• TABLE OF CONTENTS
HIDE
 Front Cover
 Copyright
 Table of Contents
 Introduction
 Acknowledgement
 Systematic descriptions
 Discussion
 Literature cited
 Appendix 1
 Appendix 2
 Appendix 3
 Back Cover






Group Title: Bulletin of the Florida State Museum Biological sciences v. 21, no. 1
Title: Variation and relationships of some Hispaniolan frogs (Leptodactylidae, Eleutherodactylus) of the ricordi group
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00095821/00001
 Material Information
Title: Variation and relationships of some Hispaniolan frogs (Leptodactylidae, Eleutherodactylus) of the ricordi group
Series Title: Bulletin - Florida State Museum Biological sciences ; volume 21, number 1
Physical Description: 46 p. : ill. ; 23 cm.
Language: English
Creator: Schwartz, Albert, 1923-1992
Publisher: Florida State Museum, University of Florida
Place of Publication: Gainesville, Fla.
Publication Date: 1976
Copyright Date: 1976
 Subjects
Subject: Eleutherodactylus -- Hispaniola   ( lcsh )
Eleutherodactylus -- West Indies   ( lcsh )
Animals -- Variation   ( lcsh )
Amphibians -- Classification   ( lcsh )
Amphibians -- West Indies   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: Bibliography: p. 41.
General Note: Cover title.
Statement of Responsibility: Albert Schwartz.
 Record Information
Bibliographic ID: UF00095821
Volume ID: VID00001
Source Institution: University of Florida
Holding Location: University of Florida
Rights Management: All rights reserved by the source institution and holding location.
Resource Identifier: oclc - 03412949
lccn - 77620825

Table of Contents
    Front Cover
        Page i
    Copyright
        Page ii
    Table of Contents
        Page 1
    Introduction
        Page 2
    Acknowledgement
        Page 3
    Systematic descriptions
        Page 4
        Page 5
        Page 6
        Page 7
        Page 8
        Page 9
        Page 10
        Page 11
        Page 12
        Page 13
        Page 14
        Page 15
        Page 16
        Page 17
        Page 18
        Page 19
        Page 20
        Page 21
        Page 22
        Page 23
        Page 24
        Page 25
        Page 26
        Page 27
        Page 28
        Page 29
    Discussion
        Page 30
        Page 31
        Page 32
        Page 33
        Page 34
        Page 35
        Page 36
        Page 37
        Page 38
        Page 39
        Page 40
    Literature cited
        Page 41
    Appendix 1
        Page 42
        Page 43
    Appendix 2
        Page 44
        Page 45
    Appendix 3
        Page 46
        Page 47
    Back Cover
        Page 48
Full Text








of the

FLORIDA STATE MUSEUM
Biological Sciences


Volume 21 1976 Number 1





VARIATION AND RELATIONSHIPS OF SOME
HISPANIOLAN FROGS (LEPTODACTYLIDAE,
ELEUTHERODACTYLUS) OF THE RICORDI GROUP

ALBERT SCHWARTZ


UNIVERSITY OF FLORIDA


GAINESVILLE









Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM, BIOxLGICAL
SCENCES, are published at irregular intervals. Volumes contain about 300 pages and
are not necessarily completed in any one calendar year.








CARTER R. GILBERT, Editor
RHODA J. RYBAK, Managing Editor


Consultant for this issue:
ERNEST E. WILIAMS














Communications concerning purchase or exchange of the publications and all manu-
scripts should be addressed to the Managing Editor of the Bulletin, Florida State
Museum, Museum Road, University of Florida, Gainesville, Florida 32611.


Publication date: Aug. 6, 1976


This public document was promulgated at an annual cost of $1647.38
or $1.647 per copy. It makes available to libraries, scholars, and all
interested persons the results of researchers in the natural sciences,
emphasizing the Circum-Caribbean region.


Price: $1.70






















VARIATION AND RELATIONSHIPS OF SOME HISPANIOLAN
FROGS (LEPTODACTYLIDAE, ELEUTHERODACTYLUS)
OF THE RICORDI GROUP

ALBERT SCHWARTZ1

SYNOPSIS: Five species of Hispaniolan Eleutherodactylus of the ricordi group are
discussed, and variation in these species is given in detail. The relationships of
these five species, both among themselves and with other Antillean members of the
ricordi group, are treated, and a hypothetical sequence of inter- and intra-island
trends is given, based upon presence or absence of digital discs and glandular areas.
Data on voice, habitat, and distribution of the included species are given.

TABLE OF CONTENTS
INTRODUCTION ------... ---................... ---- -------- ------------.---------- 2
ACKNOWLEDGMENTS AND MATERIALS -------..------............-------- 3
SYSTEMATIC DESCRIPTIONS 3-........---... --------------- ------------------- ---------- 3
Eleutherodactylus pictissimus COCHRAN 1935: 371 3------------------- -.... 3
Eleutherodactylus pictissimus pictissimus COCHRAN 1935 -..------ 4
Eleutherodactylus pictissimus apantheatus SCHWARTZ 1965: 102 ---.. 7
Eleutherodactylus pictissimus eremus SCHWARTZ 1965: 107 ---- 9
Eleutherodactylus weinlandi BARBOUR 1914: 246 -- 11
Eleutherodactylus weinlandi weinlandi BARBOUR 1914 ....----------------- 11
Eleutherodactylus weinlandi chersonesodes SCHWARTZ 1965: 115 --- 14
Eleutherodactylus weinlandi paralius new subspecies -..---.------ --------- 16
Eleutherodactylus alcoae SCHWARTZ 1971: 26 ...---- 20
Eleutherodactylus probolaeus SCHWARTZ 1965: 110 .----- 23
Eleutherodactylus warren new species --- --.---.-_-.--------.. 26
THREE ANOMALOUS SPECIMENS ......... --------------------.........-. 28
DISCUSSION -...-... .... ----------------................----- --------- --- -- 30
LITERATURE CITED --...-.....----------- ---------------- 41
APPENDIX 1 ............----------............------------- 42
APPENDIX 2 ---------------..... -....... ...------------.- -......--- ----- 44
APPENDIX 3 ...-------- ......... .-.--------------------.-- .---.------------------ 46


1 The author is a Professor in the Department of Biology, Miami-Dade Junior College, Miami,
Florida 33167. Manuscript accepted 9 September 1973.

SCHWARTZ, ALBERT. 1976. Variation and Relationships of Some Hispaniolan Frogs
(Leptodactylidae, Eleutherodactylus) of the ricordi group. Bull. Florida State Mus.,
Biol. Sci., Vol. 21(1): 1-46.







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


INTRODUCTION
The Eleutherodactylus ricordi group has a complex of species on the
West Indian islands of Cuba, Hispaniola, Mona, Puerto Rico, and the
Virgin Islands that form a compact assemblage of related forms; simi-
larities include general size, dorsal coloration and pattern, and those
characters that ally them to each other and are common to all members
of the group (Dunn 1926:210, Schwartz 1958:2). The complex of
species presently under discussion includes the Cuban species ricordi
Dumeril and Bibron, thomasi Schwartz, bresslerae Schwartz; the His-
paniolan species weinlandi Barbour, pictissimus Cochran, alcoae
Schwartz; as well as monensis Meerwarth from Isla Mona, richmondi
Stejneger from Puerto Rico, and lentus Cope from St. Thomas, St. John,
and St. Croix, Virgin Islands.
This group of species may be divided into two major subgroups:
one has pale dorsa with dark markings (to which all but two of the
above species may be assigned); the other is a pair of species (wein-
landi, richmondi) that are richly colored dorsally and lack, or have modi-
fied, dorsal dark pattern elements. Shreve and Williams (1963:332-334)
discussed the relationships between pictissimus, weinlandi, ricordi, bres-
slerae, and lentus, stating quite correctly that these forms show "sur-
prisingly few differences, even in coloration." They cogently suggested
that it might be more appropriate to regard some of the members of
the complex as subspecies (i.e., weinlandi and lentus; or pictissimus,
weinlandi, and lentus). However, for lack of other than inductive evi-
dence, they elected not to make the corresponding nomenclatural changes.
I reviewed the evidence in 1965 and retained the Hispaniolan pictissimus
and weinlandi as distinct species, although their distributions showed
some peculiarities-apparently of a mosaic nature-on Hispaniola. In
the same paper I named a new subspecies of weinlandi (chersonesodes)
from the Peninsula de SamanA and extreme eastern Republica Domini-
cana, and three new subspecies of pictissimus (apantheatus, eremus,
probolaeus). Of the four subspecies of E. pictissimus, all show adjacent
distributions or intergradation except probolaeus, which is known only
from extreme southeastern Rep6blica Dominicana and is geographically
separated from the other subspecies by a distance of about 200 km. At
that time (1965) two records of E. weinlandi were known from between
the ranges of probolaeus and the other subspecies of E. pictissimus:
Cueva de Santa Anna in the city of Santo Domingo (Mertens 1939:30);
12 km NE La Romana, La Romana Province, Repiblica Dominicana
(Schwartz 1965:119). It seemed apparent that the range of E. weinlandi
was intermediate to those of two subspecies of E. pictissimus, a rather
unusual situation.








SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


ACKNOWLEDGMENTS AND MATERIALS
In an effort to clarify the relationships of this complex of taxa on Hispaniola,
effort was made between 1968 and 1971 in both Haiti and the RepTiblica Dominicana
to secure additional specimens; this field work was under the sponsorship of National
Science Foundation grants G-7977 and B-023603. So much new material has ac-
cumulated from these collections that it is appropriate to reassess the status of the
named populations and to redefine their distributions. Through the efforts of C.
Rhea Warren a series of Eleutherodactylus was secured on Ile de la Tortue off the
northern Haitian coast; Mr. Warren's visits to that island were greatly facilitated by
the cooperation of M. Ramah Th6odore, Directeur G6n6ral Adjoint de l'Office Na-
tionale du Tourisme et de Propagande. Collections made by Richard Thomas along
the northern Haitian littoral and in the vicinity of Mirebalais have clarified some
problems but have posed others. In the field I have had the cooperation of Patricia
A. Adams, Donald W. Buden, Jeffrey R. Buffett, James R. Dennis, Danny C. Fowler,
Ronald F. Klinikowski, David C. Leber, Dennis R. Paulson, James A. Rodgers, Jr.,
Bruce R. Sheplan, William W. Sommer, James B. Strong, and Richard Thomas.
Lewis D. Ober collected an especially interesting Eleutherodactylus from Thomonde
in central Haiti.
The present paper is based primarily upon material in the Albert Schwartz
Field Series (ASFS) collected between 1962 and 1971. For my previous report
(1965) on variation in the weinlandi-pictissimus complex, I borrowed specimens
from the American Museum of Natural History (AMNH), the National Museum of
Natural History (USNM), the Museum of Comparative Zoology (MCZ), and the
Museum of Zoology, University of Michigan (UMMZ). Holotypes and paratypes
were deposited in the Carnegie Museum (CM), the Museum of Natural History,
University of Kansas (KU), and the Museum of Natural History, University of
Illinois (UIMNH). I have not re-borrowed these specimens, but data from them
are included in the lists of specimens examined and in the computations and distri-
butional analyses presented herein. I have borrowed specimens of the Cuban taxa
(bresslerae, thomasi, ricordi) from the American Museum; these specimens were
collected by me under NSF grants G-3865 and G-6252 between 1957 and 1960.
I wish to thank Charles W. Myers and George W. Foley for the loan of these speci-
mens. Not all problems of the relationships of the Antillean taxa to each other may
be presently solved, but the large quantity of new Hispaniolan material and new
data do manage to clarify these relationships to some extent and to expand our con-
cepts of the Hispaniolan taxa.
I would also like to extend my gratitude to David C. Leber for his artistic rendi-
tion of the two new taxa described in this paper.

SYSTEMATIC DESCRIPTIONS

Eleutherodactylus pictissimus COCHRAN 1935: 371

HOLOTYPE AND TYPE LOCALITY: MCZ 19846; Tardieu, Massif de la Hotte, ca 3000
ft (915 m), D6partement du Sud, Haiti.
DIAGNOSIS.-A species of the Antillean ricordi group of Eleutherodactylus char-
acterized by the combination of moderate size (snout-vent length: males to 34 mm,
females to 43 mm); dorsum brown (rarely) to pale (tan, buffy tan, yellow, pale
orange, pale gray, tannish bronze, dull orange-tan, light brown, pale tan, reddish
tan), either overlaid with a dense and very dark brown to black reticulum, with a
pair of dorsolateral stripes of the dorsal ground color ( = not contrastingly colored
when compared with the dorsal color), or pattern much reduced to fine stippling
with a prominent dark scapular chevron; hind limbs coarsely banded to vaguely
stippled with dark brown to black; inguinal and supra-axillary glandular areas present
but inconspicuous; expanded digital discs very poorly developed to absent; tympanum


1976







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


small (X=males, 2.2 mm; females, 2.7, 2.5, and 2.4 [see Appendix 1]); and tibia/
snout-vent length (X 100) moderate (males, 41.3-55.2; females, 42.0-53.3).
DISTamBUTIoN.-The Tiburon Peninsula in Haiti east to the Peninsula de Bara-
hona in the Repiblica Dominicana. It occurs within this region at high elevations
in the Massif de la Hotte (Tardieu) and the Montagne Noire (Furcy) (but is absent
from the highlands of the Sierra de Baoruco and the extremely arid Peninsula de
Barahona itself); the Cul de Sac-Valle de Neiba Plain (north of the Massif de la
Selle and the Sierra de Baoruco) east to the vicinity of Bani in the Llanos de Azua,
Repdblica Dominicana (Fig. 1). Altitudinal distribution extends from sea level (or
below, near Jimani and Duverg6) to 5800 ft (1769 m) at Furcy in the Montagne
Noire, but not higher than 1800 ft (549 m) above Barahona in the Sierra de
Baoruco.

Eleutherodactylus pictissimus pictissimus COCHRAN 1935
DIAGNOSIS.-A subspecies of E. pictissimus characterized by large size (snout-
vent length: males, 34 mm; females, 43 mm); dorsum rarely dark (brown) and
usually pale (tan, buffy tan, yellow, pale orange, tannish bronze, yellow-tan, orange-
tan, pale gray), heavily and densely covered by dark brown to black spots to form a
reticulum in many instances, but leaving a pair of dorsolateral stripes that are uni-
color with the dorsal ground color, a dark scapular chevron present but incorporated
into the dorsal pattern and not outstanding as a separate dorsal unit; sides grayish
and overlaid by the same dark dorsal pattern; hindlimbs prominent and contrast-
ingly banded, reticulate, or mottled with dark pigment, but usually without clearly
defined limb bands (if discernible, about three on the thighs and two on the crura);
concealed surfaces of femora gray; venter opalescent to whitish; and throat gray.
DISTRIBUTION.-The Tiburon Peninsula in Haiti (from Jer&mie and Moron in
the north and Les Platons and Les Cayes in the south), east to the southern slopes
of the Sierra de Baoruco (above Cabo Rojo); also on Ile-A-Vache, Haiti. Inter-
grades between pictissimus and apantheatus occur along the northern coast (and
inland to Furcy) of the Tiburon Peninsula between Qa Ira and Dufort on one hand,
and P6tionville and Port-au-Prince on the other, and along the extreme southeastern
edge of the Sierra de Baoruco near Enriquillo, Juancho, and Calet6n, Rep6blica Do-
minicana. Altitudinally it ranges from sea level (many localities, including Les Cayes,
Aquin, J6r6mie, Ca Ira) up to 1700 ft (519 m) in the Sierra de Baoruco northeast of
Cabo Rojo, and up to 3000 ft (915 m) both at Tardieu (type locality) and Les Pla-
tons in the Massif de la Hotte. Intergrading specimens (pictissimus X apantheatus)
are known at even higher elevations (5800 ft [1709 m]) from Furcy on the Mon-
tagne Noire.

REMARKS.-Of the three E. pictissimus subspecies, the nominate form
is by far the most widely distributed, both geographically and altitu-
dinally, and is also the largest. As pointed out previously (Schwartz
1965), E. p. pictissimus is the most darkly patterned of the subspecies
and lacks the faded appearance of apantheatus and eremus. The snout-
vent lengths of 119 adults are: 57 males, 23.0-33.9 mm (X 26.6); 62
gravid females, 28.5-43.2 mm (X 35.7). Additional mensural data are
presented in Appendix 1.
Considering the frog's broad distribution and its abundance through-
out much of its range, it is truly remarkable that E. pictissimus was de-
scribed so recently and from only one specimen. Although one has the
impression that E. p. pictissimus is confined to mesic situations, it ob-
















"I ,



0
"-- ?-----,--,-,-, :i-i.. iiii:.-










742 70* 0_10 05040
SK




FIGtUE 1.-Map of Hispaniola, showing localities and known distributions of three subspecies of Eleutherodactylus pictissimus as
follows: fine stippling, E. p. pictissimus; horizontal lines, E. p. apantheatus; coarse stippling, E. p. eremus. Overlapping symbols
indicate regions of intergradation. The three questioned localities represent individual specimens that are questionably associated
with E. pictissimus.







