(DRAFT) - Taxonomy
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
TAXONOMY
NAME - COQUI, GOLDEN
OTHER COMMON NAMES - COQUI and GOLDEN; COQUI DE ORO; COQUI AUREO
ELEMENT CODE -
CATEGORY - Amphibians
PHYLUM AND SUBPHYLUM - CHORDATA,
CLASS AND SUBCLASS - AMPHIBIA,
ORDER AND SUBORDER - ANURA,
FAMILY AND SUBFAMILY - LEPTODACTYLIDAE,
GENUS AND SUBGENUS - ELEUTHERODACTYLUS,
SPECIES AND SSP - JASPERI,
SCIENTIFIC NAME - ELEUTHERODACTYLUS JASPERI
AUTHORITY -
TAXONOMY REFERENCES -
COMMENTS ON TAXONOMY -
Golden Coqui
Eleutherodactylus jasperi Drewry and Jones, 1976
KINGDOM: Animal GROUP: Amphibian
PHYLUM: Chordata CLASS: Amphibia
ORDER: Anura FAMILY: Leptodactylidae
The Golden Coqui (Eleutherodactylus jasperi) is an arboreal frog
easily distinguishable from other Puerto Rican Eleutherodactylids by
its unique appearance and unusual reproductive biology. Small in size
(16.8-22.4 mm snout-vent length) the adults are sexually monomorphic.
Dorsal surfaces of head, body and limbs are smooth and lack pigmented
markings. Skin color ranges from olive to pale yellow underlain by a
translucent golden cast. The ventral skin of the abdomen is
transparent, revealing the silver-white peritoneum, gonads and heart.
Subdermal glands on dorsal surfaces of hind legs, trunk and parotoid
area are usually visible as indistinct lighter patches. Both hands
and feet have vestigial webbing between digits. Adults lack
prevomerine teeth. Oviducal young have a small well keratinized,
bifurcated egg tooth that sloughs off at about the time of birth.
Fertilized internally (10,11), females carry developing eggs in a
modified oviduct. Reproduction is ovoviviparous and the development
Taxonomy - 1 (DRAFT) - Taxonomy
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
of young is direct (young are fully metamorphosed at hatching and
subsequent birth). Young closely resemble adults in coloration and
morphology (12).
The reproductive character of the golden coqui initially
suggested the identification of a new genus, as with the African toad
(Nectophrynoides vivipara) which is similarly ovoviviparous (08). The
golden coqui is left in the present generic boundaries due to early
evidence of morphological and chromosomal affinities to other Puerto
Rican Eleutherodactylus (07,02). The species was named for Dr. Jasper
J. Loftus-Hills of New Victoria, Australia, who helped to collect the
first specimen, but died in a tragic accident a few months later.
The following specimens are available for species identification
(04):
Holotype - Field Museum of Natural History 196846, an adult male from
Sierra de Cayey, 6 km SE Cayey, Puerto Rico. Collected while calling
22 May 1973 by G.E. Drewry, K.L. Jones, J.J. Loftus-Hills and J.R.
Clark.
Paratypes - 16 adult specimens were examined. The 16 paratypes, and
the institutions housing them are:
The American Museum of Natural History - AMNH 90247-49;
The Field Museum of Natural History - CNHM 196944-45;
The Museum of Comparative Zoology, Harvard Univ. - MCZ 88388-90;
The National Museum of Natural History - USNM 198056-58;
The Los Angeles County Museum - LACM 114386;
The Museum of Southwestern Biology, Univ. of NM - UNM 27955.
