis from 

blackbird nesting areas; evaluating and controlling 

other pests infesting blackbird nests and chicks; and 

installing, cleaning, and repairing artificial blackbird 

nest structures (Román Cordero 1992, Ventosa Febles 

1991). The work is being carried out in the salt flats 

and mangroves of Boquerón Forest. The program has 

had a positive effect on the blackbirds, increasing their 

nest success and the number of birds fledged. 

Molothrus bonariensis (shiny cowbird)—Molothrus 

bonariensis ranges from Patagonia to Hispaniola. 

The species was first seen in Vieques in 1860, and 

first recorded in Puerto Rico in 1955, although it had 

apparently arrived earlier (Cruz and others 1998; 

Post and Wiley 1977a, 1977b). The species is kept 

as a cage bird and its movement has probably been 

facilitated by humans. Once in Puerto Rico, the 

cowbird, a host generalist, encountered avian species 

with no history of social parasitism (McKenzie and

Noble 1990, Wiley 1985). Cowbird puncturing of eggs 

was examined between 1982 and 1988 (Nakamura and 

Cruz 2000). The frequency of punctures was related 

to the number of cowbird eggs in the nest. Moreover, 

cowbirds punctured eggs indiscriminately, i.e., their 

own as well as those of the host. Egg puncturing 

occurred at any time during incubation, and clutches 

with damaged eggs were abandoned by the host. 

Cowbirds are good colonists because of their 

flexibility in habitat selection, diverse diet, gregarious 

behavior, and fecundity; moreover, cowbirds are 

host generalists, successfully parasitizing at least 

11 species in Puerto Rico (Cruz and others 1989; 

Wiley 1985, 1988). A study in southwestern 

Puerto Rico showed that cowbirds used Prosopis

juliflora woodland about 75 percent of the time; 

and agricultural land, littoral woodland, mangrove, 

scrub, and open fields the remainder (McKenzie and

Noble 1990). In mangroves, 42 percent of resident 

nonraptorial land bird species were affected (Wiley 

1985). The cowbirds do not parasitize birds in 

proportion to their abundance (Wiley 1988). The 

food habits and egg size of Molothrus bonariensis

are similar to those of their hosts, suggesting that 

they choose hosts based partly on those characters. 

Cowbirds locate nests by watching potential hosts in 

likely habitats, using cues such as nest building and 

territorial defense. Cowbird parasitism peaks with 

the host’s first day of egg laying. Hosts with covered 

nests, or cavities, were as vulnerable as those with 

open nests. The response to parasitism among host 

species varies (Cruz and others 1985). Host clutches 

in parasitized nests average smaller than those 

in nonparasitized nests; moreover, nest success 

at nonparasitized nests averaged higher than 

parasitized nests.

Patagieonas leucocephala (white-crowned 

pigeon)—Patagieonas leucocephala, widely 

distributed in the Caribbean, has declined throughout 

its range, including the mangroves in Boquerón 

Forest, because of habitat destruction (Wiley 1979). 

During the nonbreeding season, the birds disperse 

into hills and mountains. The unsuitability of 

some southern coastal forests may be due to the 

lack of nearby feeding habitat because previous 

foraging habitat was converted to sugar production. 

Several recommendations to increase populations 

are: maintain a closed hunting season; establish 

nesting and feeding sanctuaries in critical areas not 

currently protected; protect existing and new nesting 

and feeding habitats; establish no-hunting buffer 

zones; monitor population size, distribution, and 

breeding effort; and establish cooperative research 

and management programs with other countries. 

A compendium of information on the pigeon—

communication, breeding biology, interspecific 

interactions, food habits, and behavior—has been 

summarized in a monograph from observations 

elsewhere in Puerto Rico (Wiley and Wiley 1979). 

Seiurus novaboracensis (northern waterthrush)—

Seiurus novaboracensis was studied during four 

nonbreeding seasons in Avicennia germinans

mangrove forest (Reitsma and others 2002). The 

birds demonstrate high site fidelity within and 

between seasons. Feeding areas were small with 

considerable overlap among birds. Waterthrushes 

show high plasticity in their use of habitat during 

the nonbreeding season but may rely on mangroves 

for overwinter survival. Waterthrush densities are 

inversely associated with water level. Hurricane 
69

Georges had a major impact on mangrove habitat 

in the southwest, and forced the birds on the study 

plot to migrate to overnight roost sites in nearby 

Rhizophora mangle mangroves. 

Guánica Commonwealth Forest

Climatic Influences

Several years ago, preliminary studies using a 

linear strip census showed that species diversity, 

excluding migrant warblers and nocturnal birds, 

was greater at Guánica Forest than in the Luquillo 

Rain Forest of northeastern Puerto Rico; similarly, 

bird density counts were greater at Guánica than 

at Luquillo (Kepler and Kepler 1970). The greater

bird species diversity in dry forest communities 

as compared to wet communities was attributed to 

former climatic conditions during the Pleistocene 

when the West Indies were both larger and drier than 

today (Faaborg and Arendt 1990, 1992b). 

Monitoring of bird species at Guánica Forest 

provided information on the impacts of droughts 

on bird populations. When rainfall is insufficient at 

the end of the dry season, resident birds are unable 

to breed successfully, regardless of the quantity of 

rainfall later during the year (Faaborg and Arendt 

1992b, Faaborg and others 2000). Annual rainfall 

patterns, particularly the rainfall during the first 

6 months of each year, are related to population 

levels and survival rates (Dugger and others 2000). 

The relation between rainfall patterns and plant 

production and insect productivity are not yet known. 

In 1998, Hhurricane Georges caused considerable 

habitat damage in the Guánica Forest, including the 

traditional mist netting areas (Faaborg and others 

2000). Site damage will likely dominate other factors 

affecting bird populations for some time. Determining 

the long- term impact of the storm on populations will 

require additional monitoring. 

In general, hurricanes, which periodically traverse 

the island (fig. 8), have both direct and indirect 

impacts on bird populations (Wiley and Wunderle 

1993). Direct effects include mortality due to wind, 

rain, storm surges, and geographic displacement of 

individuals. Indirect effects include poststorm loss of 

food supplies or foraging sites, loss of nest or roosting 

sites, increased vulnerability to predation, and 

microclimatic changes. Birds respond in the shortterm through shifts in diet, foraging sites or habitats, 

and changes in reproduction. Long-term responses 

may involve greater use of secondary vegetation. 

The most vulnerable species would have a diet 

related to plant production—nectar, fruit, or seeds. 

Moreover, they would normally occupy and forage 

in closed-canopy forest. Small, local populations 

living in isolated woodland fragments would be most 

prone to extinction. Recovery of bird populations 

in posthurricane conditions would be dependent on 

the amount of damage to the vegetation as well as 

inherent species’ characteristics like the reproductive 

rate. Recruitment from undisturbed habitat in the 

vicinity would be important and highlights the need 

to protect sufficiently large areas of critical habitat to 

maintain wildlife populations in the long term. 

Post-Hurricane Georges observations of bat 

populations within the Luquillo Rain Forest showed 

that larger bat species were negatively impacted more 

than smaller ones (Jones and others 2001). Moreover, 

as with the birds, frugivorous and nectarivorous 

species were more affected than insectivorous species. 

Similar posthurricane results probably apply to bat 

species throughout the island, including the southwest.

Flora—Forest and Tree Species

Two studies dealing with recovery have been 

carried out in Guánica’s Subtropical Dry Forest. 

The first, initiated 13 years after clearcutting 1 ha of 

forest, showed that plant height reached 40 percent 

of the adjacent mature forest and that the secondary 

vegetation contained nearly 60 percent of the species 

of nearby mature forest (Dunevitz 1985). Sprouts 

and seedlings each accounted for one-half of the 

stems >1 cm. In the second, recovery was observed 

45 years after several human uses, i.e., baseball park, 

charcoal pits, farmlands, and house sites, had been 

abandoned (Molina Colón 1998). The type and rates 

of tree regeneration, plant diversity, and chemical and 

physical characteristics of the soil varied among sites. 

Two additional studies dealt with particular plant 

species. Trichilia triacantha, a small, evergreen tree 

endemic to southwestern Puerto Rico, was listed 

as endangered in 1988. A study of 109 plants in 

10 populations showed a high level of synchrony 

in flowering and fruiting, and a low production of 

fruit (Ventosa Febles 1997). Habitat loss and low 

fruit production are the principal threats to survival 
70

of the species. Given the small size and attractive 

nature of the plant, its survival could be improved if 

it were planted as an ornamental (Little and others 

1974). The second species, Tilandsia recurvata, 

is an epiphyte that grows in many areas, including 

Guánica Forest, where its abundance is correlated 

with host-tree height, basal area, and the amount of 

branching (Molina Colón 1986). Swietenia mahogani

and Pithecellobium unguis-cati are common hosts. 

The former is relatively abundant within the forest 

and the latter has large basal areas and a high degree 

of branching. The occurrence of Tilandsia recurvata

also varies with position within the trees, i.e., center 

vs. periphery and trunk vs. branches. When growing 

on Bucida buceras, greater quantities of the epiphyte 

were found on open-grown trees as opposed to those 

growing in dense stands.

Fauna—Bird Monitoring Studies

Fluctuations of populations of migratory birds 

may be due to limiting factors that occur on the 

breeding grounds, during migration, or at wintering 

sites. In 1972, annual mist netting and bird banding 

was initiated in Guánica Forest to determine species’ 

abundances and community patterns for resident and 

migratory bird populations (Faaborg and Winters 

1979, 1980; Faaborg and others 2000). Monitoring 

activities were expanded in 1989 and 1991 and have 

provided a wealth of important ecological information 

such as the impact of drought on certain species 

and their subsequent recovery, changes in species 

composition over time, and the survival and longevity 

of certain species. Some of the major findings include:

•	 Decline in migratory warblers—Long-term mist

netting of migratory warblers in Puerto Rico’s 

Guánica Forest has documented their decline from 

1973 through 1990 (Faaborg and Arendt 1992a). 

Typical explanations for declines in migrants 

include forest fragmentation and the loss of 

breeding habitat in temperate areas, and the loss 

of wintering habitat in the tropics. Guánica Forest, 

however, has not experienced human -induced 

changes in either structure or composition since 

the initiation of monitoring. These data suggest 

that explanations for declines in migrant warbler 

species based mainly on habitat destruction in 

the tropics are simplistic and may be ignoring 

important ecological patterns in the temperate zone 

(Faaborg and Arendt 1989b, 1990).

•	 Bird density declines with rainfall patterns—

Declines in the density of resident bird populations 

from 1972 to 1990 were associated with droughts 

whereas winter migrants seemed unaffected 

(Faaborg and Arendt 1989b, 1990, 1992b; Faaborg 

and others 1984). The pattern may reflect the 

fact that winter residents return during the humid 

part of the year (September and October), and, 

furthermore, do not have to feed young during 

the dry season. In contrast, the success of resident 

breeding species seems to depend on sufficient 

rainfall during May and June which normally ends 

the December through April dry season. 

•	 Foraging guilds of avian populations—Mist netting

showed that the absence of May to June rains 

most affected resident birds that breed during this 

period (Faaborg 1982, Faaborg and Arendt 1990). 