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


viously is also tolerant of much dryer regions and is 'one of the few
Antillean frogs that occurs in areas consistently exposed to salt water.
Repeated field data include E. p. pictissimus in such local situations as
piles of moist Cocos trash (both husks and fronds), rock piles, and under
individual rocks. The subspecies is associated with shaded ravines, as
at Camp Perrin, but does not shun grassy pastures (L'Eglise, Les Cayes).
Twelve were taken in a cave (Grotte de Counou Bois), 150 ft (46 m)
inside the entrance, where they were active during the morning hours
in an area of small moist stones. Above Cabo Rojo in the Sierra de
Baoruco, E. p. pictissimus occurs in Acacia woods, where it was en-
countered under rocks in dusty soil during the day and hopping about
actively at night both in Acacia and semi-mesic deciduous woods (as at
L'Eglise).
As I have pointed out (Schwartz 1965:113), E. pictissimus has never
been heard calling, despite collections made during very favorable
weather conditions and at all seasons of the year. Fowler and Sheplan
thought they heard a small, whispery call at a locality near La CiBnaga
(where E. p. apantheatus is exceptionally abundant), but they were
unable to verify their observations. I too have had the impression that
a low and indistinct call heard near L'Eglise might have been made by
E. p. pictissimus but was unable to verify the fact. In all members of
the ricordi group, the call is irregular and insect-like and easily confused
with other nocturnal noises or obscured by them. With no obvious
evidence to the contrary, I have assumed that E. pictissimus in non-vocal.
However, Shreve and Williams (1963:334), quoting field notes from
Anthony Curtis on E. pictissimus at his house in Port-au-Prince, Haiti,
stated, "It made a noise resembling the chirping of a cricket"; these
sounds were made by a frog while Curtis was holding it in his hand.
Many eleutherodactyls and other anurans make "chuckling" or other
distress calls when disturbed, and it is possible that this was the case
with Curtis' frog. As yet there is no concrete evidence that E. pictissi-
mus vocalizes under natural conditions.
Despite its broad distribution, E. p. pictissimus shows very little geo-
graphically correlated variation. Specimens from Ile-a-Vache, for in-
stance, are like those from the adjacent mainland at Les Cayes. Speci-
mens from north and south of the Massif de la Hotte also appear in-
distinguishable, although a short series from near J6r6mie (ASFS V9309-
12) has paler pigmentation and is less well marked than other specimens.
However, other frogs from the J6r6mie area are just as heavily pigmented
dorsally as the other E. p. pictissimus. There are no obvious pigmental or
pattern differences between specimens from the distal Tiburon Penin-
sula and those from the Sierra de Baoruco. It is interesting to note,







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


however, that the largest frogs are from the eastern extreme of the range
(the Sierra de Baoruco) and are consistently larger than frogs from else-
where.
SPECIMENS EXAMINED. HAITI. DiPT. DU SUD: beach area within 1 km of
J&ermie (ASFS V21291); 2 km NW J&remie (ASFS V9309-12); Place Negre nr.
J6r6mie (MCZ 33296-98); Mayette (MCZ 37571); Carrefour Sanon nr. J6r6mie
(MCZ 33818-19; MCZ 37574-85); Fond Rouge Daye nr. J6remie (not mapped)
(MCZ 37565); Bozo nr. J6r6mie (not mapped) (MCZ 37566-69); La Source nr.
J6remie (not mapped) (MCZ 37570); Perine nr. Jr6Bmie (not mapped) (MCZ
37572-73); Marfranc (MCZ 37743-49); Moron (USNM 60626); Tardieu, 3000 feet
(905 meters) (MCZ 19846); Castillon, 2500 feet (763 meters) (ASFS V24722-25,
ASFS V24971); Camp Perrin (ASFS X2526-27, ASFS X2671, ASFS X2683, ASFS
X2749-50, ASFS X2813-16, ASFS X3099-106, MCZ 33282-91); 5 mi. (8.0 km) from
Camp Perrin (MCZ 36746-47); Grotte de Counou Bois, 1 mi. (1.6 km) SW Camp
Perrin (ASFS X3254-65, ASFS X3272-73); Les Platons, 3000 feet (915 meters)
(ASFS V20410-13); Les Cayes (ASFS X3712-13, ASFS X3370-71); 1 mi. (1.6 km)
NW Les Cayes (ASFS X3822); 4.5 mi. (7.2 km) NW Les Cayes (ASFS X3802-11);
Ile-a-Vache, western end (ASFS X3553-55); 0.6 mi. (1.0 km) W Aquin (ASFS
V25900-02); Fond des Negres (MCZ 35194); Mingrette nr. MiragoAne (not mapped)
(MCZ 35195); Pemel nr. Miragoane (not mapped) (MCZ 35196); But&te nr.
MiragoAne (not mapped) (MCZ 35197-98); Paillant, 1800 feet (549 meters) (ASFS
V26218-20); 4.7 mi. (7.5 km) SW Paillant, 2500 feet (763 meters) (ASFS V26169).
DiPT. DE L'OUEST: ca. 5.5 mi. (8.8 km) NW Jacmel, 600 feet (183 meters) (ASFS
V9782-83); Ca Jaqueline nr. Jacmel (not mapped) (MCZ 34500); Thiotte (MCZ
36095-96). REPOBLICA DOMINICANA. PEDERNALES PROVINCE: 6.0 mi. (9.6 km) N
Pedernales, 800 feet (244 meters) (ASFS V30106-07); 13.0 mi. (20.8 km) N Peder-
nales, 600 feet (183 meters) (ASFS V29735); 26 km N Pedernales (ASFS V21472-
73); 20 km N Cabo Rojo, 1400 feet (427 meters) (ASFS V29710-11); 21 km NE
Cabo Rojo, L'Eglise, 1500 feet (458 meters) (ASFS V16802, ASFS V30031-37);
21 km NE Cabo Rojo, 1300 feet (397 meters) (ASFS V16795); 22 km NE Cabo
Rojo, 1700 feet (519 meters) (ASFS V16753).

Eleutherodactylus pictissimus apantheatus SCHWARTZ 1965:102
HOLOTYPE AND TYPE LOCALITY.-MCZ 43195; 6.5 mi. (10.4 km) NE Jimani, In-
dependencia Province, Rep6blica Dominicana.
DIAGNOSIs.-A subspecies of E. pictissimus characterized by moderate size
(snout-vent length: males to 33 mm, females to 35 mm); dorsum pale (tan, reddish
tan, yellow); dorsal markings black to dark brown but much reduced, usually form-
ing a fine vermiculate or reticulate pattern; dark dorsal pattern delimiting a pair of
dorsolateral stripes that are concolor with the dorsal ground color; a moderately
prominent dark scapular chevron, sides grayish, usually more heavily marked with
dark pattern elements than dorsum; hindlimbs marbled or mottled with dark pig-
ment, rarely with any banding indicated; concealed surfaces of femora whitish;
venter white to flesh; throat white.
DISTRIBUTION.-The Valle de Neiba in the Rep6blica Dominicana, from Jimani
east to the vicinity of Barahona, and thence south along the eastern coast of the
Peninsula de Barahona as far as Paraiso, and northeast to Fondo Negro. It pre-
sumably also occurs in the Haitian Cul de Sac Plain but is not yet known from that
area. Intergrades between apantheatus and pictissimus occur in two areas: in Haiti
between Ca Ira-Dufort and Port-au-Prince (and into the Montagne Noire to Furcy),
and at the extreme southeastern corner of the Sierra de Baoruco near Enriquillo,
Juancho, and Calet6n, Rep6blica Dominicana. Altitudinal distribution ranges from
sea level (localities along the eastern coast of the Peninsula de Barahona) or below
(in the Valle de Neiba at Jimani and Duverg6) to 1800 ft (549 m) near Barahona.


1976







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


REMARKS.-The large amount of additional material of E. p. apanthe-
atus confirms the pigment and pattern differences between that sub-
species and E. p. pictissimus that were noted in the original description
of apantheatus. The pale dorsal ground color, with the dorsal dark
markings reduced but still present, and the moderately conspicuous dark
scapular chevron serve to distinguish E. p. apantheatus from the nomi-
nate subspecies (see Schwartz 1965:figs. 89, 90, for illustrations of these
two taxa). In snout-vent length, males are slightly smaller than male
pictissimus (X 24.9 apantheatus, 26.6 pictissimus; largest male
apantheatus 33.1, largest male pictissimus 33.9), but female apantheatus
reach a much smaller maximum size (34.7) than do female pictissimus
(43.2), and the mean female snout-vent length is lower in apantheatus
(32.0) than it is in pictissimus (35.7). Tibia/snout-vent length (X 100)
ratios are comparable in both sexes of both subspecies (see Appendix 1
for all mensural data).
Intergrades between the two subspecies occur in two regions. In
Haiti there is a broad zone of intergradation between Ca Ira-Dufort in
the west, Furcy in the south, and Port-au-Prince in the east. Specimens
from this region are extremely variable; some are as poorly patterned
dorsally (with a prominent scapular dark chevron) as are apantheatus,
and others have the heavily patterned dorsa of western pictissimus.
The second area of intergradation lies along the southeastern edge of the
Sierra de Baoruco. In this region E. pictissimus is known from a series
of coastal localities from near Barahona in the north to Juancho in the
south. Fresh specimens from this coastal area indicate that to the south
there is a strong tendency for the dorsal patterns to become darker and
more definitive, with subsequent obfuscation of the dark scapular chev-
ron, and that specimens from Calet6n, for instance, are closer to E. p.
pictissimus than to E. p. apantheatus. There is a gap of some 45 km
between the closest records of pictissimus and the intergradient localities
to the southeast along the southern slopes of the Sierra de Baoruco, but
presumably (judging from the small lots of material from Juancho-
Calet6n-Enriquillo) these southern slopes are inhabited by the nominate
subspecies.
Since at least part of the range of E. p. apantheatus lies within the
hot and dry, and in places below sea level, Valle de Neiba (and pre-
sumably in the equally hostile Cul de Sac Plain in Haiti, whence no
material is yet available), apantheatus is much more confirmedly a local
mesophile than is pictissimus. In this desert region the holotype (from
near Jimani) was taken under a stack of moist palm fronds, and six
specimens from near Cabral were secured under palm slats adjacent to
Laguna de Rincon. Two frogs from El Iguito were taken from under








SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


moist Musa trash on a banana plantation in an otherwise extremely arid
desert. Along the eastern coast of the Peninsula de Barahona, speci-
mens have been secured under palm trash along the beach and under
Cocos trash in shaded to open cafetales (Rio Nizaito; Paraiso). The
frog is exceptionally abundant at La Ci6naga, where during the day
individuals were caught under rocks in a moist and shaded ravine bed
and at night on rocks along the bed of the stream. One individual here
was seen 6 inches (15 cm) above the ground in a fern, one of the few
instances of climbing noted for the species. One frog was taken on dry'
earth in Acacia woods southwest of Barahona, but at this locality there
were adjacent stands of moderately-shady, semi-xeric hardwoods mixed
with Acacia.
A pair of E. p. apantheatus was found in amplexus in a collecting
sack; the series of which this pair was a part had been night-collected
under extremely rainy conditions. There was a marked size difference
between the two sexes, with the male much the smaller, and the position
of the male was axillary.

SPECIMENS EXAMINED.-REPtOBLICA DOMINICANA. INDEPENDENCIA PROVINCE:
6.5 mi. (10.4 km) NE Jimani (MCZ 43195); 6.5 km W Duverg6 (ASFS V4417).
BARAHONA PROVINCE: El Iguito, 1.6 mi. (2.6 km) NE Fondo Negro (ASFS V30464-
65); 4.7 mi. (7.5 km) E Cabral (ASFS X9615-20); El Proprio Esfuerzo, del
Monte's finca, 1800 feet (549 meters) (AMNH 44523); 5 km Barahona (ASFS
V20589); 4.1 mi. (6.6 km) SW Barahona (ASFS V30406); 3.3 mi. (5.3 km) NE
La Ci6naga (KU 79807-10, USNM 150927-30, AMNH 71977-81, UIMNH 55587-
90, ASFS V30455, ASFS V30588-615, ASFS V30803); 1.9 mi. (3.0 km) W Paraiso,
600 feet (183 meters) (ASFS V30834); 1 km NE Paraiso, Rio Nizaito (ASFS
V16947-51, ASFS V30436-37); 2 km SW Paraiso (CM 38973-78).
Intergrades between E. p. pictissimus and E. p. apantheatus. REPOBLICA
DOMINICANA. BARAHONA PROVINCE: 0.5 mi. (0.8 km) NE Calet6n, 400 feet
(122 meters) (ASFS V30742-47); Enriquillo (AMNH 44677-78). HAITI. DEPT.
DE L'OUEST: Ca Ira (MCZ 34489-99); 5 km S Dufort (MCZ 33281); Morne de
Cayette (MCZ 33293-94); Diquini (USNM 117187, USNM 140216-17); Port-au-
Prince (USNM 117141, USNM 118835, USNM 120968, AMNH 55136); vicinity of
Port-au-Prince (AMNH 44009-10); Petionville (ASFS V20238, ASFS V14965); 1 mi.
(1.6 km) NW Petionville (ASFS V24379); Morne Calvaire, 1 mi. (1.6 km) SW
P6tionville, 2300 feet (702 meters) (ASFS X1303); Fort Jacques, 5 km SE P6tion-
ville (airline), north versant, Morne l'H6pital (ASFS V8309); ca. 7 km (airline)
W PFtionville, ca. 2000 feet (610 meters) (ASFS V8480); Furcy (USNM 121018,
MCZ 3123, MCZ 31820).

Eleutherodactylus pictissimus eremus SCHWARTZ 1965:107
HOLOTYPE AND TYPE LOCALITY.-MCZ 43196; 9.7 mi (15.5 km) E Azua, Azua
Province, Repiblica Dominicana.
DIAGNOSIS.-A subspecies of E. pictissimus characterized by small size (snout-
vent length: males to 28 mm, females to 31 mm); dorsum very pale (yellowish tan
or pale tan); dorsal markings much reduced to faint brownish to grayish stippling
or filigreeing, but with a prominent dark brown scapular chevron that may be much
reduced or fragmented; a pair of faint dorsolateral stripes that are unicolor with the







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


dorsal color; sides grayish to pinkish, with dark pattern elements much reduced;
hind limbs marbled or stippled and rarely with any sort of banding indicated; con-
cealed surfaces pinkish to gray; venter pinkish to dark gray; throat white to grayish.
DISTRIBUTION.-The xeric Llanos de Azua in Azua and Peravia provinces, from
sea level to 700 ft (214 m) in the Sierra de Ocoa, Repuiblica Dominicana.
REMARKS.-At the time of its description E. p. eremus was known
from only two localities. The new material, from three additional sites,
agrees very well with the diagnostic characteristics listed in the original
description. The very pale dorsal color, the faint dorsal pattern, and
the very prominent dark scapular chevron, which may be faint or frag-
mented, serve to distinguish eremus from both of the western subspecies.
In addition, eremus is the smallest of the subspecies of E. pictissimus.
The largest female eremus have snout-vent lengths of only 31 mm, in
contrast to a maximum of 43 mm for pictissimus and 35 mm for apan-
theatus. Snout-vent lengths of males of the three subspecies are: eremus
28 mm, apantheatus 33 mm, and pictissimus 34 mm. The tibia/snout-
vent length (X 100) ratio is lower in eremus than in any other sub-
species (X=eremus males 47.0 mm, females 44.1 mm; apantheatus males
48.9 mm, females 45.4 mm; pictissimus males 48.9 mm, females 46.2 mm).
All mensural data are presented in Appendix 1.
Because of the arid region occupied by E. p. eremus, this subspecies
(like apantheatus) is found most abundantly in locally mesic situations
in the Llanos de Azua and the lower foothills of the adjacent Sierra de
Ocoa. At the type locality specimens were taken in huge piles of palm
trash within an artificial oasis composed of a coconut grove and large
shade trees. The locality north of Cruce de Ocoa is similar but less
mesic. The juvenile from south of Bani was taken from under very
dry palm fronds on mud at the edge of a mangrove swamp. Once
again, the tolerance of E. pictissimus to marginal haline situations, as
on Ile-a-Vache, is demonstrated.
No intergrades between eremus and apantheatus are known; the
closest known collection localities are 30 km apart (El Iguito; 9.7 mi. E
Azua). El Iguito lies on the eastern bank of the Rio Yaque del Sur in
a locally mesic area, whereas the area between El Iguito and the Azua
locality is the hot and extremely dry western portion of the Llanos de
Azua. The species probably occurs in the intervening area, but these
populations may well be limited to locally, and very restricted, mesic
situations within this intermediate region.