Taxonomy - 2 (DRAFT) - Status
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
STATUS
Coded Status
Puerto Rico; State Listed
T: Federal Threatened
Commercial
Non-consumptive recreational
Ceremonial/Cultural
Medicinal
Ornamental
COMMENTS ON STATUS -
U.S. STATUSES AND LAWS:
The golden coqui (Eleutherodactylus jasperi) has been designated
a Threatened species pursuant to the Endangered Species Act of 1973
(50 CFR 17.11; P.L. 93-205, 87 Stat. 884; 16 U.S.C. 1531-1540),
as amended. The species has this status wherever found including
the Commonwealth of Puerto Rico. Critical Habitat has been designated
in Puerto Rico (50 CFR 17.95(d)). Three areas have been designated as
Critical Habitat: Cerro Avispa - elevations above 700 meters on the
south and southeastern slope of the mountain, Monte el Gato - entire
summit above 700 meters, and Sierra de Cayey at elevations above 700
meters. No land management recommendations are currently in effect.
This species is protected by the Lacey Act (P.L. 97-79, as
amended; 16 U.S.C. 3371 et seq.) which makes it unlawful to import,
export, transport, sell, receive, acquire, or purchase any wild
animal (alive or dead including parts, products, eggs, or offspring):
(1) in interstate or foreign commerce if taken, possessed,
transported or sold in violation of any State law or
regulation; or
(2) if taken or possessed in violation of any U.S. law,
treaty, or regulation or in violation of Indian tribal law.
It is also unlawful to possess any wild animal (alive or dead
including parts, products, eggs, and offspring) within the U.S.
territorial or special maritime jurisdiction (as defined in
18 U.S.C. 7) that is taken, possessed, transported, or sold in
violation of any State law or regulation, foreign law, or Indian
tribal law.
RESPONSIBLE FEDERAL AGENCIES:
USFWS -Responsible for the management/recovery, listing, and
law enforcement/protection of this species.
All Federal agencies have responsibility to ensure that any
action authorized, funded, or carried out by that agency is not likely
to jeopardize the continued existence of the species or result in the
destruction or adverse modification of Critical Habitat (50 CFR 402),
and to utilize their authorities to carry out programs for the
conservation of the species.
Status - 1 (DRAFT) - Status
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
STATE STATUSES AND LAWS:
STATE: Commonwealth of Puerto Rico
DESIGNATED STATUS: Endangered with Critical Habitat
ADMINISTRATIVE AGENCY: Department of Natural Resources,
Commonwealth of Puerto Rico.
STATE STATUTES: Organic Act, Sec. 155; and Wildlife Act, Sec. 85
and 91. Regulation to Govern the Management of
Threatened and Endangered Species in the
Commonwealth of Puerto Rico, 1985.
INTERNATIONAL STATUSES, TREATIES, AND AGREEMENTS:
None.
ECONOMIC STATUSES:
The unique reproductive biology and photogenic coloration of the
golden coqui will possibly result in the over collection of the
species for scientific, educational, and display purposes. The
species may also be sought for observation and photographic purposes.
Similarly bright colored frogs from the tropics, Amphodus auratus from
Trinidad and Atelopus varius zeteki from Panama, have suffered from
over collection and are close to extinction. Historically, this
species has had medicinal value in cultural ceremonies.
77/04/05:42 FR 18106/18108 - Proposed Threatened with Critical Hab.
77/11/29:42 FR 58755/ - Listed as Threatened w/ Critical Habitat
82/12/27:47 FR 42387/42388 - Five year review
87/07/07:52 FR 25523/25528 - Notice of review
Status - 2 HABITAT ASSOCIATIONS
HABITAT - TERRESTRIAL
TERRESTRIAL
LAND USE -
Residential
Transportation, communications, and Util
Cropland and Pasture
Mixed Forest Land
Bare Exposed Rock
COMMENTS ON HABITAT ASSOCIATIONS -
The golden coqui is found in a small restricted range south of
Cayey, Puerto Rico. The life zone is characterized as a subtropical
moist forest zone, receiving 1100 to 2200 mm of rainfall annually with
a mean biotemperature between 18 and 24 degrees C. Most of available
moisture, particularly during the dry season, comes from heavy dew
condensation produced at night (09,04,07).