Frugivorous birds and nectarivorous birds tended 

to fluctuate more in numbers than insectivorous 

birds. It appears that insects may provide a scarce 

but predictable resource that attracts migratory 

species and, in turn, restricts the breeding season 

of resident insectivores. This restriction exposes 

breeding birds to variable conditions early in 

the wet season. The relative unpredictability of 

fruit, seed, and flower resources seem to limit 

migratory species that use these foods, and release 

resident members of these guilds from some of the 

restrictions in breeding. 

•	 Migratory land birds in the Caribbean— A total of

24 regional and local collaborators documented 

the current status of North American land-bird 

migrants with regard to species diversity, regional 

and island distributions, general abundances, 

positive and negative factors affecting populations, 

measures needed to mitigate negative impacts on 

migrant numbers, and views on the probable future 

of migrant land birds within the region (Arendt 

1992). Habitat specialist species are showing 

signs of population decline, whereas, migrant 

habitat generalists, because of their capacity to 

adapt to a variety of habitat types, including those 

altered by human activity, may not be threatened 

with extinction, or even future decreases in 

their populations. The Caribbean Islands are 

characterized by rough topography and vegetation 

types that regenerate well after disturbance. Both 

of these attributes seem favorable for the continued 

survival of migratory land birds. 
71

•	 Avian demography (longevity)— Mist netting

and bird banding during 15 years resulted in the 

capture of 39 species, 21 of which were recaptured 

(Faaborg and Arendt 1989a). Among the oldest age 

estimates for recaptured birds were 14.5 years for a 

Todus mexicanus, and 11.6 years for a Myiarchus 

antillanum and a Margarops fuscatus. These 

survival rates are among the longest recorded. The 

basic theory that describes reproductive strategy 

for bird species is that tropical birds have small 

clutches and long life spans whereas temperate 

birds have large clutches and short life spans. 

Guánica data support this generalization (Faaborg 

and Arendt 1995). The general impression is that 

resident species in southwestern Puerto Rico “are 

rather long-lived despite the seeming harshness 

of the seasonality of the dry forests in which they 

live” (Faaborg and Arendt 1989a, p. 12). 

•	 Population fluctuations, degree of philopatry,

and sex ratios— Mist netting from 1973 to 1984 

captured from 8 to 31 birds of 10 migratory 

species, ranging from 3 to 7 species in any 

particular year (Faaborg and Arendt 1984, 1990). 

Winter resident populations showed one set of 

species that appeared fully integrated into the 

Guánica Forest bird community, e.g., Mniotilta 

varia (black-and-white warbler), Setophaga 

ruticilla (American redstart), and Seiurus 

aurocapilla; with fairly stable populations and 

high site fidelity from year to year, and another set 

of different species whose populations fluctuated 

considerably. Mniotilta varia and Setophaga 

ruticilla populations showed a dominance in 

numbers of females. 

•	 Measurement of avian populations and

morphological characters— Mist nets were used to 

monitor the size, composition, and survival rates 

of bird populations in the Guánica Forest (Faaborg 

and others 2004). Point counts and mist netting 

yielded different results for species abundances. 

Mist net samples declined rapidly over a 3-day 

period due to net avoidance. Mist netting samples 

indicate avian populations that are resident in the 

vicinity; moreover, they capture most birds within 

3 days in areas with low vegetation like Guánica 

Forest. Temporal variation in the measurement 

of select avian morphological characters, i.e., 

body mass and wing length, is minimal when the 

data are recorded at the same time of the year 

(Arendt and Faaborg 1989). In addition, measuring 

instruments used in the field are precise, sampling 

techniques are comparable, and the influence of 

particular investigators is generally minimal.

•	 Overwintering warblers and additional

monitoring— Mist netting and bird banding 

between 1989 and 2003 provided data on Mniotilta 

varia, Setopahga ruticilla, and Seiurus aurocapilla

to estimate abundance and survival rates in relation 

to rainfall in both the overwintering and breeding 

areas (Dugger and others 2004). The three species 

came mostly from the Eastern United States. 

Relationships between rainfall and habitat quality, 

and winter resident demography are complex and 

involve the effects of other species. Continued 

monitoring of wintering and permanent resident 

populations will be needed to understand how 

these populations fluctuate together. 

Fauna—Individual Bird Species

Caprimulgus noctitherus (Puerto Rican nightjar)—

Previously thought extinct, the Puerto Rican nightjar 

was rediscovered in March 1961 (Reynard 1962). 

Today, the species is known from the Guánica Forest, 

the hills around the town of Guayanilla, and Susua 

Forest (Kepler and Kepler 1973, Vilella and Zwank

1987). Also, a single individual was located in the 

hills around Parguera, and seven more were found in 

the Sierra Bermeja, including the Tinaja tract (Vilella 

and Zwank 1993b). In Guánica Forest, the species 

averages about one pair on 5 ha for elevations above 

75 m. The density is highest in deciduous forest, 

evergreen forest, and plantation uplands. Originally, 

the species was probably restricted to the lower slopes 

and Coastal Plains of the island. 

Nesting occurs from mid-April to July when 

the species deposits one or two eggs in leaf litter 

(Noble 1986). Nests were situated in forested areas 

characterized by larger amounts of leaf litter biomass, 

overhanging cover, and a more open understory and 

midstory than random sites. Incubation lasts 19 days. 

About 87 percent of the nests produced at least one 

fledgling (Vilella 1995a, 1995b). Populations in 

Guánica appear to be at equilibrium and at carrying 

capacity; moreover, existing populations may be 

greater than suggested earlier because not all available 

habitat in Puerto Rico’s southwest has been surveyed 

(Noble and others 1986, Vilella and Zwank 1987). 
72

The Guaynilla Hills were selectively cut for lumber 

and charcoal during the 1880s and today contain 

some open areas grazed by goats (Vilella and Zwank 

1987). The species has been recorded on grazing lands 

where a tree canopy exists, but apparently it avoids 

riparian habitats. Conservation of existing forests and 

acquisition of private tracts would help to ensure the 

survival of the species. 

Loxigilla portoricensis (Puerto Rican bullfinch)—

Bullfinches feed alone, in pairs, or in inter- and 

intraspecific groups up to five individuals (PérezRivera 1994). The species has an ample diet, using a 

wide variety of plants along with insects and snails. 

Although the bullfinch forages from the ground 

level into the canopy, most feeding in the Guánica 

Forest occurred in the upper stratum where it favored 

the edge of the canopy. Fruit gleaning, and insect 

searching and gleaning accounted for nearly 90 

percent of its foraging. 

Tachornis phoenicobia (Antillean palm swift)—

Tachornis phoenicobia, a previously unrecorded 

vagrant was observed in Guánica Forest, apparently 

attracted by swallows and/or insects on which they 

were feeding (Kepler 1971, 1972). They were seen in

July, the putative breeding period within their ranges. 

Vireo latimeri (Puerto Rican vireo)—Vireo latimeri

is an endemic resident in Puerto Rico that does 

not routinely disperse over long distances, at least 

within Guánica forest (Woodworth and others 1998). 

Vireo breeding season length, apparently affected 

by rainfall, varies by year (Woodworth 1997). Most 

young vireos spend a prolonged period in their natal 

territory, and later settle within one to three territory 

widths of their natal nest, factors that may increase 

their survival. Vireos are declining in Guánica Forest 

because of nest parasitism by Molothrus bonariensis

and predation by rats, feral cats, and mongooses 

(Faaborg and others 1997; Woodworth and others 

1998, 1999). Molothrus bonariensis parasitizes 

about 75 percent of the vireo nests, reducing fledging 

by more than 80 percent (Woodworth 1997). The 

combination of restricted breeding season, high 

predation and parasitism rates, and low seasonal 

fecundity suggest that the species is in danger of 

extirpation in portions of its range. Despite the highly 

seasonal environment and the stress of renesting as 

many as six times per season, vireos have relatively 

high survival rates. A longevity record for the vireo 

of 13 years and 2 months was estimated by use of 

a model (Woodworth and others 1999). Another 

model that considered the implications of parasitism 

and nest predation was used to evaluate alternative 

options for management of the species (Woodworth 

1999). Recovery efforts should begin before species’ 

numbers decline further (Faaborg and others 1997). 

Zenaida aurita (Zenaida dove)—A 10-year study 

provided information on the behavior, breeding, diet, 

and habitat of the dove and showed that the dove 

occupies a variety of habitats including forest edges, 

mangrove, dry scrub, mixed agriculture, and urban 

areas (Wiley 1991). Nests, found in all months of 

the year, were generally placed in trees but were also 

built on the ground when predators were absent. Point 

counts and nest counts carried out in Guánica Forest 

between 1986 and 1996 showed that the density 

of active nests was higher in areas dominated by 

semideciduous vegetation than areas of deciduous 

vegetation or thorn scrub (Rivera-Milan 1997). 

Population declines during the measurement period 

were correlated with lower 6-month rainfalls at the 

beginning of the year. Moreover, the 6-month rainfall 

accounted for about 70 percent of the variability 

associated with nest density estimates during the 

reproductive peak in the Guánica Forest, but only 9 

percent of the variability in patches of moist forest 

elsewhere (Rivera-Milán 1996). In the mid-70s, 

the dove was the principal bird species sought by 

the island’s 7,000 hunters (Maldonado Colón and 

Pérez Rivera 1977). Guánica is a key habitat for 

reproduction of Zenaida doves in the dry zone, and 

conservation of dry forest in the southwest is critical 

for maintaining dove population levels.

Fauna—Lizards and Geckos

Thirteen species of lizards are represented in the 

dry forests of southwestern Puerto Rico; moreover, 

Guánica Forest, with 10 species, has the richest dry 

forest lizard community on the island (Genet and 

others 2001) (table 5). The species Anolis cristatellus

and Sphaerodactylus nicholsi are widespread and 

abundant in all forest types at Guánica while Anolis 

cooki is restricted to coastal scrub. All three species 

were also recorded on F&WS refuges (table 5). Anolis 

cristatellus and Anolis cooki are morphologically 

and ecologically similar species, and the former is 

dominant where both species coexist (Genet 2002). 
73

The fragmented landscape and patchy distribution 

of Anolis cooki suggest that the range of the species 

is shrinking and that it deserves immediate 

conservation attention.

Future Activities

The refuge properties, which today total about 6 

km2, should remain a nesting and feeding sanctuary 

for avifauna (Wiley 1979). Recognizing that future 

development of Puerto Rico’s southwest may include 

hotel complexes and tourist attractions, the F&WS 

should implement an aggressive management program 

with local Commonwealth and Federal Agencies, 

nongovernmental organizations, and universities. 

Research should be aimed at improving management. 

Moreover, public support should be sought through 

educational programs that highlight the area’s wildlife 

as part of a memorable outdoor experience. 

Reforestation and secondary forest recovery on 

abandoned farmlands is the critical need to enhance 

wildlife habitat and help prevent wildfires. Scattered 

trees in savannas and grasslands and remnant 

trees surviving after forest clearing function in a 

role similar to gaps in forested areas (Belsky and 

Canham 1994, Guevara and others 1986). Birds 

and bats frequently transport fruits and seeds to 

isolated trees where they perch to consume them. 

Later, the fruits or seeds are regurgitated or dropped 

to the soil. Microclimate and soil conditions under 

and immediately surrounding scattered trees are 

more favorable for regeneration than in open fields 

dominated by grass cover. 

Reforestation in abandoned fields should use 

tree species in designs that facilitate rapid recovery. 

