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Conserving the Puerto Rican herpetofauna

Article  in  Applied Herpetology · October 2007

DOI: 10.1163/157075407782424566

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Conserving the Puerto Rican herpetofauna

Rafael L. Joglar1,* , Alberto O. Álvarez2 , T. Mitchell Aide1 , Diane Barber3 , Patricia

A. Burrowes1 , Miguel A. García2,4 , Abimael León-Cardona5 , Ana V. Longo1 ,
Néstor Pérez-Buitrago1,4 , Alberto Puente1,2 , Neftalí Rios-López1 , Peter J. Tolson6
1 Department of Biology, University of Puerto Rico, PO Box 23360, San Juan,
Puerto Rico 00931-3360
∗ Corresponding author; e-mail: [email protected]
2 Division of Wildlife, Department of Natural and Environmental Resources, P.O. Box 366147,

San Juan, Puerto Rico 00906

3 Fort Worth Zoo, 1989 Colonial Parkway, Fort Worth, Texas 76110, USA
4 Center for Applied Tropical Ecology and Conservation (CATEC), University of Puerto Rico–Río

Piedras, San Juan, Puerto Rico 00931

5 Aquatic Toxicology Program, University of California-Davis, VetMed:APC, 1321 Haring Hall,

Davis, California 95616, USA

6 Conservation Department — Toledo Zoo, Ohio 43614, USA

Abstract. With a total area of 8900 km2 , Puerto Rico is the smallest of the Greater Antilles. It is
divided in three physiographic regions or areas of relief: the mountainous interior, the karst region,
and the coastal plains and valleys. The island comprises six ecological life zones: subtropical dry
forest, subtropical moist forest, subtropical wet forest, subtropical rain forest, lower montane wet
forest and lower montane rain forest. The herpetofauna of Puerto Rico consists of 25 species of
amphibians (19 native, six introduced) and 56 species of reptiles (52 native, four introduced). The
goal of this paper is to describe some of the present studies directed towards the conservation of
Puerto Rican herpetofauna. Eleutherodactylus karlschmidti, E. jasperi and E. eneidae have not been
seen or heard since 1976, 1981 and 1990, respectively, and are probably extinct. Since 2000, the
potential causes of amphibian declines in Puerto Rico have been studied, and a synergistic interaction
between climate change (increased dry periods) and disease (chytridiomycosis) have been proposed
as an explanation for the patterns observed. Recovery efforts for Peltophryne lemur include a captive-
breeding program, reintroductions island-wide educational outreach, protection and restoration of
existing habitat, and the creation of new breeding ponds. Among reptiles, the first conservation efforts
to protect Epicrates inornatus were limited to trying to halt collection and hunting. However, current
strategies to preserve the boa include gathering basic biological information, habitat conservation,
and educational outreach. Recent efforts for the conservation of Trachemys s. stejnegeri combine
three research approaches to clarify the status of local populations: a mark-recapture-release study,
field monitoring of reproductive activity (i.e., nocturnal patrolling to identify nesting activity), and
field assessment of the potential impact of introduced species, particularly identification of predatory
species and exotic turtles. Recovery initiatives for Cyclura stejnegeri include management of invasive

© Koninklijke Brill NV, Leiden, 2007 APPLIED HERPETOLOGY 4: 327-345

Also available online - www.brill.nl/ah
328 R.L. Joglar et al.

mammals, a headstart program for hatchling iguanas, and the assessment of the etiology of a condition
causing blindness in adult iguanas. A reforestation project aimed at recovering a local herpetofaunal
assemblage after disturbances in a limestone valley in northern Puerto Rico is discussed. As population
sizes of common colonizers such as Eleutherodactylus and Anolis increased, larger forest-interior
and predatory species like Epicrates inornatus, Alsophis portoricensis and Anolis cuvieri followed.
Finally, the Mona Island marine turtle monitoring program is discussed and compared to other similar
programs in Puerto Rico. As these and other similar conservation efforts provide scientifically based
management recommendations, we hope to succeed in conserving the diverse herpetofauna that
characterizes Puerto Rico.

Key words: Amphibian; biodiversity; Caribbean; chytrid; climate change; conservation; Puerto
Rico; reptile; West Indies.

Introduction
Puerto Rico is the easternmost of the Greater Antilles and it is located between
18◦ 35′ −17◦ 55′ N and 67◦ 15′ −65◦ 35′ W. It is the smallest of the Greater Antilles with
a total area of 8900 km2 , 12.9, 8.6 and 1.3 times smaller than Cuba, Hispaniola, and
Jamaica, respectively. It is 179 km long and 58 km wide, and its highest peak is
Cerro Punta in Jayuya (elevation 1338 m). The highest peaks on Cuba, Hispaniola,
and Jamaica, respectively, are 1.5, 2.3 and 1.7 times higher than Cerro Punta. The
three largest Puerto Rican satellites islands are Vieques (138 km2 ) and Culebra
(27 km2 ) in the east, and Mona (57 km2 ) in the west.