Large areas of land have been deforested to accommodate
cultivation of crops and pasture land. Scattered areas of remnant
zonal associated vegetation are characterized by trees up to 20
meters. Many of the woody species are deciduous during the dry season
and epiphytes are common, but seldom completely cover branches and
trunks. Natural and improved pastures are common in this zone today
(01,05). Within the range of the golden coqui, there is a great
demand for high elevation land because of moderate rainfall and
temperatures. These conditions have encouraged large scale
modification of summit areas for residential development and mass
cut-slash and burn deforestation of natural vegetation for
agricultural and pasture purposes (05). With the exception of a
highway right-of-way, most of the known habitat of the golden coqui is
privately owned.
The critical factor that appears to determine the distribution of
this species within its range is the presence of large bromeliad
clusters. Bromeliads inhabited by golden coquis are found on faces of
rock cliffs, edges of short grass pastures, low-canopy forests (1-5
meters) and xeric scrub, isolated large trees, and margins of forest.
Bromeliads utilized include Vriesia, Hoenbergia, Guzmania, and
Tilanzia. Colonization of bromeliad clusters seems to be independent
of the bromeliads' substrate (04,07,03). No information is available
describing further specific habitat associations.
Drewry and Clark (04) have postulated that the distribution
limits of the golden coqui are due not to elevation but to a zone of
aridity (09,04) that acts as a barrier to dispersion. This zone is
formed by prevailing easterly winds that prevent orographic rain cloud
formation. At the edge of this zone, wind velocities are low and
heavy dew is produced every night, favoring the growth of epiphytic
vegetation.
Both distribution and abundance of the golden coqui within its
range are greatly affected by habitat disturbance. This is attributed
to the slow recolonization of bromeliads by the frogs in the areas
that have been deforested for agricultural or developmental purposes.
About half of the land area on Monte el Gato and Cerro Avispa consist
of unrecolonized pastures and lawns. No golden coquis have been
found in fairly extensive bromeliad growth at the summit of Cerro
Planada and the adjacent peak to the west. These areas have long been
planted in coffee and citrus - the planting probably accompanied by
clearing of natural vegetation by slash and burn methods.
Very little information is available describing habitat
Habitat Associations - 1 associations of the golden coqui. This information is critical in
order that those elements necessary for the conservation of the
species can be identified. This is important not only to conserve the
species within its present range, but also to explore areas for
unknown populations.
Habitat Associations - 2 (DRAFT) - Food Habits
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
FOOD HABITS
TROPHIC LEVEL -
CARNIVORE
LIFESTAGE FOOD FOOD PART
General Arthropods
Food Habits - 1 (DRAFT) - Environment Associations
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
ENVIRONMENTAL ASSOCIATIONS
G = General A = Adult
LIM = Limiting RA = Resting Adult
J = Juvenile FA = Feeding Adult
RJ = Resting Juvenile BA = Breeding Adult
FJ = Feeding Juvenile P = Pupae
L = Larvae E = Egg
RL = Resting Larvae
FL = Feeding Larvae
LIFESTAGE ENVIRONMENTAL ASSOCIATIONS
G Coastal Features: Vegetated offshore islands
G Terrestrial Features: Cliffs/ledges
G Terrestrial Features: Rock outcrops
Environment Associations - 1 (DRAFT) - Life History
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
LIFE HISTORY
FOOD HABITS:
Very little information is available describing foraging behavior
and type of food required by the golden coqui. It is assumed that the
frog forages on small insects - particularly those insects attracted
to the water in the axils of the bromeliad (02). Only a few
observations have been made of this species foraging on plants away
from the bromeliad. This is in contrast to observations of the
foraging behaviors of the parapatric coquis, E. gryllus and E.
cochranae. At night these frogs emerge to forage widely over the tree
or shrub bearing the bromeliad, and on adjacent vegetation (03).
Drewry, Jones and Clark (03,07,02) believe that most of the
frogs' needs are met within a single bromeliad or bromeliad cluster.