Simultaneously, if carefully selected, the trees will 

provide nesting and foraging habitat for wildlife 

species. Exotic vs. native species will always be an 

issue in reforestation efforts. The main objective, 

initially, should be to develop forest structure, 

providing habitat for native bird species and reducing 

the threat of fire. Native tree species should 

be favored. 

Several existing research papers have cited 

plant species (including trees) that could be used 

in reforestation efforts and simultaneously serve 

particular bird species: 

•	 Agelaius xanthomus—24 tree species mentioned 

as habitat and 26 tree species for daily activities 

(Cruz-Burgos 1999)

•	 Columba leucocephala—at least 19 tree species 

used for food and/or nesting (Wiley 1979, Wiley 

and Wiley 1979)

•	 Falco sparvarius—nests in tree cavities of Cocos 

nucifera, Machaerium lunatum, Mangifera indica, 

Persea americana, Roystonea borinquena, and 

cavities in cactus (Pérez Rivera and Cotte 

Santana 1977)

•	 Loxigilla portoricensis—uses parts of 41 species 

of plants as food sources and also forages on 

animal matter in the upper stratum in Guánica dry 

forest (Pérez-Rivera 1994)

•	 Geotrygon montana (ruddy quail dove)—uses 

14 plant species, including fruit trees, along with 

several grass species as food (Pérez-Rivera 1979)

•	 Patagioenas leucocephala—uses 10 tree species as 

food sources or nesting sites (Wiley 1979)

•	 Zenaida aurita—one study showed that the dove 

uses at least 31 plants as food and at least 35 tree 

species for nesting (Maldonado Colón and Perez 

Rivera 1977) and another study indicated that the 

dove uses 77 plant species for various purposes 

(Wiley 1991) 

•	 Sixteen bird species use the Roystonea borinquena

for nesting sites (Pérez Rivera 1977). Numerous 

bird species use 120 listed plant species as food 

(Soil Conservation Service 1973, see endnotes)..

•	 Birds and mammals—50 species of plants

[bromeliads, palms, grasses, and dicots (including 

mangroves)] used as sources of water and food 

in the nearby U.S. Virgin Islands (Department of 

Planning and Natural Resources 1988)

Future F&WS management activities should 

include: 

•	 Expansion of refuge properties in southwestern

Puerto Rico to form a network of critical habitats 

for wildlife conservation. Human population 

will continue to increase (fig. 2) and will be 

accompanied by conversion of wildlife habitat. 

An appropriate agency response would be the 

continual acquisition of all available land when 

feasible, including additional tracts in Sierra 

Bermeja (Aukema and others 2006). 
74

•	 Continuation of reforestation efforts as a means

of stimulating habitat restoration and forest 

regeneration (Parrotta and others 1997) 

•	 Management of the Cartagena Lagoon to enhance

populations of resident and migratory birds, 

including the target bird species which have 

experienced declines (Díaz-Soltero 1990)

•	 Management of the Candeleria and Fraternidad

Lagoons in the Salinas flats for populations of 

shore birds through the increase of edge habitat 

(construction of mounds or small islands) and 

possibly through modification of the water 

level regimen 

•	 Peer review of major F&WS management

proposals and plans among local and Federal 

Agencies, particularly when they may lead to 

major habitat modification

•	 Development and maintenance of a small

Caribbean reference library of relevant local 

literature and refuge studies to facilitate future 

management and research activities 

•	 Fire control through adequate training,

construction and maintenance of fire breaks, 

prescribed burning, surveillance, public education, 

and community involvement 

•	 Cooperative efforts with the municipal

governments and hotels to provide educational 

opportunities for residents and visitors to 

southwestern Puerto Rico 

•	 Eradication of alien species, e.g., monkeys, and

removal of unauthorized livestock that negatively 

impact native wildlife or their habitat

•	 Limitation of road construction and other habitat

fragmentation to reduce the invasion of exotic flora 

and fauna into areas of critical wildlife habitat 

(Clark 1993, Faaborg and Arendt 1990, Vilella and 

Swank 1993a) 

•	 Presentation of research and management

activities on a periodic basis, i.e., symposia among 

cooperators and to the public in nearby population 

centers like Boquerón, Lajas, and Magüayo 

•	 Monitoring compliance of refuge regulations,

including hunting and special use permits, i.e., 

commercial salt extraction and grass harvest 

•	 International cooperation among conservationists,

resource managers, policymakers, and the public 

to promote programs beneficial to migratory land 

birds (Arendt 1992)

Future research activities should include all 

levels of biological diversity—genetics, species, 

communities, and ecosystems (Clark 1993)—

involving most of the following: 

•	 Partnerships with domestic scientists at

government agencies and universities to carry 

out research on cultural, scientific, and historic 

resources for refuges and surrounding lands 

(Meretsky and others 2006)

•	 Compilation and publication of data from

monitoring projects, i.e., bird surveys, nesting 

activities, bird banding, and water-quality 

monitoring at Cartagena Lagoon; and identification 

of future data needs and studies

•	 Investigation of the ecology, behavior, and habitat

requirements of native wildlife species (DíazSoltero 1990)

•	 Stimulation of forest succession on the barren

upper slopes of Cerro Mariquita through 

innovative techniques, i.e., use of artificial bird 

perches (McClanahan and Wolfe 1993)

•	 Recurrent plant inventories, including the

permanent Tinaja plots and vegetation on other 

tracts, to determine changes in structure and 

species composition over time

•	 Monitoring of tree plantings to determine their

survival, growth, and future potential for habitat 

restoration, and verification of associated changes 

in habitat utilization by wildlife (Weaver and 

Schwagerl 2004, 2005)

•	 Determination of flowering and fruiting patterns by

plant species (Aukema and others 2006)

•	 Monitoring avifauna feeding, nesting, and

migratory habits (Grear 1992a, 1992b)

•	 Monitoring long-term population dynamics to

determine temporal, e.g., population fluctuations 

over time, and spatial changes, e.g., differential use 

of habitats, for both migrant and resident species 

(Arendt 1992)
75

•	 Monitoring biological diversity of animal

species other than birds, e.g., mammals, reptiles, 

amphibians, fishes, and insects; to determine their 

critical habitats, population numbers, roles in 

refuge ecosystems, and dynamics over time, i.e., in 

reponse to vegetation changes, introduced species, 

and climatic events 

•	 Propagation of rare and endangered species for

use in planting trials or as ornamentals within the 

region to enhance their survival

•	 Developing a brochure on major refuge insects,

including butterflies (Ramos 1982)

•	 Monitoring the salt operations at Cabo Rojo to

determine their impact on wildlife species, and 

simultaneously assessing the possible impacts of 

alternative management strategies for the 

salt ponds 

•	 Excavating and interpreting shell mounds for

refuge visitors

Acknowledgments

Field help was received from Osvaldo Quiñones 

Martínez, Yesmenia Mercado, Jennifer Ramiréz 

Jourdan, and Alexandra Torres Ruíz. Information was 

provided by Myrna Aponte Reyes of the Department 

of Natural and Environmental Resources, and Jorge 

Baez, Fernando Silva, and Ana I. Torres of the 

Puerto Rican Conservation Trust regarding protected 

reserves; Manual J. Cruz Calderón, Veronica MendezGallardo, and Adrianne Tossas of the Puerto Rican 

Ornithological Society on Christmas bird counts; 

and Wayne Arendt and Miguel Canals Mora on bird 

species for Guánica Forest. Evelyn Cepeda helped 

collect information, including several theses cited in 

this report. William Hernandez provided information 

on soils and geology. Maya Quiñones of the 

Geographic Information System and Remote Sensing 

Laboratory of the International Institute of Tropical 

Forestry helped develop the maps. Wayne Arendt 

verified the bird list, adding to the species’ lists. Frank 

H. Wadsworth reviewed the manuscript and offered 

helpful suggestions. The research was made possible 

through a “Partners for Wildlife” grant from the U.S. 

Fish and Wildlife Service. The work was carried out 

in cooperation with the University of Puerto Rico at 

both the Río Piedras and Mayagüez campuses. 

Endnotes

Anon. [No date]. Plants of Sandy Point, St. Croix. 4 p. Unpublished 

field list on file with: U.S. Fish and Wildlife Service, Carretera 301, 

Km 5.1, P.O. Box 491, Boquerón, PR 00622.

Anon. 1987. Birds seen on Desecheo Island, Puerto Rico during field 

trips. 2p. Unpublished field list on file with: U.S. Fish and Wildlife 

Service, Carretera 301, Km 5.1, P.O. Box 491, Boquerón, PR 00622.

Asencio López, Carmen I.; Rentas Soto, Yolanda V. 2006. Mariposas 

identificadas en el Centro Interpretativo de las Salinas en Cabo Rojo 

“Efrén Pérez Rivera.” 1 p. Unpublished list. In Spanish. On file with: 

U.S. Fish and Wildlife Service, Carretera 301, Km 5.1, P.O. Box 491,

Boquerón, PR 00622.

Bailey, David C. 1990. Foraging water depth related to individual bill 

lengths of three Calidris sandpiper species occurring at a migratory 

stopover. Manomet, MA: Manomet Bird Observatory. 19 p. + figures 

and tables. Unpublished student study. On file with: U.S. Fish and 

Wildlife Service, Carretera 301, Km 5.1, P.O. Box 491, Boquerón,

PR 00622.

Belitsky, D.W. 1978. Islandwide waterfowl investigations, final 

report. San Juan, PR: Department of Natural Resources. 80 p. 

Unpublished report. On file with: Estado Libre Asociado de Puerto 

Rico, Departamento de Recursos Naturales y Ambientales, P.O. Box 

9066600, San Juan, PR 00906–6600.

Bonilla, G. 1992. Puerto Rico waterfowl studies, annual performance 

report. San Juan, PR: Department of Natural resources. 45 p. 

Unpublished report. On file with: Estado Libre Asociado de Puerto 

Rico, Departamento de Recursos Naturales y Ambientales, P.O. Box 

9066600, San Juan, PR 00906–6600.

Chabert, J.L. 1984. Informe de estatus de las aves acuáticas de caza y 

sus habitats. San Juan, PR: Departamento de Recursos Naturales. 46 p. 

Unpublished report. In Spanish. On file with: Estado Libre Asociado de 

Puerto Rico, Departamento de Recursos Naturales y Ambientales, P.O. 

Box 9066600, San Juan, PR 00906–6600.

Collazo, Jaime A.; Thomas, Johanna; Knott, Anne; Colón, José A. 

1987. Preliminary findings of several aspects of the ecology of Snowy 

and Wilson’s plovers at the Cabo Rojo Salt Flats. 17 p. Unpublished 

interim report. On file with: U.S. Fish and Wildlife Service, Carretera 

301, Km 5.1, P.O. Box 491, Boquerón, PR 00622.

Departamento de Agricultura. 1995. Contrato de arrendameinto de 

facilidades pesqueras. 4 p. Contract. In Spanish. On file with: U.S. Fish 

and Wildlife Service, Carretera 301, Km 5.1, P.O. Box 491, Boquerón,

PR 00622.

Díaz Díaz, Carlos A. 1984. Recovery plan for the Puerto Rican 

whip-poor-will (Caprimulgus noctitherus). Atlanta, GA: U.S. Fish 

and wildlife Service. 16 p. Unpublished interim report on file with: 

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Appendix A

Alphabetical listing by scientific name of all tree species that were mentioned in the text or tables, and of other plant species mentioned only in text. Ecological information is provided only for tree species a

Common names Ecological information

Scientific name English Spanish Flower b Fruit b Habit c Origin d

months

Tree species (text and other tables)Acacia farnesiana (L.) Willd. Sweet acacia Aroma NO-FE ? D N?