SPECIMENS EXAMINED.-REPOBLICA DOMINICANA. AZUA PROVINCE: 9.7
mi. (15.5 km) E Azua (MCZ 43196, AMNH 71982-86, ASFS X8058-61, ASFS
V19359-89); 6 km S Peralta (ASFS V21105). PERAVIA PROVINCE: 16.5 mi. (26.4
km) S San Jos6 de Ocoa, 500 feet (153 meters) (USNM 150931-33); 1.9 mi.
(3.0 km) N Cruce de Ocoa (ASFS V34095-100); 4.8 mi. (7.7 km) S Bani (ASFS
V29439).








SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


Eleutherodactylus weinlandi BARBOUR 1914:246
HOLOTYPE AND TYPE LOCALITY.-MCZ 2050; Puerto Plata, Puerto Plata Province,
Rep6blica Dominicana.
DIAGNOSIS.-A species of the Antillean ricordi group of Eleutherodactylus char-
acterized by moderate size (snout-vent length: males to 30 mm, females to 40 mm);
dorsum heavily to moderately overlaid with chocolate to black marbling or vermi-
culations on a yellowish, buffy, tan, or dark brown ground color, a pair of brightly
colored (orange, golden, rich buffy) dorsolateral stripes; a small, or restricted, to
large sacral blotch, contrasting (rust, orange, reddish brown, red, orange-brown,
chestnut) with the remainder of the dorsum; hind limbs (including femora and
crura) either heavily patterned with crossbars or immaculate, the ground color
similar to that of the sacral blotch area; inguinal and supra-axillary glandular areas
present, but usually very much obscured by the dark lateral coloration; expanded
digital discs very poorly developed to absent; tympanum small (X= males 2.1 to 2.4,
females 2.4 to 2.9, of three subspecies); tibia/snout-vent length (X 100) high
(males 41.9-55.7, females 45.1-56.8).
DIsTRIBUTION.-Central and eastern Hispaniola. In the Repiblica Dominicana
from the extreme western portion of the Cordillera Septentrional in Valverde
Province, southwestward along the northern and eastern slopes of the Cordillera
Central to the vicinity of Santo Domingo, but apparently absent on the eastern
tip of the island in much of La Altagracia Province. In Haiti known from the
northern coast (Anse a Margot), in the Chaine de Mathieux and the Montagnes de
Trou-d'Eau and their northern affiliates, and in the adjacent Dominican Sierra de
Neiba (Fig. 2). Altitudinal distribution from sea level at many localities to 2600
ft (793 m) west of Vallejuelo in the Sierra de Neiba and north of Puesto Grande
in the Cordillera Septentrional, and possibly occurring at higher elevations in the
latter range (records without precise elevations from Loma Quita Espuela and
Monte Isabel de Torres).

Eleutherodactylus weinlandi weinlandi BARBOUR 1914
DIAGNOSIS.-A subspecies of E. weinlandi characterized by small size (snout-
vent length: males to 27 mm, females to 36 mm); dorsum tan, creamy yellow,
yellow, or gray, heavily overlain with a very dark brown to black reticulum so that
the ground color often is apparent only as a fine vermiculation between the ex-
tensively dark dorsal pattern; a pair of brightly colored (buffy, golden, yellow,
orange, deep yellow, orange, or orange-red) dorsolateral stripes that contrast sharply
with the general dorsal pattern join a small and restricted brightly colored (rust,
orange-brown, rusty brown, pale orange-tan, reddish, bright orange, orange, or
orange-red) sacral area; hind limbs brightly colored, the hue similar to that of the
sacral area and heavily banded or stippled (especially on the crura) with dark
brown to black; concealed surfaces of femora whitish to flesh, heavily mottled or
irregularly banded with brown or black; venter opalescent whitish to gray; ratio of
tibia/snout-vent length (X 100) high (males 41.9-55.1, females 45.1-54.3).
DIsTrmrrTION.-Northern Haiti (Anse A Margot); the Chaine de Mathieux and
the Montagnes du Trou-d'Eau and their northern affiliates in southern Haiti, and the
adjacent Sierra de Neiba in the Repiblica Dominicana; central Rep6blica Domini-
cana from Valverde and Espaillat provinces in the north, south along the northern
and eastern slopes of the Cordillera Central in Santiago and La Vega provinces,
central San Crist6bal Province to within 17 km of Santo Domingo in the Distrito
Nacional. Altitudinal distribution extends from sea level along the northern Domini-
can and Haitian coasts to maximum elevations of 2000 ft (610 m) in the Cordillera
Central (west of Jarabacoa) and 2600 ft (793 m) in the Cordillera Septentrional
(north of Puesto Grande) and in the Sierra de Neiba (west of Vallejuelo).


























o18


I II


740 100 10 40 3D

FIGURE 2.-Map of Hispaniola, showing localities and known distributions of four species of Eleutherodactylus as follows: hexa-
gon, E. warren; triangles, E. alcoae; squares, E. probolaeus; circles, E. weinlandi. Subspecies of E. weinlandi indicated by shad-
ing: fine stippling, E. w. weinlandi; coarse stippling, E. w. chersonesodes; crosshatching, E. w. paralius; overlapping symbols
indicate area of intergradation between weinlandi and chersonesodes. z
P







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


REMARKS.-Throughout its broad geographic range, E. w. weinlandi
shows very little chromatic, pattern, or size variation. The pattern and
coloration described above apply to most specimens. There is some
variation in size of the sacral blotch, which is usually very small and
restricted but is somewhat larger in a few specimens. The regular
banding or stippling on the crura is a common feature of the series (see
Schwartz 1965: fig. 93).
E. w. weinlandi occupies mesic situations, from the lowlands to ele-
vations of 2600 ft (793 m) in the mountains. In the lowlands, the
species is most commonly encountered, like E. pictissimus, in Cocos
trash piles, under logs and rocks, and in piles of cacao husks. Because
much of the area occupied by E. w. weinlandi has dark soils and the
habitats are well shaded and dark, the coloration of the frogs is remark-
ably cryptic. At night the species is active on the ground, and we have
never taken specimens on shrubs or above the surface, except on small
rocks. Occasional individuals have been taken in small earthen cavities
in ravine banks, and one frog was taken from a large pile of old and
rotten logs in a purely edificarian situation at an abandoned sugar mill
(Hojas Anchas).
E. weinlandi can vocalize, but we heard it calling on only one occa-
sion, north of Puesto Grande; the call was a double insect-like chirp or
buzz that is typical of ricordi group members.
E. w. weinlandi intergrades with E. w. chersonesodes over a compact
area in northeastern Republica Dominicana (see details and specimens
examined beyond). What is puzzling about E. weinlandi is the extreme
disjunction of the western populations. The Anse a Margot locality
(from which a series was taken in moist piles of palm-tree shavings in
a Theobroma grove) is separated by 150 km from the nearest localities
to the east (Cordillera Septentrional north of Cruce de Guayacanes).
At least some of the intervening area (the xeric Valle de Cibao in Monte
Cristi Province) is totally unsuitable for such a mesophile as E. wein-
landi. However, within the Cibao, there are locally suitable areas
(for instance, near the Laguna de Salodillo on the Dominico-Haitian
border) that appear adequate for the species, but where it does ap-
parently not occur. Even more puzzling is the apparent disjunct dis-
tribution of the population in the Chaine des Mathieux-Montagne du
Trou-d'Eau-Sierra de Neiba, no records being available for a distance of
some 80 km (nearest approximations Vallejuelo and Jarabacoa). Much of
this intervening area is mesic or semi-mesic, although it includes the xeric
Llanos de Azua (which, it may be recalled, are occupied by E. pictissi-
mus). One logical area of continuity between the eastern and western
segments of E. w. weinlandi in this region is the southern mesic slopes







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


of the Cordillera Central (north of the Valle de San Juan), but we have
not yet found the species in that region. The distributional picture in
the area of the disjunctions is further complicated by the presence of
three frogs from localities circumscribed by the three regions occupied
by E. w. weinlandi. Details of this situation will be discussed beyond.

SPECIMENS EXAMINED. -HAITI. D.PT. DU NORD: ca. 2 km inland from Anse
a Margot (ASFS V10282-302). DEPT. DE L'OUEST: 9.1 mi. (14.6 km) SE Mire-
balais (ASFS V26541); 8.0 mi. (12.8 km) E Lascahobas (ASFS V26600-01);
12 mi. (19.2 km) N Port-au-Prince (MCZ 24289). REPCOBLICA DOMINICANA.
VALVERDE Pnov.: 8 km N Cruce de Guayacanes, 1400 ft (427 m) (ASFS V1237);
5.9 mi. (9.4 km) N Cruce de Guayacanes, 1400 ft (427 m) (ASFS V32272-74).
PUERTO PLATA PROV.: Puerto Plata (MCZ 2050, MCZ 23526); Monte Isabel de
Torres (MCZ 22466); 25 km S Puerto Plata (USNM 107596, MCZ 23548-50);
1 km N pass between Santiago and Puerto Plata, 2000 ft (610 m) (ASFS V1868-
69); 1 km N La Cumbre, 2000 ft (610 m) (ASFS V18100); Hojas Anchas, 9.0
mi. (14.4 km) NE Altamira, ca. 800 ft (244 m) (ASFS V32184); 8 km E Imbert,
1100 ft (336 m) (ASFS V1690); 11 km SE Sosia (ASFS V1718-19). ESPAILLAT
PROV.: 4 km SE Sabaneta de YAsica (ASFS V1699-701); 6 km SE Sabaneta de
Yasica (ASFS V1697); 9 km W Sabaneta de YAsica (ASFS V1713); 2 km SW
Jos6 Contreras, 2000 ft (610 m) (ASFS V1886); 2 km N Puesto Grande, 1400-
2200 ft (427-671 m) (ASFS V18055, ASFS V33572); 3.5 mi. (5.6 km) N Puesto
Grande, 2600 ft (793 m) (ASFS V33522); 10 km N Puesto Grande, 1900 ft (580
m) (ASFS V18300-02); 10 mi. (16.0 km) N San Victor, 1400 ft (427 m) (ASFS
V34029); 3.2 mi. (5.1 km) SE Gaspar HernAndez (ASFS V33732-33); 3.0 mi.
(4.8 km) NW Gaspar Hernmndez (ASFS V33768). SANTIAGO PROv.: La Cumbre,
2000 ft (610 m) (ASFS V18033); 1 km S La Cumbre (ASFS V18209); 4 km S La
Cumbre (ASFS V18074-83, ASFS V18143-47); 0.5 mi. (0.8 km) SE Pedro Garcia,
1500 ft (458 m) (ASFS V32211-17); 3.4 mi. (5.4 km) SE Los Montones, Rio Bao,
1600 ft (488 m) (ASFS V33811, ASFS V33944). LA VEGA PROv.: 3 km NW La
Vega (ASFS V1782); ca. 5 km N La Vega (ASFS V4174-76); 4 km N La Vega
(ASFS V4177); ca. 3 km N La Vega (ASFS V4263-64); 1.5 mi. (2.4 km) N La
Vega (ASFS V14087); Bayacanes (ASFS V16155-56); 13 km SW La Vega (ASFS
V1734); 17 km NE Jarabacoa (ASFS V1931-32); 8 km W Jarabacoa, 2000 ft (610
m) (ASFS V18388); 1.2 mi. (1.9 km) SE Bonao, 700 ft (214 m) (ASFS X8128);
75 km N Santo Domingo, road to Santiago (=near Piedra Blanca) (AMNH
44018, AMNH 44021). DUARTE PROV.: Loma Quita Espuela (MCZ 23525). SAN
CRIST6BAL PRov.: 2 km W Esperalvillo (ASFS V14394). DISTRITO NATIONAL: 17
km NW Santo Domingo (ASFS V3145-46). LA ESTRELLETA PROV.: 5 km S
Elias Pifia, 2200 ft (671 m) (ASFS V415); 6.7 mi. (10.7 km) E Hondo Valle,
2500 ft (763 m) (ASFS V31409, ASFS V31425-26). SAN JUAN PROv.: 7 km W
Vallejuelo, 2600 ft (793 m) (ASFS V392).

Eleutherodactylus weinlandi chersonesodes SCHWARTZ 1965:115

HOLOTYPE AND TYPE LOCALITY.-MCZ 43203; 8 km W SamanA, Samana Province,
Republica Dominicana.
DIAGNOSIS.-A subspecies of E. weinlandi characterized by small size (snout-
vent length: males to 28 mm, females to 35 mm); dorsum almost completely over-
lain with a dark brown reticulum with faint indications of a pale tan ground color;
a pair of contrasting, but not particularly brightly colored (yellow-buff, yellow,
dull yellow, buffy) dorsolateral stripes that join a large, richly colored (rich orange-
brown, rich reddish brown, reddish brown, or dark orange-brown) sacral area; hind







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


limbs as richly colored as sacral area, without markings on the crura but with a
dark brown reticulum on a pinkish-white background on the concealed surfaces of
the femora, and with a band of the sacral color extending along the anterior face
of the femora; venter and throat gray to dark pearl gray; ratio of tibia/snout-vent
length (X 100) high (males 47.4-55.7, females 45.4-56.8).
DisTRIsuTIoN.-Eastern Hispaniola, including the Peninsula de Samana and
the eastern Repiblica Dominicana from northeastern San Crist6bal Province south-
eastward to central La Altagracia Province. Intergradation with E. w. weinlandi
in Duarte, Maria Trinidad SAnchez, SAnchez Ramirez, and La Vega provinces.
REMARKS.-The large number of specimens presently available of
E. w. chersonesodes confirm the diagnostic characters of the subspecies.
Juvenile individuals may have the crura with some vague stippling or
marbling, which is regularly absent in the long series of adults from both
the Peninsula de Samana and eastern Repuiblica Dominicana. This pat-
tern easily distinguishes chersonesodes from weinlandi. In addition, the
dorsum of chersonesodes is more solidly dark brown (see Schwartz
1965:figs. 93 and 94), with only faint remnants of a paler ground color;
the sacral blotch area is much more extensive in chersonesodes than in
weinlandi; and the basic colors involved in the sacral blotch area in
chersonesodes are darker and richer than those in weinlandi. The two
subspecies are approximately of the same size (and there are now long
series of chersonesodes available), and the tibia/snout-vent length ratios
are quite comparable. The darker gray ventral color of chersonesodes
is distinctive when compared with the pale ventral color of weinlandi.
E. w. chersonesodes is extremely abundant on the Peninsula de
Samand, where it occurs both in the lowlands near the coast (where one
specimen was secured from a small and decayed pile of Cocos trash
within 10 ft [3.1 m] of the mangrove border) and in the Sierra de
Samana, where natives found many more specimens than we preserved.
In the eastern non-SamanA, areas of the Rep6blica Dominicana, cherso-
nesodes occupies mesic regions and occurs in coastal Cocos groves (as
at Playa El Coco and Playa de Guaco) and in cacaotales and cafetales
associated with the lowlands of the Cordillera Oriental. The altitudinal
distribution of chersonesodes is from sea level to 1000 ft (305 m) be-
tween Sanchez and Las Terrenas in the Sierra de Samani and near Gon-
zalo in the haitises region. Although there are presently no records from
the Cordillera Oriental (which reaches a maximum elevation of 2296 ft
[700 m] in its eastern section), I have no doubt that the species occurs
there, and also that the hiatus shown on the map (Fig. 2) is artificial.
Intergradation between weinlandi and chersonesodes occurs in a com-
pact region embracing four provinces. In this lowland and very mesic
region, specimens show precisely those characteristics which are inter-
mediate between the two subspecies-vague banding or marbling on the
hind limbs, medium size of the sacral blotch, and intermediate extent


1976







BULLETIN FLORIDA STATE MUSEUM


of the dark dorsal pigmentation, as well as less rich and more bright
coloration of the sacral area.