This is interesting in light of groups of coquis - multiple adults and
juveniles of different ages - utilizing a single bromeliad cluster.
Studies on food-resource partitioning of the golden coqui are needed
to understand food items essential to its survival.
HOME RANGE/TERRITORY:
Drewry and Clark (03) and Drewry (14) describe interactions among
caged individuals or of frogs experimentally transferred in the field.
Adult golden coquis are strongly territorial against other adults of
their own kind and other small frogs. They defend their axil fiercely
against all other frogs, shoving, biting and, in the case of males,
calling before each attack. No more than one adult was typically
found in a small bromeliad, and no more than two of opposite sex in a
large bromeliad, spaced even then as far apart as possible. In
contrast the young were usually found in leaf axils near an adult,
presumably their mother. Two or three size classes of young were
often present. This is unlike other members of the genus
Eleutherodactylus in which young normally disperse from the adult
within 5 days of hatching (12,10). A "normal" group is 1 or 2 adults
and 2 or more size classes of juveniles per bromeliad, but almost
never more than 1 frog per axil. The group or "social" relationships
of the golden coqui remain to be investigated in greater detail.
PERIODICITY:
The golden coqui appear to conform to primarily nocturnal cycles
of behavior as is typical of the genus. In daylight, the frogs sit
quitely with vent just touching the water in an axil. From this
position they capture insects entering the axil or slide backward
under the water if a larger intruder appears. At night they venture
further out on the same leaf or to others nearby (03). Drewry (02)
described the seasonal vocalization pattern as occurring throughout
the spring (early March) through late fall (November and December) and
typically climaxing in intensity during the early summer (June).
Gravid females were collected from April to August. The immature
coquis of 2 or more different size and age groups suggest that these
frogs have multiple clutches each season.
MIGRATORY PATTERN:
Non-migratory.
Life History - 1 (DRAFT) - Life History
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
COVER/SHELTER REQUIREMENTS:
The golden coqui is an obligate bromeliad dweller utilizing the
security of the bromeliad axils by day and the leaves as a foraging
substrate at night. Colonization of bromeliad clusters seems to be
independent of ground, tree, or vertical rock surface placement of the
bromeliad (07). Drewry and Clark (03) rank substrates by frequency,
the most frequently encountered substrates are rock cliffs and low
canopy xeric scrub.
REPRODUCTIVE SITE REQUIREMENTS:
No specific information is available. It is presumed that since
adults were rarely observed outside of a bromeliad, amplexus and birth
of young occur either in the water filled bromeliad leaf axils or on
the leaf itself (02).
REPRODUCTIVE CHARACTERISTICS:
The golden coqui is a terrestrial breeding frog. The males call
from bromeliads from March through December (02). Fertilization of
the females is internal (11,12) and gravid females have been collected
from April to August. The eggs are heavily yolked and number from 1
to 6 (04,12). The female carries the developing eggs in a modified
fold of the oviduct during the entire gestation period of
approximately 1 month (04,10,12). Intraoviducal nutrition is
dependent on the yolk throughout development (12). Eggs undergo
direct development and hatch as tiny froglets. The young remain with
an adult for an undetermined amount of time. Observation of 2 or more
size classes of subadults in the same bromeliad with an adult suggests
that a female may reproduce more than 1 time per year (03,12).
PARENTAL CARE:
No indication of parental care has been demonstrated after the
hatching of immature golden coquis. Juveniles of this species less
than half grown - sometimes of more than 1 age and size group - were
always found in proximity to adults. This observation is in contrast
to other Eleutherodactylus species in which hatchlings disperse from
the adults within a few days of hatching (10,11). Lack of dispersion
in juvenile golden coquis has been attributed to the semi-xeric
environment rather than to parental care.