Acacia tortuosa (L.) Willd. Twisted acacia Casia YR YR? D N

Acrococomia media O.F. Cook Prickly palm Corozo YR YR E N

Adelia rincilla L. NA Cotorro FA-SP WI-SU D N

Albizia lebbeck (L.) Benth Tibet Acacia amarilla AP-SE YR D I

Amyris elemifera L. Sea amyris Tea MA-OC YR E N

Andira inermis (W. Wright) DC. Cabbage angelin Moca JA-FE+MA-SE MA-DE D N

Annona glabra L. Pond Apple Corozón SP SU D N

Annona muricata L. Soursop Guanábana JU-OC FA E N

Annona squamulosa L. Sugar Apple Anón YR YR D N

Avicennia germinans (L.) L. Black mangrove Mangle prieto YR YR E N

Bauhinia monandra Kurz Butterfly bauhinia Mariposa YR YR E I

Bourreria succulenta Jacq. Pigeon berry Palo de vaca YR YR E N

Bucida buceras L. Oxhorn bucida Ucar YR YR E N

Bunchosia glandulosa (Cav.) L.C. Rich. NA Café forestero MA-OC JU-AU E/D N

Bursera simaruba (L.) Sarg. Terpentine tree Almácigo SP-SU SP-SU D N

Byrsonima lucida (Mill.) L.C. Rich. Long Key byrsonima Palo de doncella YR YR E N

Byrsonima spicata (Cav.) HBK. NA Maricao YR YR E N

Calophyllum calaba L. Santa-maria María SP-SU SP-WI E N

Calotropis procera (Aiton) W.T. Aiton Giant milkweed Algodón de seda YR YR E I

Canella winterana (L.) Gaertner Canella Barbasco YR YR E N

Capparis amplissima Lam. NA Burro blanco YR YR E N

Capparis cyanophallophora L. Jamaica caper Burro prieto YR YR E N

Capparis flexuosa (L.) L. Limber caper Palinguan YR YR E N

Capparis hastata Jacq. NA Burro YR YR E N

Capparis indica (L.) Fawe & Rendle Linguam Sapo prieto YR YR E N

Carica papaya L. Papaya Lechosa YR YR E I

Casearia aculata Jacquin NA Cambrón YR YR D N

Casearia guianensis (Aublet) Urban Wild coffee Palo blanco YR YR E N

Cassine xylocarpa Vent. Marble-tree Coscorrón YR YR E N

Cedrela odorata L. Spanish cedar Cedro hembra YR YR D N

Ceiba pentandra (L.) Gaertner Silk-cotton-tree Ceiba DE-FE SP-SU D N

Chamaesyce articulata (Aublet) Britton NA NA YR YR D N

continued
90

Appendix A (continued)

Alphabetical listing by scientific name of all tree species that were mentioned in the text or tables, and of other plant species mentioned only in text. Ecological information is provided only for tree species a

Common names Ecological information

Scientific name English Spanish Flower b Fruit b Habit c Origin d

months

Chryobalanus icaco L. Coco-plum Hicaco YR YR E N

Citharexylum fruiticosum L. Fiddlewood Péndula YR YR E N

Citrus aurantifolia (Christm.) Swingle Lime Limón ágrio SP-JL SU-FA E I

Citrus limon (L.) Burm. f. Lemon Limón de cabro SP SU-FA E I

Clerodendron aculeatum (L.) Schlecht Haggarbush Escambrón YR YR ? N

Clusia rosea Jacquin Wild-mammee Cupey YR YR E N

Cnidoscolus aconitifolius (Mill.) I.M. Johnst. NA Papayuelo YR O E I

Coccoloba diversifolia Jacq. Doveplum Uvilla YR YR E N

Coccoloba microstachya Willd. NA Uverillo SU-FA WI E/D N

Coccoloba swartzii Meissn. NA Ortegón JU-SE JU-SE E NE

Coccoloba uvifera (L.) L. Sea grape Uva de playa YR YR E N

Coccoloba venosa L. Chicory-grape Calabreña MA-SE OC D N

Cocos nucifera L. Coconut Palma de coco YR YR E I

Colubrina arborescens (Mill.) Sarg. Coffee colubrina Abeyuelo SP-FA SP-FA E N

Colubrina elliptica (Sw.) Briz. & Stern Soldierwood Mabí JL-NO JL-JA E N

Comocladia dodoaea (L.) Urban Pioson ash Carrasco WI-SP SP-SU E N

Conocarpus erectus L. Button mangrove Mangle botón YR YR E N

Cordia alliodora (Ruiz & Pav.) Oken Onion cordia Capa prieto YR YR E N

Cordia collococca L. Red manjack Cerezo SP-SU SP-SU D N

Cordia rickseckeri Millsp. Manjack San Bartolome YR YR D? N

Cordia sulcata DC. White manjeck Moral SP-FA YR D N

Crecentia portoricensis Britton NA NA ? ? E NE

Crescentia cujete L. Calabash tree Higüero YR YR E/D N

Crescentia linearifolia Miers. NA Higüerito YR YR D N

Crossopetallum rhacoma Crantz Florida crossopetalum Coral YR YR E/D N

Croton astroites Dryander NA Marán YR YR E N

Delonix regia (Bojer) Raf. Flamboyant tree Flamboyán MA-JL YR D I

Erithalis fruticosa L. Black torch Jayajabico YR YR E N

Erythroxylum aerolatum L. Thin-leaf erythroxylum Indio OC-JU ? D N

Erythroxylum rotundifolium Lunan Brisselet Rocio SU-FA SU D N

Eugenia biflora (L.) DC. NA Hoja menuda SP-SU SP-SU E N

Eugenia ligustrina (Sw.) Willd. NA Palo de muleta MA ? E N

Eugenia monticola (Sw.) DC. Black-cherry Biriji SP-FA SP-FA E N

Eugenia pseudosidium Jacq. Wild guava Quiebrahacha YR? YR? E N

Eugenia sessiliflora Vahl NA NA SP-SU SP-SU E N

continued
91

Appendix A (continued)

Alphabetical listing by scientific name of all tree species that were mentioned in the text or tables, and of other plant species mentioned only in text. Ecological information is provided only for tree species a

Common names Ecological information

Scientific name English Spanish Flower b Fruit b Habit c Origin d

months

Eugenia woodburyana Alain NA Hoja menuda ? ? E NEEugenoa procera (Sw.) Poir. NA Hoja menuda SP-FA SP-FA E NExostema caribaeum (Jacq.) R. & S. Caribbean princewood Albarillo YR YR E NFicus benjamina L. Benjamin fig Laurel de Benjamina ? SP-FA E IFicus citrifolia Mill. Shortleaf fig Jagüey blanco YR? YR E NGaussia attentuata O.F. Cook Beccari Puerto Rico llume-palm Palma de lluvia YR YR E NEGliricidia sepium (Jacq.) Kunth ex Walp. Mother-of-cacao Mata-ratón WI-SP WI-SU D IGuaiacum officinale L. Lignum-vitae Guayacán SP-FA SP-FA E NGuaiacum sanctum L. Holywood lignum-vitae Guayacán blanco SP-FA SP-FA E NGuapira discolor (Spreng.) Little Barrehorno Barrehorno SP-SU SU-FA E NGuapira fragrans (Dum.-Cours) Little Black mampoo Corcho SP-SU SP-SU E NGuazuma ulmifolia Lam. West Indian elm Guácima SP-FA YR E/D NGuettarda elliptica Sw. Velvetseed Cucubano liso SP SU E NGuettarda scabra (L.) Vent. Greenhart Palo de cucubano YR YR E NGymnanthes lucida Sw. Oysterwood Yaití SP-FA SP-FA E NHaematoxylum campechianum L. Logwood Campeche DE-MA ? D IHelicteres jamaicensis Jacq. Cowbush Cuernecillo YR YR E NHippomane mancinella L. Manchineel Manzanillo SP-OC YR E NHymenaea courbaril L. West Indian locust Algarrobo SP-FA ? E NInga laurina (Sw.) Willd. Sweetpea Guamá YR YR E NInga vera Willd. NA Guaba YR YR E NJaquinia arborea Vahl Torchwood Barbasco WI-SU SP-FA E NKrugiodendron ferreum (Vahl) Urban Ironwood Bariaco YR YR E NLaguncularia racemosa (L.) Gaertn. White mangrove Mangle blanco YR YR E NLantana camera L. var. camara Lantana Lantana YR YR E NLeucaena leucocephala (Lam.) de Wit Tantan Zarcilla YR YR D IMachaerium lunatum (L. f.) Ducke NA Escambrón YR YR E NMachaonia portoricensis Baill NA Alfilerillo YR YR D NEMangifera indica L. Mango Mango WI-SP MA-SE E IManilkara bidentata (A. DC.) Chev. Balata Ausubo YR YR E NManilkara pleeana (Pierre) Cronq. NA Zapote de costa YR YR E NMelia azedarach L. Chinaberry Alelaila YR YR D IMelicoccus bijugatus Jacq. Kinep Quenepa AP-JU JU-SE E IMetopium toxiferum (L.) Krug & Urban Florida poisontree Papayo FE SU-FA E NMyrcia citrifolia (Aubelet) Urban NA Limoncillo del monte SP-FA SU-WI E N

continued
92

Appendix A (continued)

Alphabetical listing by scientific name of all tree species that were mentioned in the text or tables, and of other plant species mentioned only in text. Ecological information is provided only for tree species a

Common names Ecological information

Scientific name English Spanish Flower b Fruit b Habit c Origin d

months

Neea buxifolia (Hook. f.) Heimerl NA NA SP-SU SU E NOttoschulzia rhodoxylon (Urban) Urban NA NA ? ? E NEOuratea littoralis Urban NA Abey amarillo YR YR E NOxandra lanceolata (Sw.) Baill. Lancewood Haya prieto ? ? ? NParkinsonia aculeata L. Jerusalem-thorn Palo de rayo YR YR D NPectitia domingensis Jacq. Fiddlewood Capa blanco YR YR E NPersea americana Mill. Avocado Aguacate JA-AP AP-MA D IPictetia aculeata (Vahl.) Urban Fustic Tachuelo YR YR D NPilosocereus royenii (L.) Byles & Rowley Pipe-organ cactus Sebucán YR ? E NPimenta racemosa (Mill.) J.W. Moore Bay-rum-tree Malagueta AP-AU AU-OC E NPiscidia carthagenesis Jacq. Dogwood Ventura FE-JU SU D NPisonia albida (Heimer) Britton & Standley Corcho bobo Corcho bobo FE-MA SP (late) D NPisonia subcordata Sw. Water mampoo Corcho blanco SP-SU SP D NPithcellobium dulce (Roxb.) Benth Ape’s earring Guamá americano JA-MA FE-JL E IPithcellobium unguis-cati (L.) Mart. Catclaw Una de gato YR YR D NPlumeria alba L. Milktree Alelí YR YR E NPolygala cowellii (Britton) Blake Violet-tree Violeta WI MA-AP D NEPolygala penaea L. NA NA YR YR E NProckia crusis R. Brown ex L. NA NA ? SU E/D NProsopis juliflora (Sw.) DC. Mesquite Bayahonda SU-FA SU-FA D IPterocarpis officinalis Jacq. Swamp bloodwood Palo de pollo FE-SE AP-NO E NRandia aculeata L. Box briar Tintillo YR YR D NRauvolfia nitida Jacq. Bitter ash Palo amargo YR YR E NReynosia uncinata Urban NA Cascarolla YR YR? E NRhizophora mangle L. Red mangrove Mangle rojo YR YR E NRicinus communis L. Castorbean Higüerito YR YR E IRochefortia acanthophora (DC.) Griseb. Juso Juso SP-SU SP-SU D NRondeletia inermis (Spreng) Krug & Urban NA Cardobancillo YR YR E NERoystonea borinquena O.F. Cook Royal palm Palma royal YR? YR? E NSabal casuarium (O.F. Cook) Beccari Puerto Rico palmetto Palma de sombrero YR YR E NESamanea saman (Willd.) Merril Raintree Samán SP-FA FA-WI E ISapindus saponaria L. Wingleaf soapberry Jaboncillo ? ? E NSavia sessiliflora (Sw.) Willd. NA Amansa guapa YR YR D NSchaefferia frutescens Jacq. Florida boxwood Jiba YR YR E NSecuringa acidoton (L.) Rawcett & Randle NA NA SP-SU SP-SU D N

continued
93

Appendix A (continued)