Physiography and general ecology

Puerto Rico is divided in three physiographic regions or areas of relief: (1) the
mountainous interior; (2) the karst region; and (3) the coastal plains and valleys
(Cruz and Boswell, 1997; Gannon et al., 2004).
The mountainous interior is the largest of the three regions, and it is the cradle of
the main rivers of the island. This ‘backbone’ or mountainous spine is composed of
the Cordillera Central and the Sierra de Luquillo. The Cordillera Central extends in
an east-west direction from Sierra de Cayey in the southeast running almost without
interruptions to Mayagüez, which lies close to the west coast. Sierra de Luquillo,
an isolated upland region in the northeast, and the largest natural protected area of
Puerto Rico, shelters montane rainforests at lower elevations and cloud forests and
elfin woodland at the highest elevations; one of the largest tracks of elfin woodland
in the Caribbean is found in these mountains (Hedges, 1999).
The Puerto Rican karst region includes two separate zones: (1) the northern karst,
which ranges from Loiza in the northeast to Aguadilla in the northwest; and (2)
the southern karst, which extends from Juana Díaz (east of Ponce) to Guayanilla
to the west (Cruz and Boswell, 1997; Lugo, 2005). The Puerto Rican northwestern
karst topography is similar to that in Cockpit Country of Jamaica, Los Haitises of
Hispaniola, and the Viñales region of western Cuba (Hedges, 1999).
Conserving the Puerto Rican herpetofauna 329

According to the Holdridge system of classification, Puerto Rico has been divided
into six ecological life zones: (1) subtropical dry forest; (2) subtropical moist forest;
(3) subtropical wet forest; (4) subtropical rain forest; (5) lower montane wet forest;
and (6) lower montane rain forest (Ewel and Whitmore, 1973). Average annual
rainfall is 60-110; 100-220; 200-400 cm in the first three zones, respectively, with a
lower rainfall limit of 380 cm in the fourth zone, and with elevations over 1000 m in
the lower montane wet forest and lower montane rain forest. An alternative system
has been proposed by Lugo (2005), in which geological data were incorporated
with temperature, precipitation, and elevation information of the Holdridge system
to generate ten geoclimatic zones or forest types for Puerto Rico.

Conservation in Puerto Rico

Because Puerto Rico is a US Territory governed by a commonwealth, state and fed-
eral agencies and their laws are involved in the island’s conservation practices. At
the state level, the Planning Board, the Environmental Quality Board, the Depart-
ment of Agriculture, the Department of Natural and Environmental Resources, and
all 78 municipalities through their Territorial Arrangement Plan, deal with corre-
sponding conservation issues (Quevedo, in press). The main state agency actively
engaged in conservation in Puerto Rico is the Department of Natural and Envi-
ronmental Resources (DNER). Established in 1972, this agency has the difficult
responsibility of protecting and managing Puerto Rican natural resources, including
biodiversity. Among many other responsibilities, the DNER is in charge of endan-
gered species, forest reserves, wildlife refuges, and corridors. The DNER’s capacity
to accomplish these goals and responsibilities are seriously limited by (1) politi-
cal pressures, (2) budget and (3) excessive responsibilities. At the federal level, the
two most important agencies involved in conservation are (1) the U.S. Fish and
Wildlife Service (USF&WS) as part of the US Department of Interior (USDI), and
(2) the U.S. Forest Service (USFS) as part of the U.S. Department of Agriculture
(USDA). Other federal agencies such as the U.S. Environmental Protection Agency
(EPA), the USDA Natural Resources Conservation Service (NRCS) and the U.S.
Corps of Engineers, are also involved. The USF&WS is responsible for endangered
species and their habitats and for managing several federal wildlife refuges, such
as Desecheo, Vieques and Cabo Rojo. The USFS is responsible for protecting and
managing El Yunque, the largest forest reserve in Puerto Rico. Within the USFS,
the International Institute of Tropical Forestry (IITF) is in charge of research and
management of forest resources. As in the Continental USA, in Puerto Rico the
USF&WS has not been effective at listing endangered species and declaring and
protecting critical habitat for listed species. This shows that political pressures and
budget limitations for conservation agencies are also a problem at the federal level.
Puerto Rico’s history of protecting areas for conservation purposes goes back to
1876, when the Spanish Crown protected areas in El Yunque and Utuado. Laws of
the Spanish Crown protected forests, mangroves and water resources. Since then,
Puerto Rico has created a system of forest reserves that includes 21 forest reserves
330 R.L. Joglar et al.

(such as Maricao, Toro Negro, Carite, Cambalache and Susua) administrated by

the state government (DNER), El Yunque administrated by the federal government
(USFS-USDA), and San Patricio, Monte La Choca, Nuevo Milenio and Del Pueblo
are co-managed by the state government and local environmental groups that are
actively involved in managing and protecting these areas (Sanchez-Martínez, 2007).
In addition to these 26 forest reserves, Puerto Rico also has wildlife refuges (federal
and state) and natural reserves such as Mona Island and Caja de Muertos. Since all
these areas are isolated from each other, a series of corridors have been proposed to
connect some of them and increase their conservation value.
A very important asset to the conservation of land and biodiversity in the island
is the Puerto Rico Conservation Trust, a private corporation. Since 1970 it has
managed to acquire and protect 20 natural areas for conservation for a total of
21 364 acres (86.5 km2 ) which is the size of Mona Island and Culebra combined.
In addition, they offer excellent interpretative nature programs at Las Cabezas de
San Juan Natural Reserve in Fajardo, Hacienda Buena Vista in Ponce and in La
Esperanza en Manatí.
Taking all protected areas into consideration, including state and federal forest
reserves, wildlife reserves and other preserved areas mentioned above, Puerto Rico
protects 7.2% of its territory for conservation. To put this number in perspective,
the United States protects 25% of its territory, Costa Rica 34%, Cuba 32%, the
Dominican Republic 42% and the Virgin Islands, where tourism plays an important
role in the economy, 54% (Lloveras, in press). In 2007 a group of scientists
discussing the effect of climate change on Puerto Rican biodiversity and natural
resources requested the government to protect 25% of the Puerto Rican territory.
We hope that this goal is met in the near future and surpassed thereafter.