POPULATION BIOLOGY:
Very little demography information is available for the golden
coqui. Between May 1973 and August 1974, Dr. G. Drewry estimated the
following populations based on census' of vocalizations (02):
Cerro Avispa - less than 10;
Monte el Gato - 500-1000;
Sierra de Cayey - 1000-2000.
These population estimates are the only figures available.
SPECIES INTERRELATIONSHIPS:
Drewry and Clark found possible evidence of range exclusion
between the golden coqui, and 2 parapatric species of coqui - E.
gryllus and E. cochranae. Directly north and east of the outer
Life History - 2 (DRAFT) - Life History
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
boundaries of the golden coqui distribution and at elevations above
700 m, E. gryllus frequents bromeliads along the edges of tall
forests. E. cochranae is found from the lowlands to 700 m south and
west of the golden coqui range, inhabiting litter-filled axils of
bromeliads. Surveys have indicated that all 3 species of
axil-dwelling Eleutherodactylids are mutually exclusive in ranges.
Drewry (02,03) has found evidence of upward range extension of E.
cochranae in the Sierra de Cayey area. Since this area is possibly
the best habitat available (02), the spread of E. cochranae into the
golden coqui range could be a significant problem in the recovery of
the species.
OTHER LIFE HISTORY DESCRIPTORS:
None.
Life History - 3 (DRAFT) - Management Practices
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
MANAGEMENT PRACTICES
RESULT MANAGEMENT PRACTICE
Beneficial Suppressing wildfire
Beneficial Prescribed/controlled burning of habitat
Beneficial Maintaining undisturbed/undeveloped areas
Beneficial Restricting/regulating human use of habitats
Beneficial Land Acquisition
Beneficial Stocking captive-reared wild-strain animals
Beneficial Transplanting wild animals
Beneficial Restricting Poaching
Beneficial Regulating commercial harvest levels
Beneficial Controlling/Removing Native Vertebrates
Adverse Collecting
Existing Collecting
Adverse Inherent Reproductive Characteristics
Existing Inherent Reproductive Characteristics
Adverse Low Gene Pool
Existing Low Gene Pool
Adverse Rural Residential/Industrial Areas
Existing Rural Residential/Industrial Areas
Adverse Highway/Railroads
Existing Highway/Railroads
Adverse Competition
Existing Competition
Adverse Grazing
Existing Grazing
Adverse
Existing
Adverse Fire
Existing Fire
Adverse Harvesting
Existing Harvesting
COMMENTS ON MANAGEMENT PRACTICES -
The golden coqui (Eleutherodactylus jasperi) is very rare and
restricted in range to a small area about 5 km in diameter where it
inhabits dense bromeliad growth at elevations of 700 to 850 meters
(04). Drewry and Jones (04) indicated that the species is threatened
due to past and potential loss of habitat, its obligate bromeliad
dwelling mode of existence, its presumed low reproductive rate, the
potential for overcollection, and an apparent inability to disperse.
Within the range of the golden coqui, there is a great demand for high
elevation land because of moderate rainfall and temperatures. These
conditions have encouraged large scale modification of summit areas
for residential development and mass cut-slash and burn deforestation
of natural vegetation for agricultural and pasture purposes (05).
With the exception of highway right-of-way, most of the known habitat
of the golden coqui is privately owned. Habitat destruction clearly
demonstrates the need for land usage regulations within its range.
Fire is believed to be one of the most important threats to the
Management Practices - 1 (DRAFT) - Management Practices
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
golden coqui primarily because of the semi-xeric nature of the
vegetation. In 1973 one scrub area contained golden coquis in
bromeliads on the ground, in low trees and on surfaces of large
boulders. The population was reexamined following a fire. The only
golden coquis which survived the fire were those in bromeliads on rock
surfaces (07). Fire can have a catastrophic effect on golden coqui
populations, particularly where fire control has allowed a heavy
build-up of fuel. Appropriate fire management methods need to be
employed where practical to minimize adverse effects of fire.