Alphabetical listing by scientific name of all tree species that were mentioned in the text or tables, and of other plant species mentioned only in text. Ecological information is provided only for tree species a

Common names Ecological information

Scientific name English Spanish Flower b Fruit b Habit c Origin d

months

Senna polyphylla (Jacq.) Irwin & Barneby Desert cassia Ratama prieta YR YR D N 

Sideroxylon foetidissum Jacq. False mastic Tortugo amarillo YR YR E N

Sideroxylum obovatum Lam. Breakbill Lechecillo YR? YR? E N

Solanum drymophilum O.E. Schultz NA Erubia YR YR ? NE

Spondias mombin L. Hogplum Jobo WI-SU SU-WI D N

Stahlia monosperma (Tul.) Urban NA Cobana negra FE-JU SU-FA E N

Stenocereus hystrix (Mill.) R. Kiesling Organpipe cactus Pitahaya SP-SU SU E N

Sterculia apetala (Jacq.) Karst Panama tree Anacaguita SP-FA ? E I

Swietenia mahagoni Jacq. Small leaf mahogany Caoba dominicana MA-JL WI D I

Tabebuia heterophylla (DC.) Britton White-cedar Roble blanco SP+YR YR D N

Tamarindus indica L. Tamarind Tamarindo SP-FA WI-SP E I

Tecoma stans (L.) Juss. Ginger-thomas Roble amarillo YR YR E N

Tectona grandis L. f. Teak Teca AU-DE YR D I

Terminalia catappa L. Indian almond Almendro YR YR E/D I

Thespesia populnea (L.) Solander Seaside mahoe Emajagüilla SP-FA SP-FA E I

Thouinia striata Radlk. NA Ceboruquillo SP-FA SP-FA E NE

Thouinia striata var. portoricensis Serrasuela Serrasuela YR YR D NE

(Radlk) Voltava & Alain

Trichilia hirta L. Broomstick Tinacio YR YR D N

Trichilia triacantha Urban NA Bariaco FE ? E NE

Vitex divaricata Swartz White fiddlewood Higüerillo FE-JL JU-NO D N

Zanthoxylum flavum Vahl Yellow sanders Aceitillo WI-SU SP-FA D N

Zanthoxylum martinicense (Lam.) DC. White prickle Espino rubial SP-FA SP-FA E N

Zanthoxylum monophyllum (Lam.) P. Wilson Yellow prickle Palo rubio ? ? D N

Ziziphus reticulata (Vahl) DC. Cascarroya Cascarroya SU SU E N

Other Plant Species (text only)Aristida chaseae A.S. Hitchc. NA (grass) NA (graminea)

Aristida portoricensis Pilger NA (grass) NA (graminea)

Batis maritima L. Saltwort, pickleweedBromelia penguin L. Penguin bromelia Piña de ratónBulbostylis pauciflora (Liebm.) C.B. Clarke Fewflower hairsedgeCatesbaea melanocarpa Krug & Urban Tropical lilythornCenchrus ciliaris L. African foxtail

continued
94

Appendix A (continued)

Alphabetical listing by scientific name of all tree species that were mentioned in the text or tables, and of other plant species mentioned only in text. Ecological information is provided only for tree species a

Common names Ecological information

Scientific name English Spanish Flower b Fruit b Habit c Origin d

months

Cynometra portoricensis Krug & Urban False oregano Oreganillo falso

Dendrophthora bermejae Kuijt, Carlo & Aukema Mistletoe Muérdago

Heliotropium curassavicum L. Seaside heliotropeHohenbergia antillana Mez. Antilles lacebarkLeptocereus quadricostatus Bello (Britt & Rose) PitahayaLyonia truncata Urban var. proctorii Proctor’s staggerbushPanicum maximum Jacq. P.Q. Guinea grassPolygala hecatantha Urban West Indian milkwortPsychilis krugii (Bello) Sauleda Krug’s peacock orchid NE

Sesuvium portulacastrum L. Shoreline purslaneTephrosia cinerea (L.) Pers. Asshen hoarypeaTillandsia recurvata (L.) L. Small ballmossVeronia proctorii L.E. Urbatsch NA NA

Waltheria calcicola Urban RaichieWedelia lanceolata DC. Riverbank creeping oxeye N 

Zamia portoricensis Urban ? Marungüey ? ? E NE

a Scientific names for plants were used in text to avoid confusion. Many species have several common names in both languages and some species have no common name. b Months in flower or fruit, as follows: JA, FE, MA, AP, MM, JU, JL, AU, SE, OC, NO, DE; YR = entire year, or intermittently during the year; seasonal (i.e., SP = spring, SU = 

summer, FA = fall, WI = winter); ? = uncertain or data not provided; O = not fruit bearing in Puerto Rico. Sources; Little and Wadsworth 1964; Little and others 1974.

c Habit: D = deciduous; E = evergreen; E/D = evergreen, except in pronounced drought.d Origin: N = native; NE = endemic to Puerto Rico; I = introduced; ? = uncertain.
95

Appendix B

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Pteridophyta

Polypodiaceae

Azolla caroliniana Wild. – – X –

Acrostichum danaeifolium Langsd. & Fisher – – – X

Cheilanthes microphylla (Sw.) Sw. – – – X

C. trichomanolides (L.) Mett. – – – X

Marsilea ancylopoda A. Braun – – X –

Memionitis palmata L. – – – X

Nephrolepis biserrata (Sw.) Schott – – X –

N. multiflora (Roxb.) Jarret ex Morton – – – X

Polypodium heterophyllum L. – – – X

P. polypodioides (L.) Watt – – – X

Gynospermae (Cycadophyta)

Zamiaceae

Zamia portoricensis Urban – – – X

Monocotyledonae

Alismataceae

Echinodorus berteroi (Spreng.) Fassett X – X –

Sagittaria intermedia M. Micheli – – X –

S. lancifolia L. X – X –

Araceae

Anthurium crenatum (L.) Kunth – – – X

Philodendron scandens Koch & Sello – – X –

Pistia stratiotes L. – – X –

Xanthosoma brasiliense (Desf.) Engl. – – – X

Arecaceae

Acrocomia media O.F. Cook X – X –

Cocos nucifera L. X X X –

Roystonea borinquena O.F. Cook X – X –

Sabel morrissi H. Wendl. X – – –

Thrinax morrissi H. Wendl. – – – X

Commelinaceae

Commelina diffusa Burm. f – – X –

C. elegans HBK. – – X X

Cyperaceae

Bulbostylis capillaris (L.) Kunth ssp. antillana (Britton) T. Koyama X – – X

Cyperus articulatus L. – – X –

C. compressus L. – – X –

C. esculentus L. – – X –

C. giganteus Vahl – – X –

C. laevigatus L. – – X –

C. lingularis L. – – X –

C. ochraceus Vahl X – X –

C. odoratus (Presl.) Mattf. & Kuk – – X –

C. rotundus L. X – X X

C. surinamensis Rottb. X – X –

Eleocharis cellulosa Torrey – – X –

continued
96

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Cyperaceae (continued)

E. fallax Weath. – – X –

E. geniculata (L.) R. & S. – X X –

E. interstincta (Vahl) R. & S. – – X –

E. mutata (L.) R. & S. – – X –

Kyllinga brevifolia Rottb. – – – X

Pycreus poystachyos (Rottb.) P. Beauv. – – – X

Rhynchosphora nervosa (Vahl) Boeck.

ssp. ciliata (Vahl) T. Koyama – – – X

R. holochoenoides (L.C. Rich.) Herter – – – X

Scleria lithosperma (L.) Sw. – – – X

S. pterota Presl. – – – X

Graminae (or Poaceae)

Anthephora hermaprodita (L.) Kuntze X – – –

Aristida adscensionis L. X – – –

A. chaseae Hitchc. X – – –

A.portoricensis Pilger – – – X

Bambusa vulgaris Schrad. ex Wendl. – – X –

Bothriochloa pertusa (L.) A. Camus X – X X

Bouteloua americana (L.f) Scribn. X – X –

B. repens (HBK.) Scribn. & Merr. – – – X

Brachiaria adspersa (Trin.) Parodi X – – –

B. fasciculata (Sw.) S.T. Blake X – – –

B. purpurascens (Raddi) Henr. – – X –

B. subquadripara (Trin.) Hitchc. X – – X

Cenchrus ciliris L. X X X X

C. echinatus L. X X X –

Cheloris ciliata Sw. X – – –

C. inflata Link X – X –

Cynodon dactylon (L.) Pers. X – X –

Dactyloctenium aegyptium (L.) Beauv. X – X –

Dichanthium annulatum (Forssk.) Stapf. X X X X

Digitaria bicornus (Lam.) R. & S. – – X X

D. decumbens Stent X – – –

D. eggersii (Hack.) Menr. – – – X

D. insularis (L.) Mez – – X –

D. sanguianalis (L.) Scop. X – – –

Echinochloa colona (L.) Link X – X –

Eleusine indica (L.) Gaertn. – – X –

Eragrostis ciliaris (L.) R. Br. X – X –

E. tenella (L.) Beauv. ex R. & S. X – – –

Eriochloa polystachya HBK. – – X –

E. punctata (L.) Desv. – – X –

Heteropogon contortus (L.) Beauv. ex R. & S. X – – –

Hymenachne amplexicaulis (Rudge) Hees – – X –

Lasiacis divaricata (L.) Hitchc. – – – X

Leersia monandra Sw. – – – X

Leptochloa filiformis (Lam.) Beauv. X – – –

Olyra latifolia L. – – – X

Panicum maximum Jacq. X X X X

Paspalidium geminatum (Forssk.) Stapf X – X –

continued

Appendix B (continued)
97

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Graminae (or Poaceae) (continued)