SPECIMENS EXAMINED.-REPOiBLICA DOMINICANA. SAMANA PROVINCE:
3.3 mi. (5.3 km) NE SAnchez, 1000 ft (305 m) (ASFS V34126); 7.6 mi. (12.2
km) NE SAnchez, 1000 ft (305 m) (ASFS V34288-96, ASFS V34352-83, ASFS
V34839-81); 14 km E SAnchez (ASFS V2010); 22 km E SAnchez (ASFS V14120);
5 km E Las Terrenas (ASFS V21848-51); Samana (USNM 74983-92, AMNH
34201-02); west of SamanA (AMNH 34519); SamanA and Laguna (USNM 65106-
08); 6 km W Samana (ASFS V1868-71); 8 km W SamanA (MCZ 43203, ASFS
V1989-93); Laguna (AMNH 34495-96); Rojo Cabo (USNM 34261); Las Flechas
(not mapped) (AMNH 34177-80); Rio San Juan (USNM 74625, USNM 74629-40,
UMMZ 92212-13); Peninsula de SamanA (USNM 66980). EL SEIBO PRov.: Cafio
Hondo (AMNH 3297-301); Cueva de Cafio Hondo (ASFS X9278-83); Boca del
Infierno (USNM 74960-61); Sabana de la Mar (AMNH 34199, AMNH 44133);
3.5 mi. (5.6 km) S Sabana de la Mar (ASFS X7971-74); Las Cafiitas (USNM
65709, ASFS V35321); 10.5 km N Hato Mayor (ASFS V35328, ASFS V35301-02,
ASFS V35306-07); 1.4 mi. (2.2 km) E Miches (ASFS X9341); 1.1 mi. (1.8 km)
W Miches (ASFS V28786-92); 2.6 mi. (4.2 km) NE La Vacama (ASFS V28986-
91); 20.2 mi. (32.3 km) NW, 3.4 mi. (5.4 km) N La Vacama, Playa de Guaco
(ASFS V29327-76, ASFS V29415-16); 7 km W El Cuey (ASFS V17584-85).
LA ALTAGRACIA PROV.: 24.8 mi. (39.7 km) ESE Miches (ASFS X7893); Playa El
Coco, 46 km N Higiiey (ASFS V17499-505); 3.2 mi. (5.1 km) W Higiiey (ASFS
X751-57); 4.7 km NW La Enea (ASFS V948). SAN CRIST6BAL PROv.: 5.0 mi.
(8.0 km) NE Gonzalo, 1000 feet (305 meters) (ASFS V29539).
E. w. weinlandi X E. w. chersonesodes. REPtJBLICA DOMINICANA. DUARTE
PRov.: 6 km NE San Francisco de Macoris (ASFS V2947); ca. 4 km NE Pont6n,
Rio Cuaba (ASFS V3001); 6.4 mi. (10.2 km) SE Tenares, 700 ft (214 m) (ASFS
V33555); 7.5 mi. (12.0 km) NW Cruce de Pimentel (ASFS V33481-84); 9 km
NW Pimentel (ASFS V1820-22); 3 km NE Villa Riva (ASFS V1827-32). MARiA
TRINIDAD SANCHEZ PROv.: 3.3 km S Cabrera (ASFS V4253); 2.1 mi. (3.4 km)
NE Rio San Juan (ASFS V33698-702); 2 km S El Factor (ASFS V1853-59); 4 km
N Azucey (ASFS V16081-83); 1.0 mi. (1.6 km) S Cafio Hondo (ASFS V34135-39).
SANCHEZ RAMIREZ PROv.: 4.4 km E Cotui (ASFS V627-29); 1 km SE La Mata
(ASFS V18439, ASFS V18574-78, ASFS V33671). LA VEGA PROV.: 8 km S Moca
(ASFS V4332).

Eleutherodactylus weinlandi paralius new subspecies
FIGURE 3

HOLOTYPE.-USNM 194004, an adult female, from 14 km SE, 1 km N Boca
Chica, San Pedro de Macoris Province, Republica Dominicana, one of a series
taken by James R. Dennis, James A. Rodgers, Jr., and Albert Schwartz on 12
August 1969. Original number ASFS V19390.
PARATYPES.-AMNH 87216-49, CM 54132-35, MCZ 84637-40, USNM 194005-08,
same data as holotype; ASFS X9321, 12 km NE La Romana, La Romana Province,
Repuiblica Dominicana, R. Thomas, 19 July 1963; ASFS V35081, 12 km NE La
Romana, La Romana Province, Republica Dominicana, D.C. Fowler, 22 November
1971; ASFS V28596-97, 7 km S Aeropuerto Internacional de las AmBricas, Punta
Caucedo, Distrito Nacional, Repiblica Dominicana, D. C. Fowler, 1 July 1971;
MCZ 84641, 11 km S Aeropuerto Internacional de las Am6ricas, Punta Caucedo,
Distrito Nacional, Rep6blica Dominicana, D. C. Fowler, 7 July 1971; ASFS V29511-
14, 11 km E Boca Chica, Distrito Nacional, Rep6blica Dominicana, D. C. Fowler,
A. Schwartz, 6 August 1971; ASFS V22445, Santo Domingo, La Feria, Distrito
Nacional, Repdblica Dominicana, A. Schwartz, 19 December 1970; ASFS V2465,


Vol. 21, No. 1








SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


FIGURE 3.-Dorsal views of (left) Eleutherodactylus w. paralius holotypee, USNM
194004) and (right) E. warren holotypee, CM 54138).

ASFS V2467, 5.1 mi. (8.2 km) E Santo Domingo (from Rio Ozama), Distrito
Nacional, Repiblica Dominicana, D. C. Leber, 19 June 1964; CM 54136-37, USNM
194009-11, 1.8 mi. (2.9 km) S Boca del Soco, San Pedro de Macoris Province,
Repiblica Dominicana, D. C. Fowler, native collectors, 16 July 1971; AMNH
87250-51, MCZ 84642-44, 15.5 mi. (24.8 km) E San Pedro de Macoris, Rio Cu-
mayasa, La Romana Province, Repiblica Dominicana, D. C. Fowler, A. Schwartz,
12 July 1971.
DISTRIBUTION.-The southern Dominican coast, from Santo Domingo in the
west, east to La Romana Province. Intergradation with E. w. weinlandi and E. w.
chersonesodes unknown, but see discussion beyond.
DIAGNOSIS.-A subspecies of E. weinlandi characterized by large size (snout-
vent length: males to 30 mm, females to 41 mm); dorsum yellow, pale yellow-tan,
or pale buffy; dorsolateral stripes unicolor with dorsal ground color (occasionally
slightly more yellow to golden); dorsal dark pattern brown, usually very restricted
and diffuse, with only a dark scapular chevron clearly defined in many specimens;
sacral area pale rusty, brick red, rich orange, rust, or bright orange, restricted by
paler ground color anteriorly; hind limbs pale rusty, pale orange, dull reddish brown,
rust, bright orange, or tan; crura unmarked, with darker or with some vague darker
gray marbling or suffusions, occasionally with clear dark transverse bars; concealed
surfaces whitish, tan, or gray, with dark brown marbling; venter white to pale
gray; ratio of tibia/snout-vent length (X 100) low (males 44.1-50.5, females 44.8-
50.6).
DESCRIPTION OF HOLOTYPE.-An adult female with the following measurements
(in mm): snout-vent length 40.4, head length 14.8, head width 14.5, longitudinal
diameter of tympanum 3.2, longitudinal diameter of eye 5.0, distance from naris
to anterior corner of eye 4.5, femur 17.5, tibia 18.3, fourth toe 16.0, tibia/snout-
vent length (X 100) 45.3. Head slightly longer than distance from snout to poster-
ior border of tympanum; snout truncate, with nares prominent at anterior ends
of canthus rostralis; diameter of eye longer than distance from naris to anterior
corner of eye; interorbital space 5.0, equal to diameter of eye; diameter of tympanum
much less than diameter of eye; distance from tympanum to eye equal to one-third
diameter of tympanum. Digital discs present, small, largest on digits 3 and 4, that
of digit 3 the largest and equal to about one-quarter size of tympanum. Fingers
long, unwebbed, 3-4-2-1 in order of decreasing length; subarticular tubercles well
developed, prominent, pale gray. Toes relatively long, all with slight basal webbing,
4-3-5-2-1 in order of decreasing length; subarticular tubercles large, prominent,
gray. Heels do not touch when femora are held at right angles to body axis. Dor-
sum smooth, with two non-glandular warts between the angle of the jaw, tympanum,
and forelimb insertion; throat and venter smooth, belly disc feebly developed.


1976







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


Dorsal surfaces of all limbs smooth; posterior faces of thighs with low, pavement-
like granules. Inguinal and supra-axillary glands absent. Tongue large, very
slightly nicked, free behind, its greatest width equal to about one-half that of floor
of mouth. Prevomerine teeth in two long arched series, extending from outside
the choanae and adpressed against them, the two series separated from each other
medially by a distance equal to about one-quarter the diameter of a choana.
COLORATION OF HOLOTYPE.-Dorsal ground color yellow, with a dark brown
dorsal pattern composed of an irregular dark interocular bar and a moderately
prominent scapular chevron, the area between these two figures marbled irregularly
with dark brown; a pair of dorsolateral stripes, which are concolor with the dorsal
ground color; sides and mid-back irregularly marbled with dark brown, with a
restricted pale rusty sacral blotch area; upper surfaces of hind limbs pale rusty, with
the concealed surfaces marbled heavily with dark brown, the crura and pedes
virtually immaculate; forelimbs yellow with some vague grayish markings; ventral
ground color white with fine brownish stippling on throat.
VARIATION.-Eleven males have the following measurements (in mm) (extremes
and means): snout-vent length 22.0-29.5 (25.6); head length 8.7-11.7 (9.9); head
width 8.4-11.1 (9.7); tympanum 2.2-2.7 (2.4); eye 3.0-4.0 (3.4); naris to eye 2.4-
3.6 (2.9); femur 9.3-13.3 (11.1); tibia 10.6-14.6 (12.0); fourth toe 8.9-12.7 (9.6);
tibia/snout-vent length (X 100) 44.1-50.0 (46.7). Thirteen gravid and adult
females measure: snout-vent length 29.3-40.4 (35.4); head length 10.9-14.8 (12.2);
head width 10.9-14.9 (13.2); tympanum 2.4-3.3 (2.9); eye 3.9-5.0 (4.5); naris to
eye 3.1-4.7 (4.1); femur 12.2-18.0 (15.7); tibia 13.2-19.3 (16.5); fourth toe 11.8-
16.9 (14.5); tibia/snout-vent length (X 100) 44.8-50.6 (46.7).
E. w. paralius is somewhat variable in dorsal pattern. Some specimens (AMNH
87248-49) are almost as heavily patterned dorsally as E. w. weinlandi. The heavy
dorsal markings of ASFS X9321 from 12 km NE La Romana have been pre-
viously noted (Schwartz 1965:119). Taken as a group, however, both adults and
juveniles agree quite well with the pattern description of the holotype. There is no
evidence that specimens from the two range extremes (Santo Domingo; 12 km
NE La Romana) are any closer to the respective subspecies, which they approxi-
mate geographically (weinlandi in the west, chersonesodes in the east), than are
specimens from the more central portion of the range of paralius. Consequently,
there are no known intergrades between paralius and either weinlandi or chersone-
sodes.
The dorsal pattern typically consists of a much reduced dark brown reticulum
(that is usually apparent only on the dorsum) bounded by a fairly well-defined
interocular dark bar, and a dark scapular chevron, and another at about the middle
of the back. Pale dorsolateral stripes are present but usually are similar to the pale
(yellow, pale yellow-tan, pale buffy) dorsal ground color. The sacral area is faintly
colored with pale rusty, brick red, rust, rich or bright orange; the same colors occur
on the hind limbs, the crura of which are immaculate or with some vague grayish or
brownish marbling or stippling. The concealed surfaces of the femora are whitish,
tan, or gray, and extensive dark brown markings are present. The venter is white to
pale gray, as is the throat, and the venter has, in addition, some fine dark brown
stippling in most adults.
The supra-axillary and inguinal glandular areas' distinctness is variable in oc-
currence, but both areas are readily discernible in less heavily pigmented indi-
viduals.
COMPARISONS.-E. w. paralius differs from both E. w. weinlandi and E. w.
chersonesodes in having a greatly reduced dorsal pattern and larger size (see
Appendix 1). The hues involved in the sacral area and hind limbs of paralius are
less bright than in either of the other two subspecies, and the tibia/snout-vent ratio
is much less in both sexes of paralius than in both sexes of either weinlandi or
chersonesodes. Detailed comparisons are unnecessary; the three subspecies are ex-







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


ceptionally distinct in numerous details of size, pattern, and coloration. The differ-
ences are so striking that when I collected the first series from the type locality, I
was uncertain that these frogs were correctly assigned to E. weinlandi.

REMARKS.-E. w. paralius, as its name implies, occurs along the south-
ern coast of the Republica Dominicana, a region that is moderately mesic,
with residual stands of semi-mesic woods often situated on limestone
platforms or substrate. The type locality is such a place, having a rich
but open lowland forest on a diente de perro base. The woods south
of the Aeropuerto Internacional on Cabo Caucedo are somewhat more
luxuriant and lie on a raised limestone platform surrounded by sand
and rock beach. The specimens from La Romana were found in a
deep, rocky, well-forested, mesic ravine that cuts through otherwise
cultivated sugar cane fields-an altogether specialized situation. One
frog was taken from under an old, deep trash pile behind the exposed
beach on Cabo Caucedo, and the specimens from the Rio Cumayasa
were taken at night from paths in a grassy riverside pasture and on rocks
adjacent to a gravel road. The Boca del Soco frogs were secured in
Cocos trash piles at the edge of a coastal palm grove, and the single
specimen from Santo Domingo was taken from beneath a large rock in
a moist, dead grass pile near the ocean. Mertens (1939:30) reported
a specimen from Cueva de Santa Anna, which lies within the city of
Santo Domingo; he described the sacral area and hind limbs as "lebhaft
ziegelrot," in agreement with the rust color typical of paralius. The
specimens from east of Santo Domingo were secured at night in a mesic
limestone-cliff forest.
The voice of E. w. paralius has been described as "a series of irregular
bubbly 'bleeps' or 'ticks'" (type locality), and a "low intensity 'peep'"
(east of Santo Domingo). The males call erratically and are easily
disturbed, even under very favorable moisture conditions. The calling
sites include a cave (east of Santo Domingo), large rocks, and herbs
and shrubs (up to 3 ft [0.9 m] above the ground) at the type locality.
The favored shrub for calling here is Zamia. At the Rio Cumayasa a
male was secured as it called from exposed rocks adjacent to a gravel
road through semi-mesic Acacia woods. Another male was found calling
on a low shrub, 6 in (15 cm) above the ground in limestone woods 11
km E Boca Chica.
The association of E. w. paralius with limestone areas on this south-
ern coast is probably not fortuitous. Such porous limestone outcrop-
pings and platforms offer advantageous diurnal retreats for the frogs and
also support (even today) relatively luxuriant wooded growth for noc-
turnal protection and activities. However, it is also obvious that E. w.
paralius can persist in situations close to and associated with human


1976







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


activities, if there are trash and vegetational heaps that offer moist
diurnal havens.
The absence of acknowledged intergrades between paralius and either
weinlandi or chersonesodes is puzzling, because paralius presumably
comes in contact with these subspecies at the western and eastern
extremes of its range, respectively. Although only a short distance
separates the closest paralius and weinlandi localities (Santo Domingo
and 17 km NW Santo Domingo), the two taxa remain quite distinct in
these areas. The same holds true at the eastern extreme of the range,
where the localities for paralius and chersonesodes (12 km NE La
Romana and 4.7 km NW La Enea, respectively) are separated by about
25 airline km. In neither case is there an obvious geographic barrier
that might explain the separation of, and lack of intergradation between,
the subspecies.
The map (Fig. 2) shows a large hiatus in the range of E. weinlandi.
The subspecies paralius occurs in the south, weinlandi in the west, and
chersonesodes in the north and east. I do not think this hiatus actually
exists. The area is quite mesic, and there are extensive riverine gallery
forests that must support E. weinlandi. The relationships of the three
subspecies within this region will be most interesting to ascertain.

Eleutherodactylus alcoae SCHWARTZ 1971:26
HOLOTYPE AND TYPE LOCALITY.-CM 45889; 22 km NE Cabo Rojo, 1500 ft (458 m),
Pedernales Prov., Republica Dominicana.
DIAGNOSIs.-A species of the Antillean ricordi group of Eleutherodactylus char-
acterized by the combination of moderate size (snout-vent length: males to 38 mm,
females to 45 mm); dorsum yellow or flesh-colored to tan or pale tan to yellow-
green with various dark dorsal markings, the most stable of which are a dark (gray
to almost black) interocular bar and a dark scapular chevron; remainder of dorsum
mottled, marbled, or irregularly marked with dark brown pigment; dorsolateral stripes
absent; hind limbs stippled or marbled with dark (never crossbanded), concealed
surfaces brown to purplish, occasionally suffused with orange, and without a dark
reticulum or vermiculations; venter white; throat heavily to moderately stippled with
brown; throat ground color, underside of all limbs and brachia bright yellow in some
specimens; inguinal and supra-axillary glandular areas present; digital discs ex-
panded and well developed, that of digit 3 equal to about one-third to one-half
size of tympanum; tympanum large (X=males 3.1, females 3.4), and tibia/snout-
vent length (X 100) low (males 41.8-51.6, females 43.6-47.5).
DITRImurriON.-The Rep6blica Dominicana, south of the Massif de la Selle-
Sierra de Baoruco, from the Rio Pedernales east to 32 km SE Pedernales, on the
Peninsula de Barahona; may also be expected in extreme southeastern Haiti (Fig. 2).x
Altitudinal distribution from sea level to 2000 ft (610 m) in the Sierra de Baoruco.