The unique reproductive biology and attractive coloration are
likely to create a high demand for specimens for scientific,
educational and display purposes. It has been suggested that without
proper regulation, the fate of the golden coqui may closely parallel
that of the golden frog of Trinidad (Amphodus auratus) and the
Panamanian golden frog (Atelopus varius zeteki). Both frogs are
extremely photogenic and have highly restricted ranges. Both species
have been over collected and are close to extinction (07). Potential
for gold and tungsten mining exists in the mountains between Cayey and
Salinas, which could be a potential threat to the species (14).
Drewry and Clark found possible evidence of range exclusion
between the golden coqui, and 2 parapatric species of coqui - E.
gryllus and E. cochranae (03). Directly north and east of the outer
boundaries of the golden coqui distribution and at elevations above
700 m, E. gryllus frequents bromeliads along the edges of tall
forests. E. cochranae is found from the lowlands to 700 m south and
west of the golden coqui range, inhabiting litter-filled axils of
bromeliads. Surveys have indicated that all 3 species of
axil-dwelling Eleutherodactylids are mutually exclusive in ranges.
Drewry (02,03) has found evidence of upward range extension of E.
cochranae in the Sierra de Cayey area. Since this area is possibly
the best habitat available (02), the spread of E. cochranae into the
golden coqui range could be a significant problem in the recovery of
the species.
The current population of the golden coqui is quite restricted in
its distribution and thus susceptible to habitat destruction and mass
collection of specimens. As a result, with information provided by
Drs. George E. Drewry and Kirkland L. Jones, the species was
designated as Threatened (Federal Register 42(65):18106-18108
04/05/07), and Critical Habitat determined in the three summit areas
of Cerro Avispa, Monte el Gato and Sierra de Cayey above 700 meters.
APPROVED PLAN:
U.S. Fish and Wildlife Service. 1984. Golden Coqui Recovery Plan.
U.S. Fish and Wildlife Service, Atlanta, GA. 12 pp.
The following recommendations are included in the Recovery
Plan for the golden coqui:
1. Implement actions necessary to protect Critical Habitat such as
land useage regulations, controlling development (urban and
agricultural), land purchase, etc.
2. Establish specific habitat associations and conduct a status survey
Management Practices - 2 (DRAFT) - Management Practices
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
in each of the three Critical Habitat areas.
3. Survey similar habitats outside of the Critical Habitat area for
unknown populations, and evaluate the feasibility of establishing
other populations by transplantation of individuals.
4. Explore potential benefits of captive propagation as a fallback
plan or for stock to populate new areas distant from the current
sites.
5. Investigate potential of competitive exclusion of the golden coqui
from E. cochranae and implement necessary preventative/corrective
measures.
6. Develop a long term fire management plan to be employed where
practical to minimize adverse effects of fire. Fire can have a
catastrophic effect on golden coqui populations, particularly where
fire control has allowed a heavy build-up of fuel. Appropriate fire
management methods (such as prescribed burning or fire control when
conditions and fuel quanity are undesirable) need to be employed
where practical to minimize adverse effects of fire.
Recovery objectives have been recommended to be defined as 3
populations which are stable or expanding, with each population
containing at least 1000 adults, and long term essential habitat
protection through appropriate means.
Although it was not included in the original Recovery Plan,
Drewry (14) recommends an extensive formal survey to obtain current
status. This would be accomplished through examining all known
collection localities and other areas ecologically similar to the
collection localities. In view of the recent failure to locate
individuals of this species, plans and immediate steps to institute
captive propagation if/when individuals are found is also recommended.
The species has been maintained in captivity for periods of several
months before. The restricting of poaching for individual or
commercial purposes should be restricted in view of the rarity of this
species.
Management Practices - 3 (DRAFT) - References
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
References
***** REFERENCES FOR ALL NARRATIVES EXCEPT N-OCCURRENCE *****
01 Birdsey, R.A. and P.L. Weaver. 1982. The Forest Resources of
Puerto Rico. USDA For. Serv. Res. Bull. No. 85:59 pp. Inst. Trop.