Paspalum conjugatum Berg. – – X –

P. distichum L. – – X –

P. laxum Lam. X – – –

P. notatum Flugge – – X –

P. setaceum Michx. var. ciliatifolium (Michx.) Vasey X – – –

Pharus glaber HBK. – – – X

Rottboellia cochinchinensis (Lour.) Clayton – – X –

Saccharum officinarum L. – – X –

Setaria barbata (Lam.) Kunth – – X –

S. rariflora Mikan – – X –

S. setosa (Sw.) Scribn. var. leiophylla (Nees) Arechavaleta X – – –

Sporobolus indicus (L.) R. Br. – – – X

S. jacquemontii Kunth X – X – 

S. pyramidatus (Lam.) Hitchc. X – – – 

S. virginicus (L.) Kunth X – – –

Tragus berteronianus Schult. X – – –

Tricholaena repens (Willd.) Hitchc. X – – X

Urochloa maxima (Jacq.) R.D. Webster X – – –

Liliaceae

Aloe vera (L.) Burn. f. X – – –

Sansevieria trifasciata Prain X X X –

Yucca aloifolia L. X X – –

Najadaceae

Najas guadalupensis (Spreng.) Morong – – X –

Orchidaceae

Epidendrum krugii Bello – – – X

Oecceoclades maculata (Lindl.) Lindl. X – – X

Psychilis krugi (Bello) Sauleda – – – X

Tetramicra canaliculata (Aubl.) Urban – – – X

Vanilla claviculata (W. Wr.) Swartz – – – X

Pontederiaceae

Eichhornia crassipes (Mart.) Solms in DC. – – X –

Smilaceae

Smilax coriacea Spreng. – – – X

Typhaceae

Typha domingensis Pers. – – X –

Dicotyledonae

Acanthaceae

Blechum pyramidatum (Lam.) Urban – – X –

Justica comata (L.) Lam. – – X –

Ruellia tuberosa L. X X X –

Siphonoglossa sessilis (Jacq.) Gibson X – – –

Thunbergia alata Bojer ex Sims – – X –

continued

Appendix B (continued)
98

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Aizoaceae 

Mollugo verticillata L. X – – –

Sesuvium portulacastrum L. X X – –

Trianthema portulacastrum (L.) L. – – – X

Amaranthaceae

Achyanthes aspera L. X X – X

Alternanthera paronychioides St. Hil. – – X –

Amaranthus crassipes Schlecht X X – –

A. dubius Mart. X – X –

A spinosus L. – – X –

A. viridis L. – – X X

Gomphrena serrata L. X – – X

Iresine angustifolia Euphrasén – – – X

Amaryllidaceae

Hymenocallis caribaea (L.) Herb. X – X –

Anacardiaceae 

Comocladia dodonaea (L.) Urban X – – X

Mangifera indica L. X – X X

Metopium toxiferum (L.) Krug & Urban X X – –

Spondias mombin L. X – X X

Annonceae 

Annona glabra L. – – X –

A. muricata L. X – X – 

A. reticulata L. – – X –

A. squamosa L. X X X –

Oxandra lanceolata (L.) Urban – – – X

Apocynaceae

Catharanthus roseus (L.) G. Don X X – –

Pentalinon luteum (L.) Hansen & Wunderlin – X – –

Plumaria alba L. X X – X

P. obtusa L. X – – –

Rauvolfia nitida Jacq. X X X X

R. viridis Willd. Ex. Roem. & Schult. X – X –

Aquifoliaceae

Ilex sp. – – X –

Asclepidaceae

Asclepias curassavica L. – – X –

Calotropis procera (Ait.) Ait.f. X X X X

Cryptostegia grandiflora R. Brown – – X –

Matelea maritima (Jacq.) Woodson X – – –

Metastelma linearae Bello – – – X

M. fallax Schltr. X – – –

Basellaceae

Anredera leptostachys (Moq.) v. Stennis X – – X

Bataceae

Batis maritima L. X X – –

continued

Appendix B (continued)
99

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges 

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Bignoniaceae

Crescentia cujete L. X – X X

C. linearifolia Miers. X – X –

C. portoricensis Britton X – – – 

Disticus lactifolia (Vahl) DC. X – X X

Macfadyena unguis-cati (L.) A. Gentry X – X X

Spathodea campanulata Beauv. – – X –

Tabebuia donnell-smithii Rose X – – –

T. haemantha (Bert.) DC. – – – X

T. heterophylla (DC.) Britton X X X X

Tecoma stans (L.) Kunth in HBK. X – – –

Bombacaceae

Ceiba pentandra (L.) Gaertner X – X X

Boraginaceae

Bourreria succulenta Jacq. var. succulenta X X X X

B. virgata (Sw.) G. Don – – – X

Cordia alliodora (R. & P.) Oken – – X X

C. collococca L. X – X X

C. curassavica (Jacq.) R. – – – X

C. globosa (Jacq.) Kunth var. humilis (Jacq.) I.M. Johnst. X – – X

C. laevigata Lam. X – X X

C. polycephala (Lam.) I.M. Johnst. – – X – 

C. rickseckeri Millsp. X – – –

C. stenophylla Alain X – X X

C. sulcata DC. X – X X

Heliotropium angiospermum Murray X X X X

H. curvassavicum L. X X X –

H. fruticosum L. X – – X

H. indicum L. X – – –

H. procumbens Mill. X – – –

H. ternatum Vahl – – – X

Rochefortia acanthophora (DC.) Griseb. X X – X

Tournefortia hirsutissima L. – – X X

T. volubilis L. X X X –

Brassicaceae

Brassica sp. X – – –

Cakile lanceolata (Wild.) O.E. Schultz X – – –

Lepidum virginicum L. – – – X

Bromeliaceae

Bromelia pinguin L. X – X X

Hohenbergia portoricensis Mez – – X –

Pitcairnia angustifolia Aiton – – – X

Tillandsia polystachya (L.) L. – – X –

T. recurvata (L.) L. X X X X

Burseraceae

Bursera simaruba (L.) Sarg. X X X X

Tetragastris balsamifera (Sw.) Kuntze – – – X

Cactaceae

Harrissia portoricensis Britton X – – –

Hylocereus trigonus (Haw.) Safford – – X X

continued

Appendix B (continued)
100

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Cactaceae (continued)