The Puerto Rican Herpetofauna and Their Conservation

Twenty-five species of amphibians occur in Puerto Rico: 19 are native and six
are introduced (Joglar, 2005a; Rios-López and Thomas, 2007; see the appendix
for a complete list). The number of native species of amphibians in Puerto Rico
(19) is high, especially considering the size of the island. Puerto Rico has the
highest number of species of amphibians per area in comparison to the other
Greater Antillean islands (table 1) and many other islands of the world (Duellman,
1999a,b; Joglar, 2005a). However, the number of families and genera of native
amphibians is low, with only two families (Leptodactylidae and Bufonidae) and
three genera (Eleutherodactylus, Leptodactylus and Peltophryne) present. Most of
the native species belong to the genus Eleutherodactylus (89.5%) and to the family
Leptodactylidae (94.7%). Endemism is high; most native species (15/19 = 78.9%)
are endemic to Puerto Rico (Mona Island included); however, four additional
species are endemic to the Greater Puerto Rican Bank, as they are also found in
the Virgin Islands and on other islands that were connected to Puerto Rico during
low sea levels associated with glacial maxima. Six species of amphibians have been
Conserving the Puerto Rican herpetofauna 331

Table 1. Species of amphibians (native, endemic and introduced) and percentage of endemism.
Adapted from Hedges (1999) and Duellman (1999a,b). Hispan refers to Hispaniola, PR Bank to Puerto
Rican Bank, PR to Puerto Rico and Lesser A. to Lesser Antilles.
Native Endemic Percentage Introduced Total Area Species/
species species of endemism species species (km2 ) 106 km2
Cuba 52 50 96 2 54 110 922 469
Hispan 63 63 100 2 65 76 470 824
Jamaica 22 22 100 4 26 10 990 2002
PR Bank 21 21 100 6 27 9 511 2103
PR 19 15 (19) 78.9 (100) 6 25 8 990 2002
Lesser A. 10 9 90 4 14 5 840 1712
Bahamas 2 0 0 4 6 11 296 177

Table 2. Number of reptiles; a according to Uezt (2000; web page); b according to Powell et al. (1996).
Worlda West Indies Puerto Exotics Endemic Threatened/
Rico in PR species endangered
Reptiles 8002 500a 56 4 89% (42/47) 22-24%
Turtles 296 13b 7 1 0 83% (5/6)
Lizards 4610 321b 34 2 87% (28/32) 12% (4/32)
Amphisbaenians 160 12b 4 0 100% 0
Snakes 2911 107b 10 ? 90% 22% (2/9)
Crocodilians 23 4b 1 1 0 0
Tuataras 2 0 0 – – –

introduced in Puerto Rico since the 1920s (Joglar, 2005a); Puerto Rico has the
highest number of introduced species in the Greater Antilles (table 1).
Fifty-six species of reptiles occur in Puerto Rico; 52 are native and four are
introduced (Joglar, 2005b; see the appendix for a complete list). Included as native
species are five marine turtles that have a wide distribution in the West Indies and
in the rest of the world. Of the terrestrial native species, 89.4% (42/47) are endemic
to Puerto Rico and the Puerto Rican Bank (table 2).
The status of Puerto Rican amphibians has been reviewed extensively (Joglar
and Burrowes, 1996; Joglar, 1998; Burrowes et al., 2004; Burrowes and Joglar,
2005) — see next section. Eleven reptilian species are currently protected as
threatened or endangered (appendix). Some common characteristics among these
species are: (1) large body size; (2) specialists in habitat, behavior, or morphology;
(3) limited distributions; (4) human consumption or commercial value; and (5)
feared and persecuted by humans (Joglar, 2005b). For a summary of factors
affecting vulnerability, major threats, conservation activities, and recommendations,
see Wilson et al. (2006). The following sections summarize seven important efforts
regarding the conservation of Puerto Rican amphibians and reptiles; additional
efforts and conservation projects have been implemented but are not included here.
332 R.L. Joglar et al.