' Since this manuscript was completed, specimens of E. alcoae (ASFS V39545, ASFS V39976-84)
were collected at the village of Los Patos, Barahona Province, Repfblica Dominicana, immedi-
ately adjacent to the ocean on the east coast of the Peninsula de Barahona. The habitat is a
steep and cave-riddled hillside with dense hardwood forest cover, a much more mesic situation
than any other where E. alcoae has been collected. One frog was taken during the day on the







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


REMARKS.-E. alcoae was described on the basis of five frogs from
two general regions: a cave in the very arid lowlands of the Peninsula
de Barahona and rather mesic deciduous-Acacia forests at elevations be-
tween 1500 and 2000 ft (458 and 610 m) on the Sierra de Baoruco
above Cabo Rojo. We made a special search for additional E. alcoae
in 1971, and there are now 52 specimens of the species from eight lo-
calities in this region. The westernmost specimens are from 6.0 mi
(9.6 km) N Pedernales, along the Dominico-Haitian border, and the
easternmost are from 7 km N, 20.0 km SE Cabo Rojo (= 32 km SE
Pedernales). The species extends into the Sierra de Baoruco along the
road from Cabo Rojo to Aceitillar to an elevation of 2000 ft (610 m)
but appears to be rather uncommon at such a high elevation. In some
areas E. alcoae is precisely syntopic with E. p. pictissimus; and although
the two species are not easily recognized at night in the field by flash-
light, they are nevertheless quite distinct in many ways. Both reach
about the same size (it may be recalled that it is the Sierra de Baoruco
E. p. pictissimus that are the largest specimens of that subspecies), but
the tympanum of alcoae is larger in both sexes (alcoae: males 2.6-3.7,
females 4.2-5.6; p. pictissimus: males 1.8-2.8, females 2.2-3.9). Other
mean differences are also apparent for most measurements (see Appendix
1), but the tibia length means for females of both species are very com-
parable. E. alcoae may also be readily differentiated from E. pictissimus
by its well-developed digital discs.
I have noted previously that E. pictissimus is apparently mute. E.
alcoae, on the other hand, is vocal, and loud choruses of the species
were encountered on several occasions in one of the most rigorous situa-
tions for an anuran in the Antilles. On 14 August 1971 we visited an
arid limestone area with a flora of cacti and Acacia that lies 32 km SE
Pedernales. Many males were heard vocalizing (the call a single
"cheep"), despite no previous rain in this desert area. There was a brief
10 minute rain that greatly increased the number of calls, but we were
unable to secure or even locate any males actively calling. We revisited
this same locality on 22 August 1971, and once again, despite no rains
on that day (but heavy rains occurred during the late evening of 21
August), we heard many calling male alcoae. Their vocalizations were
a single, or occasionally double, "cheep," and we were successful in lo-
cating and collecting calling males from shrubs, bushes, and a tree crotch

floor of a cave, and the remainder were secured at night while the males were calling a single
(occasionally two-note) chirp from vines and shrubs up to 6 ft (1.8 m) above the ground and
from rock crevices near the cave entrance. Many more were heard than were collected. Thirty-
two eggs were deposited over-night in the plastic bag in which both males and females were
housed; whether this represents the clutch of more than one female is unknown. In color and
pattern, the Los Patos series does not differ strikingly from specimens from low elevations on
the Peninsula de Barahona; the venters are white and the throats are finely stippled with gray.
The Los Patos locality is not shown on the distribution map in the present paper.


1976







BULLETIN FLORIDA STATE MUSEUM


up to 4 ft (1.2 m) above the ground. None was found calling from the
ground, rocks, or the pocketed limestone substrate. Even more remark-
ably, two frogs were secured in amplexus, completely exposed on bare
limestone. The upper frog was grasping the lower frog anterior to the
forelimbs, and both were walking slowly with the hind limbs straight
and 'in almost tetanic contraction. Amplexing pairs of any Antillean
Eleutherodactylus are extremely difficult to locate, and rarely has am-
plexus been seen in any Antillean species (only E. cundalli and E.
nubicola in Jamaica and E. patriciae in Hispaniola as far as I am aware).
However, the peculiar stance and gait of this pair suggested that open
and exposed areas may not be the customary site of amplexus; when
the two individuals were dissected in the laboratory, it was found that
they were both males. This is the first instance of which I am aware
of homosexual amplexus being observed in Eleutherodactylus, although
it is a common occurrence in many (primarily water-breeding) anurans.
A second instance of male vocalization occurred on 21 August 1971
in the vicinity of L'Eglise at an elevation of 1500 ft (458 m) in the
Sierra de Baoruco. Here males were calling from exposed rocks, cavities
in rocks, an exposed root 3 in (8 cm) above the ground, and the stems
and leaves of shrubs. Finally, males were heard calling and collected
6.0 mi (9.6 km) N Pedernales at an elevation of 800 ft (244 m). The
frogs were calling from saplings (3 ft [0.9 m] above the ground), vine
tangles (3 ft), and on diente de perro limestone outcroppings on the
ground in a more or less shaded and semi-mesic cut-over Acacia hillside
adjacent to an open field. Many other males were heard calling on the
east side of the road at this locality, in Acacia weeds with a Bryophyllum
understory.
The locality records suggest that E. alcoae's distribution is primarily
lowland, and it is associated in these lowland regions with eroded lime-
stone. In every instance, our specimens were taken on or near pocketed
and eroded limestone, regardless of the surrounding vegetational asso-
ciations (varying from harsh open desert to semi-mesic Acacia-deciduous
woods). The species may be extremely abundant and easily secured,
once one knows the calling sites and habitat requirements. The Penin-
sula de Barahona is sandy along its eastern and western shores, but a
rugged and raised limestone ridge courses down the center of the penin-
sula. This ridge supports an extensive forest, and our collecting localities
southeast of Pedernales lie on or immediately adjacent to this ridge,
which reaches a maximum elevation of 322 m in Cerro Caballo south
of the Pederales-Oviedo road. We never collected alcoae at Oviedo,
in the now cultivated savannas in the east, nor at Pedernales in the
very arid and sandy regions in the west. Once one begins to ascend


Vol. 21, No. 1







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


the Massif de la Selle above Pedernales, however, the limestone out-
croppings begin and E. alcoae occurs.
E. alcoae is exceptionally variable in dorsal pattern and to some
extent in coloration. The dorsal ground color is pale (yellow, flesh, pale
tan, tan, or yellow-green), with dark gray to brown markings. These
markings may be more or less uniform in distribution (ASFS V30093),
or they may be more coarse and show a prominent dark interocular bar
and scapular chevron (ASFS V30044). All variations between these two
extremes occur; in one extreme instance the dorsum appeared uniformly
and faintly stippled, without an interocular bar or chevron (ASFS
V30065). The concealed surfaces are brown, but occasional specimens
have an orange wash or suffusion on these surfaces. The most striking
differences in color in E. alcoae are those associated with elevation.
Lowland (up to 600 ft [183 m] elevation) frogs are pale and much less
brightly colored than those from upland areas (1500 ft [458 m] and
above). In the latter specimens, the throat, brachia, and undersides
of the hind limbs are bright yellow, whereas in lowland specimens the
throat is white, stippled or blotched with brown, and the brachia and
limbs lack any bright pigment. It has not yet been determined whether
these differences result from deme formation in a frog with specialized
habitat requirements or from differentiation at the subspecific level in E.
alcoae populations. The fact that the known range of E. alcoae is at
present so circumscribed argues against the latter interpretation. Once
specimens are available from southeastern Haiti (where E. alcoae un-
doubtedly occurs, since we have specimens from very near the Rio
Pedernales, which here forms the international boundary), a more definite
interpretation can be made of these pigmental differences between low-
land and highland specimens.

SPECIMENS EXAMINED.-REPtBLICA DOMINICANA. PEDERNALES PROVINCE:
6.0 mi (9.6 km) N Pedemales, 800 ft (244 m) (ASFS V30091-105); 24 km NE
Cabo Rojo, 2000 ft (610 m) (ASFS V16678); 22 km NE Cabo Rojo, 1500 ft (458
m) (CM 45889); 21 km NE Cabo Rojo, 1500 ft (458 m) (USNM 166959); 21 km
N Cabo Rojo, L'Eglise, 1500 ft (458 m) (ASFS V30038-46); 8 km N, 2 km E
Cabo Rojo (ASFS V21544, MCZ 43255); 7 km N, 2.4 km SE Cabo Rojo, 200 ft
(61 m) (ASFS V29708-09); 7 km N, 17.6 km SE Cabo Rojo (ASFS V30075-78);
7 km N, 20.0 km SE Cabo Rojo, 600 ft (183 m) (ASFS V29763-65, ASFS V30059-
72).

Eleutherodactylus probolaeus SCHWARTZ 1965:110
HOLOTYPE AND TYPE LOCALITY.-MCZ 43197; 0.5 mi (0.8 km) NW Boca de Yuma,
La Altagracia Province, Repiblica Dominicana.
DIAGNOSIS.-A species of the Antillean ricordi group of Eleutherodactylus char-
acterized by the combination of moderate size (snout-vent length: males to 30 mm,
females unknown); dorsum tan, pale tan, or yellow-tan, having mottled dark brown


1976







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


to black pattern with three prominent dark areas (an eroded or truncate inter-
orbital triangle with its apex pointed posteriorly, a broad scapular chevron,
and an irregular rectangular or subquadrate mid-back blotch); dorsolateral stripes
pale, same color as dorsal ground color; hind limbs marbled dark brown, the pat-
tern of which may consist of two or three vague crossbars on the crura; concealed
surfaces of femora black or dark brown, paler (gray) medially and showing some
remnants of the pale dorsal ground color; venter opalescent; throat and underside of
limbs gray; inguinal and supra-axillary glands present; expanded digital discs well
developed, that of digit 3 equal to about one-quarter size of tympanum; tympanum
small (2.3-3.0 in males); and tibia/snout-vent length (X 100) low (47.0-51.7 in
males).
DIsTRIBUTION.-Known only from the vicinity of the type locality, La Altagracia
Province, Rep6blica Dominicana (Fig. 2).

REMARKS.-E. probolaeus was described (as a subspecies of E. pic-
tissimus) from three male specimens collected in 1963. Despite re-
peated visits to the type locality and the fact that the frog appears to
be common there, only six additional specimens (all males) have been
found. Many males have been heard calling (a trilling "brrrt" call,
followed at times by a series of "bzeut's"), but the dense woods and the
fact that the frogs call primarily from areas of dense understory makes
it extremely difficult to collect specimens. The two new localities are
both almost adjacent to the type locality.
The area where E. probolaeus occurs is remarkably specialized. All
specimens have been taken from the flat top of a limestone ridge that
parallels the coast behind the village of Boca de Yuma. The forest is
semi-mesic, with many high trees, lianas, epiphytes, often a dense under-
story consisting of Bryophyllum and Zamia, as well as other small herbs
and shrubs. The old road between Higiiey and Boca de Yuma ascended
this ridge and then descended on the southern face, so that the crossing
was accomplished in 2.5 km, much of which was steep. A new road has
been completed between these two settlements. It reaches Boca de
Yuma from the west, ascends, and then passes along the ridge top for
several kilometers. In 1971 we did not find E. probolaeus along this
new stretch of road, which passes through virtually undisturbed forest
similar to that at and near the type locality, just a few kilometers to
the east. The frogs were not heard calling in this region, although on
the same night many were heard calling at the type locality. Likewise,
in the slightly more xeric, but far from arid, deciduous woods near the
coast between Punta Cana and Juanillo, we did not hear or encounter
E. probolaeus, although this area also appears at least grossly suitable.
I can offer no explanation for the apparently very circumscribed distri-
bution of E. probolaeus. It appears to be a recent arrival from else-
where, but the species seems not to be closely allied to E. pictissimus
(see below), and certainly not to E. weinlandi. E. probolaeus may







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


demonstrate a relict distribution in this southeastern area, but even this
suggestion suffers from the very small area it is known to occupy.
I described E. probolaeus as a subspecies of E. pictissimus, even
though there is a hiatus of some 185 km to the east of the closest
known E. pictissimus station (south of Bani). We now know that the
intervening area is in large part occupied by E. weinlandi paralius, a
species with which E. probolaeus cannot logically be associated. Al-
though the patterns of pictissimus and probolaeus are somewhat similar
(dark dorsal markings on pale dorsal ground and dorsolateral stripes
that are colored similarly to the back), the pattern details are quite
different (Schwartz 1965:fig. 92). The digital discs in probolaeus are
larger than those of pictissimus, and probolaeus is vocal whereas pictis-
simus appears to be mute. Although probolaeus is a stockier frog, this
difference is not apparent when comparing the tibia/snout-vent length
ratios in males of the two species (Appendix 1). Everything considered,
it seems likely that probolaeus is an eastern Hispaniolan endemic whose
history has not been closely allied to that of pictissimus (see discussion
below).
Calling male E. probolaeus have been taken from a shrub limb and
a Bryophyllum leaf, both 2 ft (0.6 m) above the ground. Bryophyllum
and Zamia both are common calling sites for males, and the frogs have
been taken between 1 and 6 ft (0.3 and 1.8 m) on these plants. A single
male was collected on a pile of dead and leafless slash adjacent to a
path through the forest at a height of 4 ft (1.2 m) above the ground.
Interestingly, many males were heard calling on nights following hot
and dry days, whereas on nights after rainy and overcast days the
number of calling males was decreased.
In an effort to locate a related (to E. probolaeus), non-Hispaniolan
species of the Eleutherodactylus ricordi group, I have examined a series
of 26 E. monensis Meerwarth, taken by Richard Thomas in 1965 at
Anclaje Sardinero, on Isla Mona. This inlet lies some 70 km to the east
of the extreme southeastern tip of Hispaniola, and thus it would seem
reasonable from a geographic standpoint that E. probolaeus might be
related to E. monensis. The two species are similar, but no more so
than either is to other species within the complex. Examination of eight
male E. monensis shows that the Mona species appears not to reach
so large a size as probolaeus (26.4 versus 29.8 mm) and has lower means
in all head and limb segments measurements (Appendix 1). The tibia/
snout-vent length ratio also is slightly lower in monensis than in probo-
laeus. Thomas's field notes on E. monensis indicate that the dorsum is
light brown to tan (somewhat paler on the sides) with chocolate brown
dorsal markings the most constant of which are an interocular bar and







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


a scapular chevron. The venter is brown to pinkish-gray, and the con-
cealed surfaces are tan, heavily marbled with brown and occasionally
suffused with reddish brown. The discs are large, as in probolaeus,
the voice was reported by Thomas to be a "short, whispering trill of 2 or
3 notes, repeated several times." Diurnal retreats include rock and palm
trash piles as well as isolated rocks, and at night the frogs were seen on
the ground in a Cocos grove. One frog was taken while on a leaf about
1 ft (0.3 m) above the ground in this same grove, but the species is
primarily terrestrial. Although E. monensis and E. probolaeus resemble
each other in several ways, they are distinct and are probably no more
closely related to each other than either is to, say, E. pictissimus or E.
alcoae.
Since Puerto Rico lacks a spotted member of the ricordi complex,
it seems much more likely that E. monensis has been derived from His-
paniola than from the richly-colored and unmarked Puerto Rican E.
richmondi. Schmidt (1926:153-154) allied monensis with an unnamed
Hispaniolan species (thus following Stejneger 1904:564) and also placed
it in a sequence of weinlandi-monensis-richmondi-lentus as a series of
vicarious species. Although Schmidt's sequence may not be correct, still
the relationships of these four species (and many others in the Greater
Antilles) are unequivocal.

SPECIMENS EXAMINED.-REPOBLICA DOMINICANA. LA ALTAGRACIA PROV.:
0.5 mi (0.8 km) NE Boca de Yuma (MCZ 43197-98, ASFS V964); 1 mi (1.6 km)
NW Boca de Yuma (ASFS V28948-49, ASFS V29050); 2 km NW Boca de Yuma
(ASFS V17610-12).