Leptocereus quadricostatus (Bello) Britton & Rose X X X X

Melocactus intortus (Mill.) Urban X X – X 

Opuntia borinquensis Britton & Rose X – – –

O. cochenillifera (L.) Miller – – X –

O. dillenii (Ker-Gawl.) Haw. X – X X

O. repens Bello X X X X

O. tricantha (Willd.) Sweet X – – –

Pilosocereus royenii (L.) Byles & Rowley X X X X

Campanulaceae

Sphenoclea zeylanica Gaertn. – – X –

Canellaceae

Canella winterana (L.) Gaertner – X – –

Capparaceae

Capparis amplissima Lam. ssp. portoricensis – – X X

C. baducca L. X – – X

C. cyanophallophora L. – – – X

C. flexuosa (L.) L. X – X X

C. hastada Jacq. X X X X

C. indica (L.) Fawc. & Rendle X X X X

Cleome gynandra L. – – X –

C. spinosa Jacq. X – – –

C. stenophylla Klotzsch – – – X

C. viscosa L. X – – –

Celastraceae

Crosopetalum rhacoma Crantz X – X X

Elaeodendrum xylocarpum (Vent.) DC. X – – X

Gyminda latifolia (Sw.) Urban – – – X

Schaefferia frutescens Jacq. – – – X

Ceratophyllaceae

Ceratophyllum demersum L. – – X –

Chrysobalanaceae

Chrysobalanus icaco L. – X – X

Clusiaceae

Clusia gundlachii Stahl – – – X

C. rosea Jacq. X – X X

Garcinia hessi (Britton) Alain – – – X

Combretaceae

Bucida buceras L. X X X X

Conocarpus erectus L. X X – –

Laguncularia racemosa (L.) Gaertn. – X – –

Quisqualis indica L. X – – –

Terminalia catappa L. – X X X

Commelinaceae

Callisia repens (Jacq.) L. – – – X

Commelina diffusa Burn. F. – – – X

C. elegans Kunth in HBK. X – X –

continued

Appendix B (continued)
101

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Compositae

Acanthospermum hispidum DC. X – X X

Bidens alba (L.) DC.var. radiata (Sch.Bip.) Ballard in Melchert – – X –

B. cyanpiifolia Kunth – – – X

Borrichia arborescens (L.) DC. – X – –

Eclipta prostada (L.) L. X X X –

Emilia fosbergii Nicolson X – – –

Eupatorium odoratum L. X – – –

Gnaphalium indicum L. – X – –

Lagasoea mollis Cav. X – – X

Mikania micrantha HBK. X – X X

Partherium hysterophorus L. X – X X

Pectis cilaris L. X – – –

P. carthusianorum Less. X – – –

P. linifolia L. X – – –

Pluchea carolinensis (Jacq.) G. Don X – – –

P. odorata (L.) Cass. – – X –

Pterocaulon virgatum (L.) DC. – – – X

Synedrella nodiflora (L.) Gaertn. – – X X

Tridax procumbens L. X X X X

Verbesina encelioides (Cav.) Benth. & Hook. X – – –

Vernonia cinerea (L.) Less. X – X X

V. proctorii Urbatsch – – – X

Wedelia lanceolata DC. X X – X

Convolvulaceae

Convolvulus nodiflorus Desr. X – – X

Cuscuta americana L. – – – X

C. indecora Choisey X – X –

Evovulus alsinoides (L.) L. var. grisebachianus Meissn. X – – –

E. convolvuloides (Wild. Ex Schultes) Stern – – X –

E. nummularius (L.) L. – – X –

Ipomoea alba L. – – – X

I. guamoclit L. – – X – 

I. indica (Burm.f) Merr. var. acuminata (Vahl) Fosb. X – – X

I. nil (L.) Roth X – – –

I. ochracea (Lindl.) G. Don X – – –

I. steudelii Millsp. X – – X

I. triloba L. X – – X

Jacquemontia pentanthos (Jacq.) G. Don – – X –

J. tamnifolia (L.) Griseb. X – – –

Merremia aegyptia (L.) Urban X – – –

M. dissecta (Jacq.) Hall.f. X – – –

M. quinquefolia (L.) Hall.f X – – X

M. umbellata (L.) Hall.f. X – X X

Turbina corymbosa L. Raf. – – – X

Crassulaceae

Bryophyllum pinnatum (Lam.) Oken X X X X

Cruciferae

Rorippa portoricensis (Speng.) Stehle – – X –

continued

Appendix B (continued)
102

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Cucurbitaceae

Cucumis anguria L. X X X X

Doyerea emetocathartica Grossourdy – – – X

Melothria pendula L. X – – –

Momordica charantia L. X – X X

Ericaceae

Lyonia truncata Urban var. proctorii Judd – – – X

Erythroxlaceae

Erythroxylum areolatum L. X X X –

E. brevipes DC. – – – X

E. rotundifolium Lunan – – – X

E. urbanii O.E. Schulz in Urban – – – X

Euphorbiaceae

Acalypha bisetosa Bert. X – – –

A. portoricensis Muell. Arg. X – – X

Adelia ricinella L. – – X X

Alteramnus lucidus (Sw.) Rothm. – – – X

Argythamnia candicans Sw. X – – X

Caperonia palustris (L.) St. Hil. – – X –

Chamaesyce articulata (Aublet) Britton – – – X

C. glomerifera Millsp. X – – –

C. hirta (L.) Millsp. X – X –

C. hypericifolia (L.) Millsp. X – X –

C. hyssopifolia (L.) Small – – X –

C. prostata (Ait.) Small X – – –

Cnidoscolus aconitifolius (Mill.) I. M. Johnston X – – –

Croton astriotes Dryander – – – X

C. betulinus Vahl X – – X

C. discolor Willd. X – – –

C. glandulosus L. X – – –

C. humilis L. X – – –

C. lobatus L. X – X –

Dalechampia scandens L. X – – X

Euphorbia herterophylla L. X – – –

E. lactea Haw. X X – –

Gymnanthes lucida Sw. – – – X

Hura crepitans L. X – X –

Jatropha curcas L. – – X X

J. gossypifolia L. X – X X

J. hernandifolia Vent. X – – –

Margaritaria nobilis L.f. – – – X

Pedilanthus tithymaloides (L. Poiteau) ssp. tithymaloides – – – X

Phyllanthus amarus Schumach. X – X X

Ricinus communis L. X – X X

Securinega acidoton (L.) Rawcett & Rendle X – – –

Savia sessiliflora (Sw.) Willd. – – – X

Tragia volubilis L. X – X X

continued

Appendix B (continued)
103

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Flacourtiaceae

Casearia aculata Jacq. – – X X

C. decandra Jacq. – – – X

C. guianensis (Aublet) Urban – – – X

C. sylvestris Sw. – – X X

Prockia crucis L. – – – X

Samyda dodecandra Jacq. – – – X

Goodeniaceae

Scaevola plumieri (L.) Vahl – X – –

Guttiferae

Calophyllum calaba L. X – – X

Hippocrateaceae

Hippocratea volubilis L. – – – X

Krameriaceae

Krameria ixina L. – X – X

Labiatae

Hyptis capitata Jacq. – – X –

H. suaveolens (L.) Poit. X – – –

Leonotis nepetifolia L. X X X –

Ocium sanctum L. X – – –

O. campechianum Mill. – – – X

Lauraceae

Cassytha filiformis L. – – X X

Licaria parvifolia (Lam.) Kostern. – – – X

Nectrandra coriacea (Sw.) Griseb. – – – X

Lemnaceae

Lemna aequinoctialis Welw. – – X –

L. perpusilla Torry – – X –

L. polyrhiza (L.) Schleiden X – X –

Leguminosae - Caesalpinioidae

Bauhinia monandra Kurz – – – X

Caesalpinia bonduc (L.) Roxb. – X – –

Cassia chamaecrista L. X – – –

C. fistula L. – – X –

C. obtusifolia L. X – – –

Chamaecrista glandulosa var. swartzii

(Wikstr.) Irwin & Barneby – – X –

C. nictitans (L.) Moench – – – X

Hymenaea courbaril L. X X X X

Parkisonia aculeata L. X X X –

Senna obtusifolia (L.) Irwin & Barneby X – X –

S. occidentalis (L.) Link – – X –

S. polyphylla (Jacq.) Irwin & Barneby X – X X

S. siamea (Lam.) Irwin & Barneby – X –

Stahlia monosperma (Tul.) Urban X X X –

Tamarindus indica L. X X X X

continued

Appendix B (continued)
104

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Leguminosae - Faboidae 

Abrus precatorius L. X – X X

Aeschynomene americana L. – – X –

A. sensitiva Swartz – – X –

Alysicarpus vaginalis (L.) DC. X – X X

Andira inermis (W. Wright) Kunth X – X X

Centrosema virginianum (L.) Benth. X – X –

Clitoria ternatea L. X – – X

Coursetia caribaea (Jacq.) Lavin – – – X

Cracca caribaea (Jacq.) Benth X X – X

Crotalaria incana L. – – X –

C. falcata Vahl ex DC. – – – X

C. lotifolia L. – – – X

C. pallida Ait. – – X –

C. retusa L. X – X –

Desmodium glabram (Mill.) DC. X – – –

D. incanum DC. – – – X

D. procumbens (Mill.) Hitchc. X – – X

D. tortuosum (Sw.) DC. X – – –

Galactia dubia DC. – – – X

G. striata (Jacq.) Urban X – – –

Gliricidia sepium (Jacq.) Kunth ex Walp. – – – X

Indigofera suffruticosa Mill. – – X X

I. tinctoria L. X – – –

Lonchocarpus domingensis (Pers.) DC. – – X –

Macroptilium lathyroides (L.) Urban X – X –

Pictetia aculeata (Vahl) Urban X – X X

Piscidia carthagenensis Jacq. X X – –

Poitea paucifolia (DC.) Lavin X – – –

Pterocarpus officinalis Jacq. – – X –

Rhynchosia minima (L.) DC. X – X –

R. reticulata (Sw.) DC. – – X –

Sesbania emerus (Aubl.) Urban – – X –

Sesbania sericea (Willd). Link X X X –

Stylosanthes hamata (L.) Taub. X – – X

S. viscosa Sw. – – – X

Tephrosia cinerea (L.) Pers. X – X –

T. senna HBK. X – – –

Teramnus labialis (L.f.) Spreng. X – – –

Zornia reticulata J.E. Smith X – – X

Leguminosae - Mimosoideae

Acacia farnesiana (L.) Willd. X – X X

A. retusa (Jacq.) Howard – – X X

A. tortuosa (L.) Willd. – – – X

Albizia lebbeck (L.) Benth. X X X X

A. procera (Roxb.) Benth. – – – X

Dasmanthus virgatus (L.) Willd. X – X X

Inga vera Willd. – X X –

Leucaena leucocephala (Lam.) de Wit X X X X

continued

Appendix B (continued)
105

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Leguminosae - Mimosoideae (continued)

Minosa casta L. – – – X

M. ceratonia L. – – X –

M. pudica L. – X X X

Neptuna plena (L.) Benth. in Hook – – X –

Pithecellobium dulce (Roxb.) Benth. X X X X

P. ungis-cati (L.) Mart. X X X –

Prosopis juliflora (Sw.) DC. X X X X

Samanea saman (Willd.) Merr. X – X X

Loganiaceae

Spigelia anthelmia L. – – X –

Loranthaceae

Dendropemon purpureus (L.) Krug & Urban – – X –

Malphighiaceae

Bunchosia glandulosa (Cav.) L.C. Rich X – – –

Byrsonima lucida (Mill.) L.C. Rich. X – – –

B. spicata (Cav.) HBK. X – – X

Heteropteris purpurea (L.) Kunth X – – X

H. laurifolia (L.) A. Juss. X – – X

Stigmaphyllon emarginatum (Cav.) A. Juss. X – X X 

S. floribundum (DC.) C. Anders. – – X –

S. periplocofolium (Desf.) A. Juss. – – X X

S. tomentosum (Desf.) Ndz. – – X X

Malvaceae

Abutilon umbellatum (L.) Sweet X – – –

Bastardia viscosa (L.) HBK. X – – X

Hibiscus phoeniceus Jacq. X – – X

Gossypium barbadense L. var. barbadense – X – X

Malachra capitata (L.) L. – – X –

M. alceifolia Jacquin X – X –

M. urens Poiteau ex Ledeb. & Aldestram – – X –

Malvastrum americanum (L.) Torrey X – – –

M. corchorifolium (L.) Garcke – – X X

M. coromandelianum (L.) Garcke X – – –

Pavonia spinifex (L.) Cavanilles – – – X

Sida abutilifolia Mill. X – – X

S. acuta Burm. f. X X – X

S. alba L. X – X –

S. ciliaris L. – – X –

S. cordifolia L. X – – –

S. glabra Mill. X – – –

S. glomerata Cav. X – X X

S. jamaicensis L. X – – X

S. pyramidata Desp. ex Cav. – – – X

S. rhombifolia L. X – – X

S. salviifolia Presl. X – X –

S. spinosa L. X – X –

S. stipularis Cav. – – – X

S. urens L. – – X –

Sidastrum multiflorum (Jacq.) Fryxell X – X –

continued

Appendix B (continued)
106

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Malvaceae (continued)

Thespesia populnea (L.) Solander ex Correa X X X –

T. grandiflora DC. – – X –

Urena lobata L. ssp. lobata – – X X

Wissadula amplissima (L.) R.E. Fries X – X X

W. periplocifolia (L.) Presl. – – X X

Meliaceae

Azadirachta indica A. Juss. – – X X

Cedrela odorata L. X – – –

Guarea guidonia (L.) Sleumer – – X –

Melia azedarach L. X – X –

Swietenia mahagoni (L.) Jacq. X X – X

Trichilia hirta L. X X X X

Menispermaceae

Cissampelos pareira L. – – – X

Menyanthaceae

Nymphoides indica (L.) Kuntze – – X –

Molluginaceae

Mullugo verticulata L. X – – –

Moraceae

Cecropia schreberiana Miq. – – – X

Ficus americana Aubl. – – X –

F. benjamina L. X – – –

F. citrifolia Mill. X – X X

Moringaceae

Moringa oleifera Lamarck – – X –

Myrsinaceae

Ardisia obovata Hamilt. – – – X

Myrtaceae

Eugenia biflora (L.) DC. – – – X

E. foetida Pers. – X – –

E. lingustrina (Sw.) Willd. – – – X

E. monticola (Sw.) DC. – – – X

E. procera (Sw.) Poir. – – – X

E. pseudopsidium Jacq. – – – X

E. rhombea (Berg.) Krug & Urban – – – X

E. sessiliflora Vahl X X – X

E. woodburyana Alain X – – X

Myrcia citrifolia (Aublet) Urban – – – X

M. splendens (Sw.) DC. – – – X

Myrciaria floribunda (West ex Wild.) Berg – – – X

Pimenta recemosa (Miller) J.W. Moore var. racemosa X – – X

Psidium guajava L. – – – X

Nyctaginaceae

Boerhavia coccinea Mill. – – – X

B. diffusa L. X – – X

B. erecta L. – – – X

Commicarpus scandens (L.) Standl. X – X X

continued

Appendix B (continued)
107

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Nyctaginaceae (continued)