Puerto Rican Amphibians, Pathogens and Climate Change

Puerto Rico was one of the first countries in the world which noticed what has
become an international phenomon known as ‘declining amphibian populations’.
We started monitoring amphibian populations in 1988 at El Yunque and, since then,
have monitored more than 20 sites with an emphasis on the highlands. We have
not found Eleutherodactylus karlschmidti, E. jasperi, or E. eneidae since 1976,
1981 and 1990, respectively, in spite of continuous field expeditions. An interesting
population effect is observable at our elfin forest site in the Caribbean National
Forest (El Yunque), where E. portoricensis and E. coqui seem to be recovering
from population declines observed in the early 1990s. However, populations of three
other species (E. wightmanae, E. richmondi and E. locustus) have not recolonized
some areas at El Yunque where they were abundant in the past, and are still very
Please, need data
scarce in that forest. Fortunately, these species still occur in some parts of the
island, and we are monitoring them carefully. Since 2000, we have been studying
the potential causes of amphibian declines in Puerto Rico, and have proposed
a synergistic interaction between climate change (increased dry periods) and
disease (chytridiomycosis) as an explanation for the patterns observed (Burrowes
et al., 2004). Our data suggest that, during times of drought, frogs may clump in
humid patches of the forest where the fungus is most likely to occur. This could
provoke a situation where frogs, stressed from lack of water and clumping, become
immunologically deprived in an area where the fungus is potentially abundant. As a
consequence, infections by chytrid may increase, generating an epidemic that results
in amphibian declines. Recent work led by a graduate student at our lab (Ana Longo)
has tested part of this hypothesis. We established a controlled experiment with
six terraria in which the number of refugia, food, light and temperature remained
constant. Water was applied to the soil on only one half, and later to one fourth of
the experimental terraria, while controls received water over the entire surface. The
use of retreat sites by high-mountain E. coqui was monitored daily. Frog dispersion
within the terraria changed significantly as a result of the water treatments (ANOVA
F1,35 = 13.6; P = 0.001). In the experimental terraria, frogs moved from dry to
humid sides within three days of drought. Thus, in spite of their territorial behavior,
frogs facing limited water supplies clumped in the humid retreat sites, whereas
control frogs used refuges on both sides. The next step of this experiment is to
inoculate frogs with equal numbers of chytrid zoospores to determine if those forced
to clump by a limited water regime are more susceptible to chytridiomycosis. We
expect to have results on this work in the near future to better understand patterns
of infection and prevalence of this disease among Puerto Rican frogs.
Fieldwork involving amphibian monitoring and chytrid detection at various sites
on the island (El Yunque, Patillas, Toro Negro and Maricao) suggests that chytrid
is abundant throughout the highland forests of Puerto Rico. Efforts to detect chytrid
among five amphibian species (157 individuals) at six localities in the lowlands have
yielded negative results. Our monitoring program includes four different highland
forests in Puerto Rico where we estimate frog densities by species, and sample
Conserving the Puerto Rican herpetofauna 333

for the incidence of chytridiomycosis during dry versus wet seasons, in specific
microhabitats, and as a function of gender and ontogenetic stage. An interesting
pattern of cyclic decline, survival, and recuperation of the heartiest species is
becoming apparent. This has led us to question whether a synergy between climate
and disease is playing a role in the prevalence of the fungus in Puerto Rico, as
has been described in Australia (Berger et al., 2004; Retallick et al., 2004). To
answer this question we have expanded geographical and taxonomic monitoring of
amphibian populations and chytrid detection, and have included fine-scale weather
data collection at some localities. Finally, we expect to complete historic sampling
at other localities to test disease dispersion hypotheses for the chytrid fungus in
Puerto Rico.
In 2004, with Marcelino Hernandez from the Dominican Republic, we initiated
collaborations that included a workshop on monitoring amphibians (by R.L. Joglar),
followed by several days of fieldwork in the Cordillera Central. This work resulted
in baseline population data for the localities of Ebano Verde (1440 m) and Valle
Nuevo (2500 m) and the first records of chytrid for the Dominican Republic in three
species of anurans: Eleutherodactylus pituinus, E. patriciae and Hyla vasta. Luis
M. Diaz, from Cuba, participated in this training activity, and we hope to develop
similar collaborations in the near future.

The Puerto Rican Crested Toad (Peltophryne (Bufo) lemur)

This species once occurred on Puerto Rico and Virgin Gorda. Habitat loss and
introduced species, such as Bufo marinus, are major causes for the toad’s decline
and have led to a listing as Threatened (U.S. Fish and Wildlife Service, 1992) and
Critically Endangered by the International Union for Conservation of Nature and
Natural Resources (IUCN, 2004). This is the only toad native to Puerto Rico, and
is easily distinguished from introduced forms by its unique head crests. It lives
in arid to semi-arid climates in karst limestone formations. In Puerto Rico, toad
populations once were divided into two distinct populations, one in the north and
one in the south. Mitochondrial DNA analyses suggest that these two populations
have been separated for up to 1 million years and are genetically distinct (CBSG,
2005). Unfortunately, northern toads have not been seen in the wild since 1988, and
biologists consider the population extirpated (Johnson, 1999). Currently, the only
known wild population is the southern form, which resides in small ponds located
in the Guanica National Forest (CBSG, 2005).
In an effort to save this species from extinction, a Species Survival Plan (SSP) was
officially created in 1984 through the auspices of the American Zoo and Aquarium
Association (AZA). A reintroduction program is a large component of the recovery
plan for this species. Each year, captive toads from zoos and aquaria in the United
States and Canada are bred, and tadpoles are sent to Puerto Rico for release.
Reintroduction efforts began on a small-scale, with the release of northern toadlets
in 1982, before the SSP was formed. Between 1982 and 1985, approximately 1300
334 R.L. Joglar et al.