Eleutherodactylus warren new species
FIGURE 3
HOLOTYPE.-CM 54138, a gravid female, from vicinity of Palmiste, Ile de la
Tortue, Haiti, one of a series obtained by native collectors for C. Rhea Warren
between 15 and 27 August 1970. Original number ASFS V20253.
PARATYPES.-ASFS V20254-57, ASFS V20263-64, CM 54139, MCZ 84645-46,
USNM 194013-14, same data as holotype; USNM 194012, same locality as holotype,
native for C. R. Warren, 26 January 1968.
DIAGNOSIS.-A species of the Antillean Eleutherodactylus ricordi group char-
acterized by the combination of moderate size (snout-vent length: males to 26 mm,
females to 33 mm); dorsum yellow-brown to medium brown with black dorsal
markings consisting primarily of the remnants of an interocular bar and a scapular
chevron and some scattered dark markings on the back and (more heavily) sides,
adjacent to a pair of dorsolateral stripes that are colored similar to the dorsal ground
color; hind limbs concolor with dorsum and with concealed surfaces marbled
purplish-brown; crura with some vague grayish-brown crossbar remnants; venter
white; some dark brown stippling on throat; digital discs moderately expanded, that
of digit 3 equal to or slightly less than one-quarter size of tympanum; supra-axillary
and inguinal glandular areas apparently absent; tympanum small (X=males 2.4,
females 2.6); and tibia/snout-vent length (X 100) high (males 51.0-51.4, females
47.6-50.2).








SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


DIsTnRI TnoN.-Known only from lie de la Tortue off the northern Haitian coast
(Fig. 2).
DESCRIPTION OF HOLOTYPE.-A gravid female with following measurements (in
mm): snout-vent length 33.3, head length 12.1, head width 12.4, longitudinal
diameter of tympanum 2.6, longitudinal diameter of eye 4.2, distance from naris to
anterior corner of eye 3.4, femur 15.0, tibia 16.0, fourth toe 13.5, tibia/snout-vent
length (X 100) 48.0. Head width slightly greater than distance from snout to
posterior border of tympanum; snout truncate with nares conspicuous at anterior
ends of canthus rostralis; diameter of eye longer than distance from naris to anterior
corner of eye; interorbital space 3.5, less than diameter of eye; diameter of tympanum
much less than diameter of eye, distance from tympanum to eye equal to about
one-third diameter of tympanum. Fingers long, unwebbed, 3-4-2-1 in order of
decreasing length; subarticular tubercles well developed, prominent, white. Toes
relatively long, slightly webbed at their bases, 4-3-5-2-1 in order of decreasing
length; subarticular tubercles large, prominent, white. Heels overlap when femora
are held at right angles to body axis. Dorsum smooth to slightly wrinkled, snout
minutely tuberculate, upper eyelids tuberculate, one large non-glandular wart be-
tween the angle of the jaw, tympanum, and forelimb insertion; throat and venter
smooth, belly disc feebly developed. Dorsal surfaces of all limbs smooth to finely
rugose; posterior faces of thighs with low, small, pavement-like granules. Inguinal
and supra-axillary glandular areas absent. Tongue large, only very slightly nicked,
free behind, its greatest width equal to about one-half that of floor of mouth. Pre-
vomerine teeth in two long arched series (extending from just outside the lateral
margins of the choanae and separated from the choanae by a distance equal to about
one-eight of a choana) that are separated medially by a distance equal to about
three-quarters the diameter of a choana.
COLORATION OF HOLOTYPE.-Dorsum medium brown, overlaid with irregular
black markings, the most prominent being a narrow and much dissected interocular
bar and the remnants of a scapular chevron, as well as a few scattered dark blotches
and spots. Dorsal zone separated from more heavily dark-blotched sides by an
indistinct pair of dorsolateral stripes that are not colored differently from the dorsal
ground color. A black supratympanic crescent which sends a dark bar to the center
of the tympanum. Sides heavily blotched with black, particularly above the
forelimb insertion and behind the tympanum. Lores totally black, thus no dark
loreal stripes. Fore- and hindlimbs concolor with dorsum; forelimbs with irregular
dark and medium brown stippling and marbling; hind limbs with vague crural
mottling, the concealed femoral surfaces with a purplish-brown area, more or less
solid but with intrusions and spotting of the dorsal medium brown ground. Pes
irregularly mottled with dark brown. Venter white, throat and undersides of fore-
and hindlimbs darkly stippled with brown. Iris metallic bronze, suffused with red-
dish below.
VARIATION.-Three male E. warren have the following measurements (in mm)
(extremes and means): snout-vent length 24.7-25.5 (25.2); head length 9.5-10.4
(9.9); head width 9.7-10.2 (10.0); tympanum 2.2-2.7 (2.4); eye 3.0-4.1 (3.6);
naris to eye 2.8-3.0 (2.9); femur 12.0-12.5 (12.3); tibia 12.7-13.1 (12.9); fourth
toe 10.5-11.8 (11.3); tibia/snout-vent length (X 100) 51.0-51.4 (51.3). Nine
gravid and adult females measure: snout-vent length 30.0-33.3 (31.5); head length
11.2-12.1 (11.6); head width 11.6-12.4 (12.0); tympanum 2.5-2.7 (2.6); eye 3.7-4.5
(4.2); naris to eye 3.2-3.7 (3.5); femur 13.2-15.0 (14.1); tibia 14.7-16.0 (15.2);
fourth toe 12.6-13.7 (13.1); tibia/snout-vent length (X 100) 47.6-50.2 (48.5). The
series of paratypes includes one subadult specimen (USNM 194012) with a snout-
vent length of 19.8 mm.
As noted in the diagnosis, the dorsal ground color varies between yellow-brown
and medium brown, with dorsal black markings. The black markings consist prin-
cipally of a narrow interocular bar, or its remnants, and a scapular chevron, or its
remnants. One individual (ASFS V20264) shows the low extreme of dorsal dark


1976








BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


pigmentation, since it lacks an interocular bar and the scapular chevron is re-
stricted to a pair of small discrete spots; the dorsum otherwise has no pattern.
The dorsolateral stripes are more or less evident in all specimens, but they are not
contrastingly colored in comparison with the dorsal ground color. The area between
the stripes is variously marbled, blotched, or stippled with dark pigment, but never
so much that the scapular chevron is obscured. The sides below the dorsolateral
stripes are usually more heavily pigmented with dark blotches than the dorsum itself,
with a supratympanic crescent and a large dark area above the forelimb insertion
commonly present. The venter is white with fine and uniform dark brown stippling
on the throat and the undersides of the limbs. The iris in all specimens is as
described for the holotype. No specimen shows supra-axillary or inguinal glandular
areas; these may be truly absent, but in many species they are obscured by dark
lateral pigmentation, which may be the case in E. warren.
COMPARISONS.-E. warren differs from E. weinlandi by lacking a heavy dorsal
dark pattern, although E. w. paralius resembles E. warren in the general absence
of a dorsal pattern. The two species are easily distinguished in life by the brighter
colors in the sacral blotch area and on the hind limbs in weinlandi. In addition,
warren has expanded digital discs, which are feebly developed or absent in
weinlandi. This same character will also separate E. warren from E. pictissimus.
E. warren differs from both E. alcoae and E. probolaeus in having small (versus
expanded) digital discs, a much smaller tympanum than alcoae, and a much different
pattern than probolaeus. E. warren is much smaller in size than alcoae, as is also
apparently true of probolaeus (female probolaeus are unknown).
Remarkably, E. warren is closest in coloration and pattern to E. monensis. The
latter species has slightly larger digital discs than E. warren and also has a supra-
axillary glandular area, which is absent in warren. E. monensis reaches a larger
size (males to 26.4, females to 37.5) than E. warren (males to 25.5, females to
33.3), and the tibia/snout-vent length means are higher in warren (males 51.3,
females 48.5) than in monensis (males 47.9, females 44.5). Although the two
species are quite close in many respects, I regard them as separate species rather
than subspecies, primarily because of the wide separation (460 km) between the
two islets upon which they occur. The problem of their relationships, as well as
that of warren to other Hispaniolan members of the complex (weinlandi, pictis-
simus), is compounded by the specimens noted below.
E. warren is named in honor of C. Rhea Warren, whose collecting activities on
Ile de la Tortue have been responsible for our greatly increased knowledge of the
herpetofauna of that island. Only our other Eleutherodactylus (E. inoptatus Bar-
bour) is known to occur on Tortue; however, that species is very large and thus
there is little possibility of confusing the two.

THREE ANOMALOUS SPECIMENS
There remain for discussion three extremely puzzling specimens.
Each is a subadult female from a separate locality, so that a series of
specimens is unavailable for analyses.
1) The first of these specimens (ASFS V1258) is from 19 km SE
Martin Garcia, 600 ft (183 m), Santiago Rodriguez Province, Repiblica
Dominicana (see Fig. 1 for localities of anomalous specimens). I had
earlier (Schwartz, 1965:113-114) considered this specimen to be E.
pictissimus from a locality far removed from the balance of the range
of the species and separated from southern E. pictissimus by an inter-
vening population of E. w. weinlandi. I see no reason at this time to
change my assignment. The specimen in life was pale, with a prominent







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


scapular chevron and prominent dorsolateral stripes that were not
concolored to the tan dorsal ground color. In most respects the frog
resembles E. p. apantheatus, but its locality is far removed from the
known range of that subspecies and separated from it by the intervening
high Cordillera Central. We made a special effort in 1971 to collect
both at the locality where the frog was taken (along a deeply entrenched
stream with gallery forest in an otherwise xeric area) and an area along
the Rio Guarabo, which is nearby and close to Los Quemados. The
Guarabo is a large river with extensive rich gallery forest traversing
desert scrub. At neither locality did we find even one more frog of this
type, despite optimal habitat and weather conditions. The fact is in-
escapable that there is a population of E. pictissimus in the northern
Rep6blica Dominicana (I reject the hypothesis that this single froglet
is a fortuitous introduction), but only further intensive and extensive
collecting in that region may reveal its taxonomic status.
2) The second frog (ASFS V22430) is from Thomonde, 400 ft (122
m), D6pt. de l'Artibonite, Haiti, collected by Lewis D. Ober in a crab
hole beneath very high grass at the edge of a small stream which flows
into the Riviere Thomonde. The locality lies some 25 km northwest
of a known locality for E. w. weinlandi (8.0 mi. E Lascahobas). The
dorsum is pale tan, and there is a black dorsal pattern (of the general
E. weinlandi type), with bold dorsolateral stripes, that are still lighter
(rather than brighter) than the dorsal ground color. The concealed
surfaces are violet, overlaid with a black reticulum; the crura are marbled
with grayish to brownish. There is no bright sacral area (as is typical
of weinlandi, and the hindlimbs are not brightly colored (also as in
weinlandi). It may well be that the Thomonde specimen is merely a
peculiarly patterned E. weinlandi, but it differs in so many features
from typical E. weinlandi that I am reluctant to make this assumption.
The two localities are separated by the Riviere de 1'Artibonite, and it is
possible that this large river separates two distinctive subspecies of E.
weinlandi in this region. Without further specimens from north of the
river, no further comments are possible.
3) The third specimen (ASFS V10223) is from Cap-Haitien, D6pt.
du Nord, Haiti, collected by Richard Thomas from a pile of rocks on a
more or less mesic hillside. The dorsal ground color is tan, and the
dorsolateral stripes are prominent and an even paler tan (just as in the
Thomonde specimen). The ground color of the sides and femora is
gray and that of the crura tan. There is no brightly colored sacral
blotch area. The dorsum is heavily marked with black vermiculations,
and the scapular chevron is obvious, despite the densely patterned dor-
sum. In many ways, this specimen strongly resembles the Martin Garcia


1976







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


specimen, the locality of which is about 100 km away. The Cap-
Haitien locality is 90 km from Thomonde, the third locality for anoma-
lous specimens.
An especially puzzling aspect of the Cap-Haitien specimen is that it
lies between Anse a Margot and Cruce de Guayacanes, both of which are
known localities for E. w. weinlandi. If the Cap-Haitien specimen is
anything other than E. w. weinlandi (and it does not agree with my
concepts of that taxon), the implication is that E. pictissimus, or some
other ricordi group species, occurs along this northern coastal area.
None of these specimens bear any resemblance to E. warren; this makes
the situation even more complicated, since the species whose range on
the mainland comes closest to Ile de la Tortue is E. weinlandi (Anse a
Margot). This northern Haitian littoral is so poorly known, especially
as far as its anuran fauna is concerned, that only by more detailed
collecting will we be able to understand the relationships of the frogs
in this northwestern area.

DIscussIoN
The five species (pictissimus, weinlandi, alcoae, probolaeus, warren)
discussed in this paper form a complex of species within the ricordi
group of Antillean Eleutherodactylus. Although these species seem more
closely related to each other than to any other Hispaniolan species,
they nevertheless show a degree of variation in both pigmental and
structural characters that suggests they represent part of an even larger
complex of species in the West Indies. The fact that Mertens (1939:30)
used the combination Eleutherodactylus lentus weinlandi, thereby com-
bining E. weinlandi with the Virgin Islands E. lentus, suggests a close
phylogenetic relationship of weinlandi to the more eastern species.
Shreve and Williams (1963:333) also noted that the relationships of
pictissimus and weinlandi to the Cuban forms (ricordi and bresslerae)
were difficult to evaluate, as was evident especially from their statement
that bresslerae "looks strikingly like weinlandi." They suggested that
pictissimus and weinlandi might well be conspecific, since the only
differences between them involved details of color and pattern. Study
of recently collected material of these two taxa has done nothing to
change the original picture, and there is still a very real possibility that
pictissimus and weinlandi might best be regarded as subspecies. This
association becomes even more likely with the removal of E. probolaeus
from E. pictissimus.
I have seen no evidence of intergradation in color or color pattern
between pictissimus and weinlandi, which would serve as an argument
against a subspecific relationship. On the other hand, that their geo-






SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


graphic ranges do not overlap could be viewed as a counter-argument.
I have the impression that E. pictissimus generally is far more tolerant
of xeric conditions than E. weinlandi. E. pictissimus occurs in the ex-
tremely hot and dry Valle de Neiba, probably in the equally hot, dry,
and contiguous Haitian Cul de Sac Plain, and in the Llanos de Azua.
In these xeric regions, pictissimus avoids the general desert areas, an
attribute that is necessary for the successful occurrence of any anuran
in such hostile habitats. E. weinlandi, on the other hand, is partial to
very mesic situations, and (except for the situation noted below) I
know of no records of this species outside of moist-to-wet forest or
pseudoforest (coffee and cacao groves, Cocos groves, etc.). The ex-
ception is the area along the southern Dominican coast that is inhabited
by E. w. paralius. This region is less moist than those usually occupied
by E. weinlandi; on the other hand, this coastal area is still well forested,
with much of it being underlain by pocked limestone, a situation that
favors occupation by terrestrial frogs like E. weinlandi. If the habitat
preferences and tolerances of the two species form a continuum, from
very xeric to very mesic, then E. pictissimus occupies the lower end of
this spectrum and E. weinlandi the upper, with some overlap of toler-
ances of both species toward the central portion of the continuum.
None of these data implies de facto separation of weinlandi and pictis-
simus at the species level, but they do suggest that these two taxa are
adapted for quite different sorts of habitats.
Disregarding for the moment the peculiar northern mosaic of locali-
ties for E. pictissimus and E. weinlandi in Haiti and the Rep6blica
Dominicana, the two species approach each other without dilution of
characteristics in two general areas. E. weinlandi is the frog of the
Montagnes du Trou-d'Eau and its associated ranges in Haiti and the
Sierra de Neiba in the Repiblica Dominicana. These ranges form the
northern edge of the Cul de Sac-Valle de Neiba plain, at least the latter
of which is occupied by E. pictissimus. The nearest approximations
of these two species in this region is 12 mi N Port-au-Prince (weinlandi)
and Port-au-Prince (pictissimus), and at 6.5 km W Duverg6 (pictissimus)
and 6.7 mi E Hondo Valle (weinlandi), an airline distance of 37 km.
Between the last two localities lies the high range of the Sierra de Neiba,
which reaches an elevation of 7416 ft (2261 m).
The second area of geographic approximation of the two species lies
between Peravia Province and the Distrito Nacional. E. pictissimus
occurs 4.8 mi S Bani, and E. weinlandi is known from 17 km NW Santo
Domingo; the intervening distance is 47 km airline. It should be re-
called that in both this and the previously detailed approximation, the
subspecies involved are E. w. weinlandi and either E. pictissimus apan-






BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


theatus or E. p. eremus. The nominate subspecies of E. weinlandi is
dark and colorful, and contrasts sharply with the pale and vaguely pat-
terned subspecies of E. pictissimus. The region about Bani is ecologi-
cally a transitional region between the xeric Llanos de Azua to the west
and the more mesic southern portion of the Republica Dominicana. The
area represents the eastern range limits for some species (Ameiva line-
olata Dum6ril and Bibron, Leiocephalus semilineatus Dunn, and Sphae-
rodactylus brevirostratus Shreve), and in other cases is the area of
intergradation for several subspecies (western xerophilic Ameiva chry-
solaema boekeri Mertens and eastern mesophilic A. ch. procax Schwartz
and Klinikowski; western Diploglossus stenurus weinlandi Cope and
eastern D. s. rugosus Cope; western Anolis distichus ravitergum Schwartz
and eastern A. d. ignigularis Mertens). Transitional areas such as this
may also serve to separate species or subspecies in other cases. The
intervening region between the known records of weinlandi and pictissi-
mus (primarily southern San Crist6bal Province) comprises mesic low-
lands and uplands, and appears to be much more suitable for E. wein-
landi than E. pictissimus. Collecting in this intervening area still has
yielded no specimens of either taxon.
The above discussion deliberately ignores the problem inherent in
the three widely-spaced localities in northern Haiti and the northern
Republica Dominicana of what I interpret as E. pictissimus (or at least a
species more closely allied to E. pictissimus than to E. weinlandi). Since
these northern localities of pictissimus are rather effectively separated
from the parent population of pictissimus by the intervening occurrence
of E. weinlandi in the Montagnes du Trou-d'Eau and Sierra de Neiba,
there seems to be no reasonable way to associate them geographically
with southern E. pictissimus. These northern frogs may have reached
their respective areas via the xeric Valle de San Juan, which extends
northwestward from the Llanos de Azua (occupied by E. p. eremus)
to the Dominico-Haitian border at BAnica-Pedro Santana on the Rivibre
de l'Artibonite, and thence (in Haiti) onto the Plateau Central (specifi-
cally the Plateau de Hinche). The Thomonde locality for E. pictissimus
lies within the Plateau Central. The proposed history of E. pictissimus
in this area suffers from the fact that, despite extensive collecting in the
Valle de San Juan, we have never found either E. pictissimus or E.
weinlandi; the Plateau Central in Haiti is one of the least known regions
of that country, and negative evidence for occurrence there of E. pictis-
simus cannot be taken as fact.
The two other northern localities of E. pictissimus are less easily
explained. There seems today to be no pathway for the species to have
reached Cap-Haitien in Haiti and the Martin Garcia region in the







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


northern Repiblica Dominicana. The situation is made even more com-
plex by the fact that both these places lie between two known localities
for E. weinlandi-Anse a Margot and north of Cruce de Guayacanes.
It is possible that the Cap-Haitien and Martin Garcia localities were
invaded from the south along the deeply entrenched valley of the Rivi-
6re de l'Artibonite, and thence into the watershed of the Rio Massacre,
from which the frog may have spread both to the east and west, there
occupying xeric areas that were unsuitable for the already established
E. weinlandi. This is pure conjecture-there are no specimens of either
species from this presumed area of invasion, and indeed the species may
no longer occur there. Certainly the border area from south of Villa
Anacaona to Monte Cristi seems much more suitable ecologically for
E. pictissimus than E. weinlandi, but we have never taken either species
of this pair in several months' collecting in this region.
Mertens (1939) first proposed that the Hispaniolan fauna might be
catalogued into two old faunas that were localized on the two islands
(north and south), which have subsequently been fused to form the
present island of Hispaniola along the Cul de Sac-Valle de Neiba plain.
This basic tenet has been amplified by Williams, Thomas, and myself,
and there is little doubt that the present low-lying (and in places below
sea level) plain separates two rather different herpetofaunas. There has
been some faunal interchange since the closure of the paleostrait (and
probably before it) resulting from chance trans-strait dispersal by some
tolerant species. Invading species have had varying success once they
reached the other island. I have no doubt that E. pictissimus is a south
island species, since it occurs throughout that region, except at higher
elevations in the Sierra de Baoruco and from sea level to an elevation
of 5800 ft (1769 m) at Furcy. E. pictissimus invaded the north island,
where it gave rise to a distinctive subspecies (eremus), whose range
extends as far east (Bani) as the limits of suitable xeric habitat permit.
Apparently E. pictissimus also invaded much of the northern areas of
Haiti and the Repdblica Dominicana, but perhaps with less success; the
records are too few and scattered to generalize.
E. weinlandi, on the other hand, is completely a north island species,
occupying mesic regions primarily in the eastern part of Hispaniola. With
few exceptions, the range of E. weinlandi is fairly compact. The Anse
h Margot record is one exception. The major exception, however, in-
volves the population in the Montagnes du Trou-d'Eau and the Sierra
de Neiba, an area that is some 80 km disjunct from the next closest E.
weinlandi locality (Vallejuelo and Jarabacoa). As I previously pointed
out, the logical area for continuity between the Sierra de Neiba and more
eastern populations of E. weinlandi is not across the Cordillera Central


1976







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


between Jarabacoa and Vallejuelo, but rather along the southern slopes
of the Cordillera Central between La Cumbre and San Juan. Unfor-
tunately, these regions are poorly known herpetologically. As far as
the isolated Anse a Margot record in northern Haiti is concerned, I
suspect that E. weinlandi will be found to occur along the northern
flanks of the Cordillera Central (where it is already known from the
Rio Bao near Los Montones), and thence into the Massif du Nord in
northern Haiti. The distribution of E. weinlandi in this region may no
longer be continuous, but this seems a likely avenue for invasion from
the east of the northern Haitian littoral.
In order to place E. weinlandi and E. pictissimus in their proper
perspective within the ricordi complex in the Greater Antilles, the mor-
phological characters (presence of supra-axillary and inguinal glands,
presence of postfemoral glands, presence of throat glands, development
of digital discs) of these two species and the other with which they are
associated have been summarized in Appendix 3. Measurements for all
taxa included in this study are given in Appendix 1 (for the Hispaniolan
and Mona taxa) and Appendix 2 (for the non-Hispaniolan taxa). In
addition, the following species are known to be vocal: weinlandi, al-
coae, probolaeus, richmondi, and monensis. The other species are pre-
sumably mute, although pictissimus and thomasi have been assumed to
make sounds. I have had no field experience with warren, but Mr.
Warren reported no vocal activities on the part of this frog. E. bres-
slerae has been collected only once (under very favorable moisture
conditions) but was not heard calling; E. ricordi has been collected
often but has never been heard vocalizing. My experience with E. lentus
is limited to St. Thomas, which at the time of my visit was very dry.
Grant (1937:505) did not mention calling lentus on St. Croix, nor did
Richard Thomas and David C. Leber who collected the species there.
Examination of Appendix 3 shows that of the included 11 species,
the Puerto Rican E. richmondi is most distinctive. Although the species
lacks supra-axillary, inguinal, or postfemoral glandular areas, males are
unique in having a large single (or somewhat divided) glandular area
on the throat. E. richmondi also differs from other members of this
group in having basically a brightly-colored (rich brown, reddish tan,
bright chestnut) dorsum, often set off laterally by a pair of fine golden
to yellow or cream dorsolateral stripes that separate the dark brown
sides (see Schmidt 1928 fig. 20, for an excellent delineation of this
species). Like E. weinlandi, E. richmondi has a reddish or rusty sacral
area, but unlike weinlandi, richmondi has an unmarked (versus pat-
terned) dorsum. E. richmondi is vocal and lacks well developed digital
discs. In addition, the species is a mesophile and almost exclusively







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


terrestrial, although I have taken calling males from low shrubs (1 ft
[0.3 m]) and other individuals on grass and leaves of herbs 6 in (15 cm)
above the ground. By taking the above characteristics into considera-
tion, I cannot consider E. richmondi a central member of the frog group
under discussion; rather it seems to be a peripheral member of this com-
pact group of frogs that is in some ways specialized when compared
with other members of the complex.
The remaining members of the group fall into two major categories:
(1) those with well developed digital discs, and (2) those with the discs
poorly developed or absent. In this complex I consider presence of well
developed discs primitive and their absence specialized, as appears
to be true of the entire genus, in fact, since discs are regularly well
developed in the Antillean members of the auriculatus group (see
Schwartz 1969). This primary dichotomy separates those species with
well developed discs (ricordi, thomasi, alcoae, probolaeus, bresslerae,
monensis) from those with the discs poorly developed or absent (lentus,
weinlandi, pictissimus, warreni.
Tracing the postulated history of each of these lines further (see Fig.
4 for a graphic representation of these relationships), I regard the pres-
ence of three glandular areas (supra-axillary, inguinal, postfemoral) as
primitive, and loss of any of these as specialized. Of the disced mem-
bers, only ricordi occasionally shows postfemoral glands, whereas all
other members lack them. A third dichotomy in this sequence is the
absence or presence of supra-axillary and inguinal glands. E. bresslerae
lacks these glandular areas, and monensis has them reduced. Supra-
axillary and inguinal glands are present in probolaeus, alcoae, and
thomasi (occasionally absent in thomasi).
In the non-disced sequence, only lentus has all three glandular areas
present, and thus its relationships to the non-disced species are com-
parable to those of ricordi to the disced species. The postfemoral
glands have been lost in all other non-disced species. E. warren has
lost both supra-axillary and inguinal glandular areas, but pictissimus and
weinlandi have retained these two areas. Thus, within the disced and
non-disced series, there is a progressive loss of glandular areas.
Examining these two lines of divergence within the group in the light
of geography, we find the following. The primitive members of each
line (lentus, ricordi) occur either on islets that are far removed from
the balance of the group (lentus on St. Thomas, St. John, and St. Croix
in the Virgin Islands, east of Puerto Rico), or they have a restricted
and primarily upland distribution (ricordi only in the uplands of the
Sierra Maestra, Sierra de la Gran Piedra, and the Cuchillas de Toa in
Oriente Province in eastern Cuba) (Fig. 5). Of the non-disced species,







36 BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1
PICTISSIMUS WEINLANDI PROBOLAEUS ALCOAE

WARREN BRESSLERAE THOMASI
/ ONENSIS ----
l no glands all glands present
all glands present /\
no glands no postfemoral glands RICORDI
LENTUS
no postfemoral glands


discs reduced or absent discs present


RICHMONDI
discs present 7
no discs or glands
throat glands in males



FIGURE 4.-Dendrogram showing suggested relationships of 11 species of Antillean
Eleutherodactylus, as discussed in text. The dashed lines to monensis indicate the
problematical relationships of that species, which is closer to the line with discs re-
duced or absent than to that with discs present. The relationship to warren seems
more logical than that to bresslerae.


all except lentus are from Hispaniola (including warren on Tortue).
The fact that warren occurs on a Hispaniolan satellite and lacks glandu-
lar areas suggests this species has had a long independent history from
gland-bearing pictissimus and weinlandi on the mainland. There is no
serious problem with this distributional picture except for the remote
position of primitive lentus. This series' proposed history would be more
satisfying were there a certain member of the complex on Puerto Rico,
which lies between the Virgin Islands and Hispaniola. It is possible
that richmondi is that member, but it appears too highly specialized
to be derived or derivable from lentus. That the Virgin Islands have
had a history both allied to, but different from, that of Puerto Rico is
unquestioned, and these islets have many species of amphibians and
reptiles in common with Puerto Rico. On the other hand, they have
several species that are restricted to and very distinct from the Puerto
Rican herpetofauna, including Amphisbaena fenestrata Cope, Sphaero-
dactylus parthenopion Thomas, and (if we include the islets just east
of Puerto Rico with the Virgin Islands) Anolis roosevelti Grant. Thus
we need not expect complete similarity between the Virgin Island and












7o






2 .- 1 --22-








20"-

20' 70



FIGURE 5.-Map of Cuba, showing localities for three species of Eleutherodactylus as follows: triangles, E. ricordi; squares E.
bresslerae; circles, E. thomasi. Subspecies of E. thomasi are indicated by TH (E. t. thomasi); TR (E. t. trinidadensis); and ZA 0
(E. t. zayasi). Frogs from the westernmost four localities of E. t. thomasi show intergrading tendencies between that subspecies (
and E. t. trinidadensis.







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


Puerto Rican herpetofaunas. It seems probable that lentus is a primi-
tive relict species that has persisted on three of the many Virgin
Islands, but whose close Puerto Rican relatives have since disappeared.
Presence of some member of this complex on Puerto Rico would also
enhance the probability that the Hispaniolan members of the group were
derived from Puerto Rico; although I feel certain that this was the case,
the intermediate Puerto Rican member of the group apparently no
longer exists.
A proposed history of this frog group's disced members is some-
what more difficult and puzzling. Assuming that ricordi is the most
primitive of the disced species, there is no problem in accounting for the
presence of two disced species (bresslerae, thomasi) on Cuba. Of these
two species, thomasi is more widely distributed, occurring from Ma-
tanzas Province in the west to Camagiiey Province in the east (Fig. 5).
The Matanzas population (E. t. zayasi) is apparently disjunct, and E. t.
trinidadensis in the Sierra de Trinidad in southern Las Villas Province
also appears to be a geographic isolate. Only in the Sierra de Cubitas,
the Sierra de Najasa (E. t. thomasi) and the mountains near the Las
Villas-Camagiiey border does there appear to be any continuity among
the known records from E. thomasi. Interestingly, I regard the material
from the Las Villas-Camagiiey line (Cueva de Caguanes. Yaguajay)
and from Loma de Cunagua, near Mor6n in northwestern Camagiiey
Province, as intermediate between thomasi and trinidadensis, although
closer to the former in both characters and geography. Thus the
distribution of E. thomasi seems to be discontinuous. This may be due
partly to lack of suitable habitat in some of the intervening regions
(thomasi is associated with forested limestone areas) and partly to re-
traction of distribution coincident with changing ecology.
E. bresslerae has a very limited distribution in extreme eastern Cuba;
the species is known from two stations in the Cuchillas de Toa (Fig. 5),
the complex montane mass that occupies much of the eastern interior of
Cuba. Although there are only two records of ricordi from these same
mountains, it is probable that bresslerae and ricordi are sympatric
throughout much of this area. In general, ricordi elsewhere is an upland
species, whereas bresslerae is known from at least one coastal (but well
forested) locality.
The two Hispaniolan disced members (alcoae and probolaeus) may
be interpreted as peripherally derived from the Cuban species, and, in
fact, probolaeus resembles bresslerae in dorsal pattern details. The
restricted distributions of alcoae and probolaeus suggest that as species
they are either quite recently derived or have distributions increasingly
restricted. The latter seems much more likely. This implies there was an






SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


early invasion of disced frogs from Cuba that became adapted to life in
limited areas which were unoccupied by already resident species. This
almost certainly is the situation with alcoae, which inhabits the rigorous
Peninsula de Barahona. The problem of probolaeus is somewhat differ-
ent, because it seems not to occur in large areas of eastern Hispaniola
that appear suitable and unoccupied by any member of this complex.
Both alcoae and probolaeus are associated with limestone areas.
E. monensis poses the only serious problem with my proposed se-
quence of disced forms. As I previously pointed out, this species can-
not logically be derived from any extant Puerto Rican species (rich-
mondi) and is much more plausibly associated with a species from
Hispaniola. Since monensis has the glandular areas reduced, it seems
most likely to be in the line of evolution that led to bresslerae. How-
ever, these two species are quite dissimilar (see Schwartz 1960:fig. 6
for bresslerae, Schmidt 1928:fig. 21 for monensis), and I find it very
difficult to associate them. Of all the involved species, monensis most
closely resembles warren in general aspect. Although discs are present
in monensis, they are reduced and only slightly larger than those of
warren. Perhaps the logical explanation is that monensis and warren
represent insular remnant populations of an otherwise extinct primitive
disced species.
Whatever the history of such peculiar species as E. monensis and
E. warren, it seems evident that Hispaniola has been invaded twice
by members of this complex of frogs. One invasion of non-disced
members came from the east and resulted in weinlandi on the old His-
paniolan north island and pictissimus on the south island. The second
invasion was from Cuba in the west, where all members of this group
have enlarged digital discs. The second invasion resulted in two species
(alcoae, probolaeus), one each on the north and south islands, and both
presently have very limited distributions.
Members of this complex are all moderate- to large-sized frogs of
the ricordi group. Within the complex, the largest males vary in snout-
vent lengths (rounded off to the nearest mm) between alcoae (38) and
monensis and warren (26). Between these two extremes are ricordi
(35), pictissimus (34), thomasi (32), weinlandi, probolaeus, and bres-
slerae (30), and lentus (28). Female size extremes vary from bresslerae
(46) to warren (33), with alcoae (45), pictissimus and thomasi (43),
ricordi and weinlandi (40), monensis (38), and lentus (37) being of
intermediate size.
The most striking sexual differences in maximum snout-vent length
(in mm) are shown by bresslerae (16) and monensis (12), and the
least sexual dimorphism in warren (7) and ricordi (5). Generally, the







BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


non-disced members shows less sexual dimorphism (between 7 and 10)
than do the disced members (8 to 16). In the latter group E. ricordi
is exceptional, because the largest members of each sex differ by only
5 mm. These data suggest that sexual dimorphism is greater in the
(primitive) disced members of the complex (with the exception of
ricordi), and less pronounced in the more specialized discless members.
It is interesting that the problematical monensis has a sexual dimorphic
difference of 12 mm, greater than that of any discless member species
and within the range of the species with discs.
Comparison of the two groups' dorsal coloration and pattern shows
the following. In those species with discs, dorsolateral stripes are absent
in ricordi, alcoae, and monensis and present, but unicolor with the dorsal
ground color, in bresslerae, probolaeus, and thomasi. In thomasi stripes
are variable in the nominate subspecies and E. t. trinidadensis, but
occur in all E. t. zayasi. In the non-disced members, dorsolateral stripes
are absent in warren, present and unicolor with the dorsum in pictissi-
mus and lentus, and present but brightly colored in weinlandi. Thus,
dorsolateral stripes are absent in the advanced forms (bresslerae, probo-
laeus, alcoae, thomasi) and the two presumably primitive species (ricordi,
monensis) among the disced group, are present in all non-disced species
except warren (including the basal lentus), and are brightly colored as
well as present in weinlandi.
As pointed out in the introduction, this entire frog assemblage has
one of two dorsal colors and patterns. One group has generally pale
dorsa, more or less heavily overlaid with a dark pattern that includes a
scapular chevron and an interocular bar (although these elements may
be greatly obscured by additional dark dorsal patterning). To this
group belong (among the species without discs) warren, pictissimus,
and lentus, as well as all members of the group possessing discs. E.
lentus tends to have brighter (yellow to golden or orange tan) dorsa
than other members of the species without discs and thus tends toward
weinlandi in dorsal color and pattern. Of this latter group, bresslerae
tends to be brighter dorsally, especially posteriorly. The second style
of dorsal coloraton is shown by weinlandi, which has a bright dorsal
ground color that may be heavily overlaid with dark brown to black,
as in the subspecies chersonesodes. The clear and bright sacral blotch
of weinlandi shows some resemblance to the brighter dorsal colors in
bresslerae.
In the above summary, I have attempted to indicate the relationships
between this complex of 10 Greater Antillean species of the Eleuthero-
dactylus ricordi group. They form a compact and obviously closely
related complex of species within that group of Eleutherodactylus that







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


is represented by the most species in the West Indies. I have had the
advantage of seeing all but two of these species warrenn, monensis)
in the field, and I have handled living warren. Such familiarity with
Eleutherodactylus in the field is essential in understanding their re-
lationships, because only by knowledge of the colorations, habits, voice,
and patterns of these frogs in life can one gain even partial insight into
their relationships.