Guapira fragrans (Dum.-Cours) Little – – X X

Neea buxifolia (Hook. f.) Heimerl – – – X

Pisonia aculeata L. – – – X

P. albida (Heimerl) Britton ex Standl. X – – –

P. subcordata Sw. X – X –

Nymphaeaceae

Nymphaea ampla (Salisb.) DC. – – X –

N. odorata Dryand. in Ait. – – X –

N. pulchella DC. – – X –

Ochnaceae

Ouratea littoralis Urban – – – X

Oleaceae

Forestiera segregata (Jacq.) Krug & Urban – – – X

Jasminum fluminense Velloso X X X X

Onagraceae

Ludwigia erecta (L.) H. Hara X – X –

L. leptocarpa (Nuttal) H. Hara – – X –

L. octovalvis (Jacquin) Raven – – X –

L. peploides (HBK.) Raven – – X –

Papaveraceae

Argemone mexicana L. – – X X

Passifloraceae

Passiflora edulis Sims – – X –

P. suberosa L. X – – X

Phytolaccaceae

Petiveria alliacea L. – – X X

Rivina humilis L. X X X X

Trichostigma octandrum (L.) H. Walt. – – X X

Piperaceae

Peperomia humilis A. Dietr. – – – X

P. obtusifolia (L.) A. Dietr. – – – X

Polygalaceae

Polygala cowellii (Britton) S.F. Blake X – – –

P. penaea L. – – – X

Securidaca virgata Sw. – – – X

Polygonaceae

Antigonon leptopus Hook. & Arn. X – – –

Coccoloba diversifolia Jacq. X – – X

C. microstachya Willd. X – – X

C. swartzii Meissn. X – – –

C. uvifera (L.) L. X X – –

C. venosa L. – – – X

Polygonum acuminatum HBK. – – X –

P. glabrum Willd. in L. – – X –

P. punctatum Ell. – – X –

P. segetum HBK. – – X –

continued

Appendix B (continued)
108

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Portulacaceae

Portulaca grandiflora Hook. – – – X

P. halimoides L. X – – –

P. oleracea L. X X X X

P. pilosa L. X – – X

P. quadrifida L. X – X X

Talinum fruticosum (L.) A.L. Juss. X – X X

T. paniculatum (Jacq.) Gaerth. – – – X

T. triangulare (Jacq.) Willd. X – – –

Ranunculaceae

Clematis dioica L. – – – X

Rhamnaceae

Colubrina arborescens (Mill.) Sarg. X X – –

Gouania lupuloides (L.) Urban – – – X

Krugiodendron ferreum (Vahl) Urban X X – X

Reynosia uncinata Urban X – – –

Ziziphus reticulata (Vahl) DC. X – – X

Rhizophoraceae

Rhizophora mangle L. X X – –

Rubiaceae

Chiococca alba (L.) Hitchc. – – – X

Diodia apiculata (Willd.) K. Schum. X – – X

D. rigida Cham. & Schldl. X – – X

Erithalis fruticosa L. – X – X

Exostema caribaeum (Jacq.) Roem. & Schult. X – – X

Guettarda elliptica Sw. X – – –

G. scabra (L.) Vent – – – X

Hamelia patens Jacq. – – X X

Hedyotis corymbosa (L.) Lam. Tabl. – – – X

Machaonia portoricensis Baill. – – – X

Psychotria microdon (DC.) Urban – – – X

P. pubescens Sw. – – – X

Randia aculeata L. X X X X

Rondeletia inermis (Spreng.) Krug & Urban – – – X

Spermacoce assurgens Ruiz & Pavon X – – –

S. confusa Rendle & Gillis X – – –

S. prostrata Aubl. X – – –

S. repens (DC.) Fosberg & Powell X – – –

S. verticillata (L.) Meyer X – X –

Rutaceae 

Amyris elemifera L. – – – X

Citrus aurantifolia (Christm.) Swingle X – X –

C. limon (L.) Burm. f. X – X –

C. sinensis (L.) Osbeck – – X –

Zanthoxylum flavum Vahl X – X –

Z. martinicense (Lam.) DC. X – – X

Z. monophyllum (Lam.) P. Wilson X – X X

continued

Appendix B (continued)
109

Plant species at the Cabo Rojo and Laguna Cartagena National Wildlife Refuges

Refuge a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Sapindaceae

Allophyllus racemosus (Sw.) – – – X

Cardiospermum corundum L. X – – –

Cupania americana L. – – X X

Dodonaea americana L. – X – –

Hypelate trifoliata Sw. – – – X

Melicoccus bijugatus Jacq. X – X X

Paullinia pinnata L. – – X X

Sapindus saponaria L. – – X X

Serjania polyphylla (L.) Badlk. – – X X

Thouinia striata Radlk. var. portoricensis (Radlk.) 

Votava & Alain X – – X

Triphasia trifolia (Burm.f) P. Wilson X – – –

Sapotaceae

Bumelia obovata (Lam.) A. DC. X X X –

Chrysophyllum oliviforme L. – – – X

Manilkara pleeana (Pierre) Cronq. – – – X

Sideroxylon foetidissimum Jacq. X – X X

S. obovatum Lam. – X – X

Scrophulariaceae

Bacopa monnieri (L.) Pennell – – X –

Capraria biflora L. X X X X

Scoparia dulcis L. X – X –

Solanaceae

Capsicum frutescens L. – – X X

Datura inoxia Mill. X X – –

D. metel L. – – – X

D. stramonium L. – – X –

Goetzea elegans Wydler X – – –

Physalis angulata L. X – X –

Solanum americanum var. nodiflorum L. – – X –

S. campechiense L. – – X –

S. drymophilum O.K. Schultz – – – X

S. persicifolium Dunal X – X X

S. rugosum Dunal – – – X

S. torvum Sw. – – X X

Sterculiaceae

Ayenia insulicola Cristóbal X – – –

Guazuma ulmifolia Lam. X – X X

Helicteres jamaicensis Jacq. X – – X

Melochia nodiflora Sw. – – X X

M. pyramidata L. X X X X

M. tomentosa L. X X X X

M. villosa (Mill.) Fawcett & Rendle – – X –

Sterculia apetala (Jacq.) Karst X – – –

Waltheria indica L. X X – X

Theophrastaceae

Jacquinia arborea Vahl X X – –

continued

Appendix B (continued)
110

Plant species at the Cabo Rojo and Laguna National Wildlife Refuges

Refuges a

Cabo Rojo

Laguna 

Cartagena

Family and species H S L T

Tiliaceae

Corchorus aestuana L. X – – –

C. hirsutus L. X – – –

C. orinocensis Kunth in HBK. X – – –

C. siliquosus L. X – – –

Triumfetta semitriloba Jacquin – – X –

Turneraceae

Piriqueta ovata (Bello) Urban X – – X

Turnera diffusa Willd. X – – X

T. ulmifolia L. X – – –

Umbelliferae

Hydrocotyle verticillata Thunberg – – X –

Urticaceae

Pilea microphylla (L.) Liebm. – – X – 

Verbenaceae

Bouchea prismatica (L.) Ktze X – – X

Citharexylum fruitcosum L. X – X X

Clerodendrum aculeatum (L.) Schlecht X – – –

Lantana camara L. X X X X

L. involucrata L. X X X –

L. reticulata Pers. – – X –

L. urticifolia Miller – – X –

Lippia micromera Schaer. var. helleri (Britton) Mold. – – – X

L. nodiflora (L.) Michx. X – X –

L. strigulosa Martens & Gal. – – – X

Petitia domingensis Jacquin – – – X

Priva lappulacea (L.) Pers. – – X –

Stachytarpheta jamaicensis (L.) Vahl X X X X

Tamonea boxiana (Mold.) Howard – – X X

T. spinosa Sw. X – – X

Vitex divaricata Sw. – – – X

Viscaceae

Phoradendron anceps (Spenng.) Krug & Urb. X – – –

P. quarangulare (Kunth) Griseb. X – X X

P. randiae (Bello) Britton – – – X

P. tetrapterum Krug & Urban – – X –

Vitaceae

Cissus trifoliata (L.) L. X – – X

C. verticillata (L.) Nicolson & Jarvis – – – X

Zygophyllaceae

Guaiacum officinale L. X X – X

Kallstroemia maxima (L.) Hook. & Arn. X – – X

Tribulus cistoides L. X – – – 

Sources: Cabo Rojo Refuge (McKenzie 1986, see endnotes); Laguna Cartagena Refuge (Breckon 1998; 

Diaz-Soltero 1990; Proctor 1996, see endnotes); both refuges (Joseph Schwagerl, personal observations).

a H = Headquarters tract, S = Salinas tract, L = Lagoon tract, and T = Tinafa tract.

Appendix B (continued)
Weaver, Peter L.; Schwagerl, Joseph J. 2009. U.S. Fish and Wildlife Service refuges

and other nearby reserves in southwestern Puerto Rico. Gen. Tech. Rep. IITF-40. 

San Juan, PR: U.S. Department of Agriculture Forest Service, International Institute 

of Tropical Forestry. 110 p.

The main purpose of this paper is to summarize information about the Cabo Rojo and Laguna 

Cartagena National Wildlife Refuges in southwestern Puerto Rico. The Cabo Rojo (Headquarters 

and Salinas tracts) and Laguna Cartagena (Lagoon and Tinaja tracts) occupy 1174 ha and are 

part of the 10 231-ha Caribbean system of nine refuges. Their major geologic and physiographic 

features are Sierra Bermeja, the oldest mountain range in Puerto Rico, and the floodplains 

of the Lajas Valley. Together, the refuges combined contain 9 geologic features and 26 soils’ 

types. Rainfall averages about 1000 mm/year and hurricanes occur periodically. Among the 

several vegetation types, including are mangroves, salt flats, littoral woodland, mesquite and 

semievergreen woodland, coastal shrub, deciduous woodland, and pasture. At least 644 plant 

species grow on the refuges. Eight of these species are considered rare and endangered, and have 

formal recovery plans. Refuge mammals include two species of native bats and six introduced 

species—two monkeys, a mongoose, two rats, and a mouse. Both the Cabo Rojo Salt Flats and 

Cartagena Lagoon are renowned as wintering habitat for migratory birds where >200 resident 

and migratory birds have been observed, including 12 island endemics. Moreover, several other 

species are listed as rare and endangered, or of conservation concern, or as target species for 

management. Among the reptiles are seven lizards, two geckos, and three turtles (one terrestrial 

and two marine). Five amphibians (one toad and four frogs) and 18 species of fish (fresh and 

brackish water) are also present. Six of the wildlife species have formal recovery plans. 

The major management activities on the refuges include protection against fires, habitat 

restoration, public education, special uses, and water management at the Cartagena Lagoon and 

the Salinas salt flats. Habitat recovery has been stimulated through fire control, through livestock 

removal, and by planting >9,000 trees of nearly 80 species. Public education encompasses nearly 

20,000 visits annually to interpretative centers at the Headquarters and Salinas tracts; moreover, 

another 2,600 visitors frequent the refuges to observe wildlife and take nature photographs, and 

nearly 4,900 others participate in educational and interpretative programs. Special use permits 

have been issued to 174 users since 1979, mostly for administrative matters or environmental 

research. Finally, water management at the Cartegena Lagoon remains a problem, partly because 

of historical events and partly due to current difficulties. 

In addition to the U.S. Fish and Wildlife Service (F&WS) refuges, six other major protected 

areas occupy about 6800 ha along the southwestern coast: Punta Guaniquilla Natural Reserve, 

Boquerón Wildlife Refuge, Los Morrillos de Cabo Rojo, Boquerón Forest, Parguera Natural 

Reserve, and Guánica Forest. An additional 23 bird species not recorded on the refuges have been 

observed in these protected areas. The F&WS refuges and the protected areas provide critical 

wildlife habitat in Puerto Rico’s southwest—an area that is experiencing a rapid increase in 

human population. 

Keywords: Cabo Rojo Wildlife Refuge, Cartagena Lagoon Wildlife Refuge, fauna, flora, 

climate, geology, soils, management.
112

The Forest Service, U.S. Department of Agriculture 

(USDA), 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.

The USDA prohibits discrimination in all its programs and activities 

on the basis of race, color, national origin, age, disability, and 

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

religion, sexual orientation, genetic information, political beliefs, 

reprisal, or because all or part of an individual’s income is derived 

from any public assistance program. (Not all prohibited bases 

apply to all programs.) Persons with disabilities who require 

alternative means for communication of program information 

(Braille, large print, audiotape, etc.) should contact USDA’s 

TARGET Center at (202) 720-2600 (voice and TDD).

To file a complaint of discrimination, write to USDA, Director, 

Office of Civil Rights, 1400 Independence Avenue, SW, 

Washington, DC 20250–9410 or call (800) 795-3272 (voice) or 

(202) 720-6382 (TDD). USDA is an equal opportunity provider 

and employer.