toadlets were released in the Cambalache National Forest, in northern Puerto Rico.
Subsequently, more than 90 000 southern tadpoles were released into the Guanica
National Forest from 1987 to 2005 (Lentini, 2000; B. Johnson, pers. comm.).
The Guanica release site is an isolated man-made pond, geographically separated
from the wild population. Although no formal studies have been conducted on the
survival of the captive-hatched tadpoles, none are thought to have survived at the
Cambalache release site (B. Johnson, pers. comm.). However, in 2003 and 2005,
captive-born adult toads returned to the southern release site to breed (M. Canals,
pers. comm.), demonstrating the establishment of a second population.
Recovery goals for this species also include island-wide education outreach,
protection and restoration of existing habitat, creation of new ponds to support six
self-sustaining populations (three in the north and three in the south), and research
(U.S. Fish and Wildlife Service, 1992). Recovery efforts are primarily directed
through the USFWS, Department of Natural and Environmental Resources (DNER)
and the AZA. Additional recovery group partners consist of the University of Puerto
Rico, Juan Rivero Zoo, Ciudadanos del Karso and Inciativa Herptológica, Inc.
Restoration of existing habitat and creation of new ponds has been a continuous
process. In 1998, a secondary translocation pond was built in Guanica to serve as an
emergency refuge for tadpoles in the event of a large-scale disaster or premature
water loss at the last remaining natural breeding pond. Further, in 2000, two
ponds (4000 and 9000 sq. ft.) were built adjacent to the release site in Guanica.
Unfortunately, infiltration of saltwater has occurred at both ponds and repairs need
to be made as soon as a planned hydrology assessment has been completed. In 2005,
three small ponds were built in Arecibo (northern Puerto Rico) on private property
and northern captive-born toad tadpoles were released there in April 2006.
Future research projects will include intensive monitoring of all ponds during and
after the breeding season, with a focus on habitat use by tadpoles and newly meta-
morphosed toadlets, predation and competition, dietary studies of wild and captive
tadpoles and toads, hydrological assessments of all pond sites, amphibian monitor-
ing in Quebradillas, and a dietary and health assessment study to characterize health
threats prevalent in Bufo marinus that are cohabiting with Peltophryne lemur.
During the past twenty years, the Puerto Rican Crested Toad Recovery Program
has become a model for amphibian conservation. Much has been learned about this
elusive toad, but biologists recognize that many more questions must be answered
before this species can fully recover.

Conservation of the Puerto Rican Boa (Epicrates inornatus)

The genus Epicrates is represented by nine species in the West Indies (Tolson and
Henderson, 1993). In Puerto Rico, the genus is represented by the Virgin Island boa
(E. monensis granti), the Mona Island boa (E. m. monensis) and the Puerto Rican
boa (E. inornatus). The Puerto Rican boa is the largest snake (maximum known
SVL = 2.2 m) inhabiting the Puerto Rico Bank. This species was protected by the
Conserving the Puerto Rican herpetofauna 335

U.S. Endangered Species Act of 1973, and a recovery plan was completed in 1986.
In 2004, the Department of Natural and Environmental Resources changed its status
to Vulnerable, but it was still considered Endangered by the U.S. Fish and Wildlife
Service.
The first conservation effort was to protect boas from being collected and hunted
(USFW, 1986). More than three decades after its designation as endangered, the
ecology and natural history of these snakes still are not well documented. Thus,
different conservation strategies are being developed. Recent conservation strategies
can be categorized as: (a) gathering of basic biological information; (b) habitat
conservation; and (c) outreach.
Biological information available consists of studies of distribution, habitat pref-
erences, and life history in the Caribbean Natural Forest (Reagan and Zucca, 1982),
diet and foraging behavior (Rodríguez-Durán, 1996; Puente-Rolón and Bird-Picó,
2004; Rodriguez and Reagan, 1984; Wiley, 2003), spatial ecology and habitat use
(Puente-Rolón and Bird-Picó, 2004; Wunderle et al., 2004) and captive breeding
(Bloxam, 1981). Also, surveys for locations of unknown populations have been con-
ducted (Bird-Picó, 1994). Current research on the species focuses on habitat pref-
erences and requirements, reproductive ecology, survivorship and thermoregulatory
behavior. The goal is to contribute scientifically sound management recommenda-
tions to the development of a recovery plan for the boa.
Habitat loss and landscape fragmentation have become another concern in the
conservation of this species. Habitat destruction is increasing, and may disrupt
natural population dispersal and gene flow. Due to its protected status, translocation
(i.e., movement of wild individuals from one part of their range to another) has
become a common practice when boas are found in human settlements. A study on
how the translocation influences thermoregulation, movement and survivorship is in
progress. Preliminary data show that translocated snakes expand their home ranges
when compared to non-relocated individuals (Puente-Rolón, unpubl. data). Other
conservation efforts include the development of mechanisms for the identification of
potential habitat by local and federal agencies and the acquisition of forested areas
in the northern karst region by a non-profit organization known as Ciudadanos del
Karso. Preliminary landscape-level analysis of habitat fragmentation and potential
habitat available for the boas in the northern karst area has been performed by the
U.S. Fish and Wildlife Service to establish land conservation strategies.

Conservation of the Puerto Rican Freshwater Turtle (Trachemys s. stejnegeri)

We know almost nothing about the ecology of West Indian freshwater turtles
(Schwartz and Henderson, 1991). Turtles of the genus Trachemys might be the
least-studied vertebrates in the West Indies. Trachemys s. stejnegeri is the only
freshwater turtle native to Puerto Rico. Although its population status is currently
undetermined, early studies indicate potential threats to natural populations. For
instance, Rivero (1998) suggested that both a limited distribution and possible
336 R.L. Joglar et al.