For. Rio Piedras, PR.
02 Drewry, Dr. George E. August 26, 1983. Personal communication to
Tammie Nakamura. U.S. Dept. of Interior, USFWS, OES Washington,
D.C. 20240.
03 Drewry, G.E. and J.R. Clark. 1977. Microhabitat, competition,
climatic relationships and distribution. Part III. U.S. Dept. of
Interior, USFWS, OES Washington, D.C. 20240.
04 Drewry, G.E. and K.L. Jones. 1976. A new ovoviviparous frog,
Eleutherodactylus jasperi (Amphibia, Anura, Leptodactylidae) from
Puerto Rico. J. Herpet. 10(3):161-165.
05 Ewel, J.J. and J.L. Whitmore. 1973. The ecological life zones of
Puerto Rico and the U.S. Virgin Islands. USDA For. Serv. Res.
Paper ITF-18. 72 pp. Inst. Trop. For., Rio Piedras, PR.
06 Goin, C.J., O.B. Goin, and G.R. Zug. 1978. Introduction to
Herpetology. Freeman, San Francisco, CA. 99 pp.
07 Jones, K.L. 1975. Proposed Endangered species account for the
Golden Coqui. Personal communication to Dr. James D. Williams.
Dept. of Biology, Southern Methodist Univ., Dallas, TX 752704.
08 Noble, G.K. 1926. An analysis of the remarkable case of
distribution among the Ambhibia - with descriptions of new genera.
Amer. Mus. Novitates. 212:1-24.
09 Rivero, J.A. 1963. Notes on the distribution of some Puerto Rican
frogs, with a discussion of the possible origin of
Eleutherodactylus locustus. Carib. J. Sci. 3(2):81-85.
10 Rivero, J.A. 1978. Los anfibios y reptiles de Puerto Rico.
Editorial Universitaria. Universidad de Puerto Rico. Mayaguez,
PR. 22 pp.
11 Townsend, D.S., M.M. Stewart, F.H. Pough and P.F. Brussard. 1981.
Internal fertilization in an oviparous frog. Science 212:469-471.
12 Wake, M.H. 1978. The reproductive biology of Eleutherodactylus
jasperi (Amphibia, Anura, Leptodactylidae) with comments on the
evolution of live-bearing systems. J. Herpet. 12(2):121-133.
13 Anon. 1980. Checklist of Reptiles and Amphibians. Assoc. of
Systematic Collectors, Lawrence, KS.
14 Drewry, G.E. 1986. Personal communication. Biologist, U.S.
Fish and Wildlife Service, Washington, D.C. 20240.
***** REFERENCES FOR N-OCCURRENCE NARRATIVE ONLY *****
01 Drewry, Dr. George E. August 26, 1983. Personal communication to
Tammie Nakamura. U.S. Department of Interior, USFWS, OES.
Washington, D.C. 20240.
02 Drewry, G.E. and J.R. Clark. 1977. Microhabitat, competition,
climatic relationships and distribution. Part III. U.S.
Department of Interior, USFWS, OES. Washington, D.C. 20240.
03 Drewry, G.E. and K.L. Jones. 1976. A new ovoviviparous frog,
References - 1 (DRAFT) - References
Species COQUI, GOLDEN
Species Id ESIS204001
Date 14 MAR 96
Eleutherodactylus jasperi (Amphibia, Anura, Leptodactylidae) from
Puerto Rico. J. Herpet. 10(3):161-165.
04 Jones, K.L. 1975. Proposed Endangered species account for the
Golden Coqui. Personal communication to Dr. James D. Williams.
Department of Biology, Southern Methodist University, Dallas, TX
752704.
05 Drewry, G.E. 1986. Personal communication. Biologist, U.S.
Fish and Wildlife Service, Washington, D.C. 20240.
References - 2