small population sizes could warrant the protection of these turtles. However,
environmental agencies (i.e., Department of Natural and Environmental Resources,
and U.S. Fish and Wildlife Service) have been unable to provide legal protection
for T. s. stejnegeri due to a lack of data on the status of populations. On the
other hand, this species was classified as near-threatened in the Red List of the
International Union for the Conservation of Nature and Natural Resources (IUCN)
(Hilton-Taylor, 2000). Therefore, comprehensive studies are needed to characterize
the nature and extent of the factors threatening the long-term persistence of the
Puerto Rican freshwater turtle. Baseline data from such studies are necessary to
direct effective management and conservation efforts.
Recent efforts for the conservation of the Puerto Rican freshwater turtle (León
and Joglar, 2005) combined three main research approaches to clarify the status
of local populations: (1) a medium-term mark-recapture-release study, (2) field-
monitoring of reproductive activity (i.e., nocturnal patrolling to identify nesting
activity) and (3) field-assessment of the potential impact of introduced species (i.e.,
field identification of predatory species and exotic turtles).
Research findings provided a comprehensive description of the population ecol-
ogy and reproductive biology of the Puerto Rican freshwater turtle. Temporal pat-
terns of sightings are proportional to reproductive activity. Specifically, relative
abundance peaks during the nesting season from April to June. Presumably, females
migrate to limited nesting areas during the reproductive season and are then more
frequently sighted. Moreover, identified threats to reproductive success and recruit-
ment of early life stages include, intense egg predation by the exotic Indian mon-
goose (Herpestes javanicus), habitat alteration and establishment of exotic freshwa-
ter turtles in natural ecosystems inhabited by the Puerto Rican freshwater turtle.
We recommend two management actions based on current research. First, erad-
icating Herpestes javanicus from nesting areas is an urgent need. Eradication will
have a direct positive effect on the recruitment of early life stages. Second, enforce-
ment of exotic trade laws by local agencies could help in phasing out illegal sales
of the red-eared slider (Trachemys scripta elegans). The exotic red-eared slider has
been released and is widespread in Puerto Rican wetlands inhabited by native tur-
tles. This observation calls for stopping the introduction of exotic turtles in West
Indian ecosystems due to potential detrimental effects of interspecific competition
and hybridization in local turtle populations. For instance, recent massive introduc-
tions of T. scripta elegans into Europe have had negative impacts on the survival
rates of the European pond turtle (Cadi and Joly, 2004).

Recovery Initiatives for the Mona Island Iguana (Cyclura stejnegeri)

Iguanas of the genus Cyclura are arguably the most endangered lizard in the world
(Alberts, 1999, 2004). Their distribution is limited to the West Indies, with species
or subspecies typically restricted to few or single islands or cays. Cyclura stejnegeri
is found only on Mona Island, with estimated population densities ranging from
Conserving the Puerto Rican herpetofauna 337

0.33 individuals/ha (Wiewandt, 1977) to 0.96 ± 0.47 individuals/ha (Pérez-Buitrago

and Sabat, 2000). These densities are considered very low for this group (Iverson,
1977). Furthermore, a major concern has been the scarcity of immature iguanas
(5-10%) in the wild population (Wiewandt, 1977; Pérez-Buitrago and Sabat, 2000).
The Mona Island iguana has been listed as threatened and a recovery plan
approved (Diaz, 1984). Since then, we have implemented several recovery actions
aimed at the management of invasive mammals, a headstart program for hatchling
iguanas, and the assessment of the etiology of a condition causing blindness in
several adult iguanas.
Feral goats and pigs have been controlled for decades on Mona Island using
hunters, with a reported total of 314 goats and 49 pigs killed every hunting season
from January to April (García et al., 2000). Since estimated population numbers
of these animals are unknown, we cannot determine satisfactorily the efficacy of
this management strategy. However, hunting success has not changed significantly
throughout the years (García et al., 2000); this suggests a stable prey population
since feral goat populations increase rapidly in island ecosystems (e.g., Rudge et
al., 1970; Baker and Reeser, 1972). On the other hand, the most important iguana
nesting sites found on the coastal terrace and in two of the interior forest depressions
have been fenced to protect them from trampling and feral pig predation. From 1996
to 2005, intensive hunting and trapping of feral cats have been conducted on Mona
Island (García et al., 2001; López and García, unpubl. data). Although we have
eliminated a total of 118 cats, we seek to eradicate this predator. The feral cat diet
was composed of insects, small mammals, birds and reptiles, including Mona Island
iguana hatchlings. The eradication of feral cats in Mona Island is an attainable goal,
but more efforts need to be devoted to the hunting activity and the introduction of a
feline-specific virus may be necessary.
Headstarting is a conservation initiative aimed at increasing the survival of
hatchlings and juveniles by keeping them in captivity until they reach a size at
which they are protected from early age-class predators. The headstart program for
the Mona Island iguana started in 1999 in response to Perez-Buitrago (2000) finding
only 13% hatchling survival during the first five months. For the headstart program,
hatchlings were collected and kept in captivity under natural conditions until they
reach a target size of 25 cm and 950 g. To date, 132 headstarted iguanas have been
released into the wild. They are showing adequate survival (40%) and adaptation.
Although this is a conservative value because only individuals that are found are
considered, it agrees with other headstarted iguana survival values (9–40%; Alberts
et al., 2004; Wilson et al., 2004). Therefore, the implementation of the headstart
program for the Mona Island iguanas seems to be an appropriate conservation
strategy for augmenting the number of juveniles in the wild population.
Several blind iguanas had been observed in Mona Island (Haneke, 1995). There-
fore, health screening to assess this condition was conducted in 1998 and 1999 by
staff from the Toledo Zoo. Nine of 34 iguanas examined in the field (26%) were
blind or partially blind in at least one eye, due mainly to cataracts (Hudson and Al-
338 R.L. Joglar et al.

berts, 2004), suggesting that the blindness was an age-related condition (Reichard,
Tolson and García, unpubl. data). However, subsequent fieldwork (2000-2005) has
determined that 28% of iguanas found (n = 160) were totally or partially blind
(Pérez-Buitrago, unpubl. data). Therefore, a comprehensive and conclusive assess-
ment of this condition is urgently needed.
In conclusion, we have in a relatively short period been able to address the major
threats to this species. In addition, intensive ecological monitoring and research
are being conducted (Pérez-Buitrago, unpubl. data). These data will be useful to
assess the progress of the Mona Island Iguana Program. Thus, we are confident of
the recovery of this species within a decade — if these management strategies are
maintained or (preferably) intensified.

Recovery of the Herpetofauna in a Restored Karst Valley in Puerto Rico

The main goals of a habitat restoration project are to reestablish the original
biota and natural ecological functions (Parker, 1997; Palmer et al., 1997; Zedler,
2001; Block et al., 2001). Studies that have monitored the recovery of animal
communities in habitat-restoration projects, however, have focused mainly on
birds, mammals, and macro- and microfauna in soils; the herpetofauna is rarely
considered, particularly on tropical islands. On Caribbean islands, which lack the
large mammals present in the mainland tropics, reptiles and amphibians constitute
the top predators (Thomas and Kessler, 1996), and are the most abundant and
conspicuous vertebrates (Reagan, 1996; Duellman, 1999b). Given the increased
evidence of population declines in amphibians and reptiles worldwide (Pough
et al., 2001), largely due to habitat destruction, we need to develop effective
habitat restoration projects capable of recovering and maintaining the Caribbean
herpetofauna.
As an attempt to recover a local Puerto Rican herpetofaunal assemblage, we
reforested a limestone valley in Sabana Seca, Toa Baja, Puerto Rico. The study
area is in the Subtropical Moist Forest life zone (Ewel and Whitmore, 1973).
Average annual rainfall is 1700 mm. Rainfall is mildly seasonal, with most of
the rain falling from May to November (Eusse and Aide, 1999). A similar annual
pattern exists for temperature, with warmer months between May and November.
In January 2000, a deforested valley (160 × 20 m) was reforested with 516 plants
of 22 native woody species for a final density of 1666 plants/ha. In April 2002,
plant survivorship was 93.6% and mean plant height was 1.78 m. We studied four
reference sites, which were within 500 m of each other, and which consisted of
a deforested valley that represented the pre-reforested conditions, another valley,
which was reforested naturally approximately 30 to 40 years ago, and a karst
hillside and hilltop, which have been forested for >65 years. At all sites, we
sampled the species richness, monitored changes in community assemblages during
a 12-month period, and collected data on vegetation structure and microclimate.
Although the main objective for the reforestation of this valley was to create a
Conserving the Puerto Rican herpetofauna 339

forested habitat for the endangered Puerto Rican boa (Epicrates inornatus), we
wanted to evaluate the success of this reforestation project in recovering an entire
herpetofaunal assemblage.
We found a total of 17 species; the karst hillside had the highest number of
species (12), whereas the deforested valley had the lowest (2). In the reforested
valley, the herpetofaunal species richness increased rapidly from three to eleven
species, but the herpetofaunal assemblage composition recovered more slowly when
compared to the reference sites. In response to increases in the population sizes
of colonizers (e.g., Eleutherodactylus spp. and Anolis spp.), forest interior and
predatory species colonized the reforested valley by the end of the study. Among
the predatory species were the Puerto Rican boa (E. inornatus), the racer (Alsophis
portoricensis) and the arboreal giant anole (Anolis cuvieri). By planting native trees,
we increased vegetation cover and heterogeneity. The increase in woody species
changed the microclimate, specifically the variation in temperature and humidity,
and conditions are converging on the conditions of references sites. These changes,
along with the short distance to intact forest (species sources), have contributed to
the rapid recovery of herpetofaunal species richness, assemblage composition and
trophic structure.

Puerto Rican Marine Turtle Monitoring Programs

The first marine turtle studies in Puerto Rico started in 1974 on Mona Island
(Pinto-Rodríguez, 1991). Since then several marine turtle monitoring programs have
been developed or continued on Mona Island, Vieques, Culebra, Luquillo, Fajardo,
Maunabo, Rincón and Piñones. One of these programs stands out because of its
long time span, large turtle population and publication record: The hawksbill turtle
monitoring program at Mona Island.
Marine turtles were studied on Mona Island by Jean Thurston and Thomas
Weiwandt in 1974, Molly Olson in 1984 and Anastasia Kontos from 1985 to 1987.
These studies concluded that beach erosion and nest predation by pigs were the
major threats to Eretmochelys imbricata nesting on the island (Pinto-Rodríguez,
1991). These studies gave origin to management practices on Mona Island from
1989 to 1990 that included fencing the beach from Playa Mujeres to Sardinera to
prevent nest predation by pigs. A second phase of marine turtle monitoring was
initiated on Mona in 1989 by Carlos E. Diez (DNER) and Robert P. van Dam.
Results of their wok have shown that: (1) Mona and Monito Islands are among the
few known remaining locations in the Caribbean where hawksbill turtles exist in
considerable densities; (2) the large juvenile population of hawksbill turtles around
Mona and Monito consist of long-term residents that exhibit strong site fidelity
for periods of at least several years; (3) Mona’s hawksbill resident population is
composed of individuals from multiple nesting populations in the Wider Caribbean
(evidence from genetic data); (4) the conservation of the juvenile population of
hawksbill turtles at Mona can contribute to sustaining healthy nesting populations
340 R.L. Joglar et al.

throughout the Caribbean; and (5) Mona Island’s hawksbill population is the largest
under U.S. jurisdiction, and the second largest in the Caribbean, after Barbados
(Bowen et al., 1996; Possardt et al., 2007; Diez and van Dam, 2002; van Dam and
Diez, 1996, 1997a, 1997b, 1998a,b, 1999). Management practices such as beach
fencing in Mona and other conservation measures in different areas of the Caribbean
seem to be having a positive effect on nest numbers (Robert van Dam, pers. comm.).
There has been a significant increase in the number of hawksbill nests in the last
decade: 450 to 500 nests between 1997 and 2001; 850 to 925 nests between 2002
and 2004; and over 1000 nests between 2005 and 2006 (Carlos Diez, pers. comm.).
There are several additional marine turtle monitoring programs active in Puerto
Rico. Unfortunately their results and findings have not been published in peer
reviewed journals, making it difficult to review these studies. One of these programs
is the leatherback turtle monitoring program in Luquillo and Fajardo which started
in 1985. With more than 20 years of continuous monitoring, this is the longest
marine turtle monitoring program in Puerto Rico. Hector Horta and other DNER
personnel are responsible for this accomplishment. Luis Crespo directs another
marine turtle monitoring program in Maunabo.

Concluding Remarks
Conservation of the world’s natural resources and biodiversity is one of the greatest
challenges humanity faces at present. The obstacles encountered by conservation
biologists and environmentalists in Puerto Rico are not very different from those
elsewhere. However, among our Caribbean neighbors, we may be distinguished
by having the highest population density, level of industrialization, and rate of
urban development. Thus, the threats of habitat loss and contamination are of
more immediate concern. Unfortunately, an abysmal difference remains between
the funds and efforts allocated to ‘economic development’ and those dedicated to
the conservation of nature. Finding a balance that will allow for true sustainable
yield is the key to preserving our planet. However, this will entail major changes at
all levels of our societies (politics, economy, environmental policy, etc.), but most
importantly, such changes will require modifications of human conduct. The latter,
we can only achieve through education. Teaching the general public the value of
our biodiversity is imperative, for people will only strive to conserve that which
they consider significant.

Author contributions. All authors after the first are listed alphabetically. RLJ
conceived and designed the manuscript. Specific topics were written by the fol-
lowing authors: (1) Introduction, Puerto Rican herpetofauna, conservation, Puerto
Rican marine turtle monitoring programs and appendices — RLJ; (2) Amphibians,
pathogens and climate change — PAB, RLJ and AVL; (3) Puerto Rican crested
toad — DB; (4) Puerto Rican Boa — AP; (5) Puerto Rican freshwater turtle —
Conserving the Puerto Rican herpetofauna 341

ALC and RLJ; (6) Mona Island iguana — MAG, NPB, AOA and PJT; and (7) her-
petofaunal community — NRL and TMA. RLJ and PAB edited the manuscript.

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Accepted: August 29, 2007 (BSW).

Appendix. Amphibians and reptiles in Puerto Rico (#: introduced species in

Puerto Rico).
Bufonidae
Peltophryne lemur Bufo marinus #
Leptodactylidae
Eleutherodactylus antillensis E. brittoni E. cochranae
E. cooki E. coqui E. eneidae
E. gryllus E. hedricki E. jasperi
E. karlschmidti E. locustus E. monensis
E. portoricensis E. richmondi E. unicolor
E. wightmanae E. juanriveroi Leptodactylus albilabris
Ranidae
Rana catesbeiana # Rana grylio #
Hylidae
Hyla cinerea # Osteopilus septentrionalis # Scinax rubra #
 Conserving the Puerto Rican herpetofauna 345

Testudines
Caretta caretta Chelonia mydas Dermochelys coriacea
Eretmochelys imbricate Lepidochelys olivacea Trachemys scripta #
Trachemys s. stejnegeri
Gekkonidae
Hemidactylus brookii Hemidactylus mabouia Phyllodactylus wirshingi
Sphaerodactylus gaigeae S. klauberi S. levinsi
S. macrolepis S. micropithecus S. monensis
S. nicholsi S. roosevelti S. townsendi
Teiidae
Ameiva alboguttata A. desechensis A. exsul
A. wetmorei
Scincidae
Mabuya mabouya
Anguidae
Diploglossus pleei
Polychrotidae
Anolis cooki A. cristatellus A. cuvieri
A. desechensis A. evermanni A. gundlachi
A. krugi A. monensis A. occultus
A. poncensis A. pulchellus A. roosevelti
A. stratulus
Iguanidae
Cyclura nubila # C. stejnegeri Iguana iguana #
Amphisbaenia
Amphisbaena bakeri A. caeca A. schmidti
A. xera
Typhlopidae
Typhlops granti T. hypomethes T. monensis
T. platycephalus T. richardi T. rostellatus
Boidae
Epicrates inornatus E. monensis
Colubridae
Alsophis portoricensis Arrhyton exiguum
Crocodylia
Caiman crocodilus #
Extinct reptiles
Anolis rooselvelti extinction circa 1930
Cyclura pinguis old extinction (>500 years) in Puerto Rico; extant populations in the BVI
Leiocephelus etheridgei old extinction (>500 years)
Leiocephelus partitus old extinction (>500 years)
Monachelys monensis old extinction (>500 years)

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