LrrERATURE CrrED

Barbour, T. 1914. A contribution to the zoogeography of the West Indies, with
especial reference to amphibians and reptiles. Mem. Mus. Comp. Zool., 44(2):
209-359, 1 pl.
Cochran, D. M. 1935. New reptiles and amphibians collected in Haiti by P. J.
Darlington. Proc. Boston Soc. Nat. Hist., 40(6):367-376.
Dunn, E. R. 1926. Additional frogs from Cuba. Occ. Pap. Boston Soc. Nat. Hist.,
5:209-215.
Grant, C. 1937. Herpetological notes with new species from the American and
British Virgin Islands, 1936. Agri. Univ. Puerto Rico, 21(4):503-522, 4 pls.
Mertens, R. 1939. Herpetologische Ergebnisse einer Reise nach der Insel His-
paniola, Westindien. Abh. senckenberg. naturf. Ges., 449:1-84, 10 pls.
Schmidt, K. P. 1926. The amphibians and reptiles of Mona Island, West Indies.
Field Mus. Nat. Hist., Zool. Ser., 12(12):149-163, 6 figs.
1928. Scientific survey of Porto Rico and the Virgin Islands. New
York Acad. Sci., 10(1):1-160, 52 figs., 4 pls.
Schwartz, A. 1958. Four new frogs on the genus Eleutherodactylus (Leptodactyli-
dae) from Cuba. Amer. Mus. Novitates, 1873:1-20, 4 figs.
.1960. Nine new Cuban frogs of the genus Eleutherodactylus. Read-
ing Public Mus. Art Gallery (Sci. Publ.), 11:1-50, 6 figs.
.1965. Geographic variation in two species of Hispaniolan Eleuthero-
dactylus, with notes on Cuban members of the ricordi group. Stud. Fauna
Curaqao and Caribbean Is., 22(86):98-123, 7 figs.
1969. The Antillean Eleutherodactylus of the auriculatus group.
Stud. Fauna Curagao and Caribbean Is., 30(114):99-115.
.1971. A new species of Eleutherodactylus (Amphibia, Leptodactyli-
dae) from Hispaniola. Ann. Carnegie Mus., 43(2):25-31, 1 fig.
Shreve, B., and E. E. Williams. 1963. The herpetology of the Port-au-Prince
region and Gonave Island, Haiti. Part II. The frogs. Bull. Mus. Comp. Zool.,
129(5):302-342, 5 pls.
Stejneger, L. 1904. The herpetology of Porto Rico. Report U. S. Natl. Mus.,
1902, pp. 549-724, 197 figs.


1976








BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


APPENDIX 1.-NINE MEASUREMENTS (MEANS AND EXTREMES) OF SIX SPECIES OF
Eleutherodactylus FROM HISPANIOLA AND ISLA MONA.


1 N=number of individuals.
2 Measurements are of largest males.
3 Measurments are of gravid, adult females.


Taxon Sex
p. pictissimus M2

F3

p. apantheatus M

F

p. eremus M

F

w. paralius M

F

w. chersonesodes M

F

w. weinlandi M

F

warren M

F

alcoae M

F

probolaeus M

monensis M

F


Snouth-vent
length
26.6
(23.0-33.9)
35.7
(28.5-43.2)
24.9
(21.6-33.1)
32.0
(29.7-34.7)
25.0
(22.7-27.7)
31.0
(30.6-31.3)
25.6
(22.0-29.5)
35.4
(29.3-40.4)
23.8
(20.8-27.6)
29.8
(24.8-34.9)
21.7
(20.0-27.4)
32.6
(27.2-35.8)
25.2
(24.7-25.5)
31.5
(30.0-33.3)
33.1
(26.2-37.5)
38.6
(33.5-44.5)
27.9
(25.8-29.8)
23.4
(20.4-26.4)
33.1
(28.0-37.5)


Head
length
9.9
(8.7-12.5)
13.0
(10.8-16.1)
9.6
(8.3-12.5)
11.7
(10.6-12.6)
9.5
(8.5-10.7)
10.8
(10.3-11.4)
9.9
(8.7-11.7)
12.2
(10.9-14.8)
9.1
(7.7-11.0)
10.9
(9.4-13.8)
9.3
7.7-10.4)
11.5
9.1-13.2)
9.9
(9.5-10.4)
11.6
(11.2-12.1)
12.5
(10.1-14.2)
14.9
(13.7-16.6)
11.2
(10.3-12.7)
9.5
(8.2-10.8)
13.1
(10.3-15.2)


Head
width
9.6
(8.2-11.6)
12.9
(10.9-16.0)
9.8
(8.4-13.2)
12.0
(11.2-13.0)
9.7
(8.7-11.1)
11.3
(10.9-11.8)
9.7
(8.4-11.1)
13.2
(10.9-14.9)
8.9
(7.4-11.0)
10.7
( 9.3-13.7)
9.1
(7.1-11.0)
11.5
(9.0-13.2)
10.0
(9.7-10.2)
12.0
(11.6-12.4)
12.8
(10.9-14.2)
15.0
(13.6-17.0)
11.3
(10.5-12.0)
9.8
(8.5-11.1)
13.9
(11.5-16.1)


Tympanum
2.2
(1.8- 2.8)
2.7
( 2.2- 3.9)
2.2
(1.9- 2.8)
2.5
( 2.3- 2.8)
2.2
(1.9- 2.5)
2.4
( 2.0- 2.6)
2.4
( 2.2- 2.7)
2.9
(2.4- 3.3)
2.1
( 1.8- 2.5)
2.4
( 1.9- 3.1)
2.1
( 1.7- 2.5)
2.4
(1.8- 2.8)
2.4
( 2.2- 2.7)
2.6
( 2.5- 2.7)
3.1
( 2.6- 3.7)
3.4
(3.0- 4.1)
2.6
( 2.3- 3.0)
2.3
( 2.0- 2.7)
3.0
( 2.7- 3.6)







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


APPENDIX 1.-(CONTINUED)

Tibia/SV
Fourth length
Eye Naris-eye Femur Tibia toe (X 100)
3.4 3.0 11.2 12.8 11.2 48.9
( 2.9- 4.0) ( 2.4- 4.1) ( 9.7-14.5) (10.8-15.6) ( 9.2-14.4) (41.3-53.2)
4.3 4.1 15.5 17.3 15.5 46.2
( 3.4- 5.8) ( 3.0- 4.8) (12.2-19.8) (13.5-21.6) (12.2-19.2) (42.0-53.3)
3.4 3.1 11.3 12.2 10.9 48.9
( 2.8- 4.4) ( 2.5- 4.3) ( 9.7-16.2) ( 9.7-16.2) ( 9.2-14.8) (45.4-55.2)
4.0 3.7 14.0 14.5 13.3 45.4
( 3.6- 4.2) ( 3.2- 3.9) (13.0-15.2) (13.1-16.0) (12.2-14.6) (43.7-48.2)
3.4 3.0 10.0 11.9 11.6 47.0
( 2.8- 3.7) ( 2.4- 3.8) ( 9.7-12.5) (10.6-14.1) ( 9.8-12.8) (42.6-51.6)
3.9 3.6 13.0 13.7 12.3 44.1
( 3.5- 4.4) ( 3.5- 3.7) (12.5-13.4) (12.9-14.6) (11.9-12.7) (42.0-46.6)
3.4 2.9 11.1 12.0 9.6 46.7
(3.0- 4.0) ( 2.4- 3.6) ( 9.3-13.3) (10.6-14.6) ( 8.9-12.7) (44.1-50.5)
4.5 4.1 15.7 16.5 14.5 46.7
( 3.9- 5.0) ( 3.1- 4.7) (12.2-18.0) (12.2-18.0) (11.8-16.9) (44.8-50.6)
3.4 2.5 11.2 12.3 10.9 51.5
( 2.4- 4.1) ( 2.2- 3.1) ( 9.2-14.3) (10.1-15.1) ( 8.6-14.7) (47.4-55.7)
4.1 3.1 13.9 15.2 13.4 50.6
( 2.7- 5.1) ( 2.5- 4.2) (11.6-18.0) (13.0-18.3) (10.9-15.8) (45.4-56.8)
3.3 2.7 10.6 12.1 10.8 52.8
( 2.6- 3.8) ( 2.2- 3.4) ( 8.8-13.6) ( 9.6-15.1) ( 8.7-12.7) (41.9-55.1)
4.0 3.5 14.4 15.8 14.0 50.0
( 3.2- 4.8) ( 2.6- 4.2) (11.3-16.2) (12.5-17.8) (11.2-15.9) (45.1-54.3)
3.6 2.9 12.3 12.9 11.3 51.3
( 3.0- 4.1) ( 2.8- 3.0) (12.0-12.5) (12.7-13.1) (10.5-11.8) (51.0-51.4)
4.2 3.5 14.1 15.2 13.1 48.5
( 3.7- 4.5) ( 3.2- 3.7) (13.2-15.0) (14.7-16.0) (12.6-13.7) (47.6-50.2)
4.2 4.0 14.5 15.5 12.3 47.9
( 3.5- 4.9) ( 3.0- 4.9) (12.4-16.8) (12.7-17.8) (10.5-14.4) (41.8-51.6)
4.8 4.9 16.7 17.8 14.1 46.1
( 4.2- 5.6) ( 4.4- 5.5) (15.2-18.9) (15.9-19.4) (12.7-15.2) (43.6-47.5)
4.2 3.7 12.6 13.8 11.7 49.5
( 3.9- 4.5) ( 3.4- 4.2) (11.3-13.7) (12.8-15.2) (10.7-12.9) (47.0-51.7)
3.4 2.7 10.4 11.2 10.2 47.9
( 3.1- 3.8) ( 2.3- 3.1) ( 9.0-11.3) (10.1-12.0) ( 9.3-10.8) (44.3-50.0)
4.5 4.1 13.8 14.7 13.6 44.5
( 4.1- 5.0) ( 3.3- 5.0) (12.2-14.6) (12.8-16.1) (11.5-15.9) (41.4-49.2)


1976








BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


APPENDIX 2.-NINE MEASUREMENTS (MEANS AND EXTREMES) OF FIVE Eleuthero-
dactylus SPECIES FROM THE VIRGIN ISLANDS, PUERTO RICO, AND CUBA.

Snout-vent Head Head
Taxon Sex N1 length length width Tympanum
lentus M2 16 24.9 9.5 9.2 2.2
(21.8-27.9) ( 8.4-10.8) ( 7.4-10.7) ( 1.9- 2.5)
F3 10 33.6 12.1 12.4 2.7
(31.2-38.3) (11.2-13.9) (11.4-13.7) ( 2.4- 3.1)
richmondi M 25 27.7 10.7 10.4 1.9
(26.4-29.0) (10.3-11.5) ( 9.9-11.0) ( 1.5- 2.1)
F 17 39.6 15.1 15.2 2.7
(36.2-41.7) (13.2-16.1) (14.0-16.2) ( 2.1- 3.1)
bresslerae M 7 25.3 10.3 9.9 2.3
(21.9-30.4) ( 9.0-12.0) ( 8.3-11.6) ( 2.0- 2.8)
F 1 45.8 17.6 17.2 3.5
ricordi M 23 26.6 10.5 10.3 2.3
(19.6-34.6) ( 7.8-13.6) ( 7.5-13.4) ( 1.6- 2.8)
F 28 33.3 13.1 12.7 2.7
(25.4-40.0) ( 9.5-19.6) ( 9.0-15.0) ( 1.8- 3.2)
t. zayasi M 28 29.0 11.4 11.2 2.6
(26.9-31.8) (10.3-12.6) ( 9.9-11.9) ( 2.2- 3.3)
F 15 38.8 14.6 13.9 2.8
(37.5-40.9) (14.1-15.0) (13.2-14.7) ( 2.6- 3.2)
t. trinidadensis M 24 27.8 10.9 10.6 2.8
(24.8-31.1) ( 9.8-12.0) ( 9.5-11.9) ( 2.3- 3.3)
F 18 34.0 13.6 13.4 3.0
(31.5-39.5) (11.8-15.0) (11.5-14.6) ( 2.7- 3.2)
t. thomasi M 10 25.3 10.3 9.9 2.3
(24.5-26.3) ( 9.8-10.8) ( 9.2-10.4) ( 2.1- 2.4)
F 23 38.9 14.7 14.8 2.9
(31.9-43.4) (12.3-16.5) (12.4-16.7) ( 2.5- 3.3)

1 N=number of individuals.
2 Measurements are of largest males.
3 Measurements are of gravid, adult females.







SCHWARTZ: HISPANIOLA FROGS, RICORDI GROUP


APPENDIX 2.-(CONTINUED)

Tibia/SV
Fourth length
Eye Naris-eye Femur Tibia toe (X 100)
3.5 2.7 11.1 11.9 10.9 47.8
(2.8- 4.3) ( 2.2- 3.1) ( 9.6-12.2) (10.3-13.3) ( 9.7-12.0) (44.7-51.6)
4.4 3.7 14.7 15.5 14.2 46.2
(3.5- 4.9) ( 3.2- 4.4) (12.7-16.6) (14.3-17.8) (12.8-17.0) (42.4-49.3)
4.3 2.8 12.3 13.0 12.7 47.0
(3.9- 5.0) ( 2.4- 3.1) (11.3-13.2) (12.4-13.5) (11.7-13.6) (44.4-49.5)
5.5 4.3 17.8 18.9 18.0 47.8
(4.9- 6.0) ( 3.6- 5.5) (16.6-20.0) (17.6-21.8) (16.6-20.2) (44.6-51.4)
3.8 3.3 11.4 11.8 10.5 46.7
(3.0- 4.0) ( 2.6- 4.1) ( 9.7-13.1) (10.3-14.4) ( 8.7-11.9) (44.7-47.6)
6.0 6.1 20.4 21.3 18.1 46.5
3.8 3.4 12.3 13.3 12.3 50.2
(2.8- 5.0) ( 2.2- 4.6) ( 9.0-16.1) ( 9.1-18.7) ( 9.1-18.7) (44.4-54.0)
4.6 4.2 15.4 16.7 15.2 49.6
(3.6- 6.0) ( 3.1- 5.1) (11.6-19.0) (12.2-20.0) (12.1-18.5) (45.1-53.6)
3.9 3.7 13.3 14.3 12.1 49.2
(3.2- 4.4) ( 3.0- 4.6) (11.7-14.3) (13.3-15.5) (11.3-13.1) (46.5-52.1)
4.9 4.9 17.2 18.4 15.8 47.5
(4.5- 5.7) ( 4.4- 5.2) (16.1-18.4) (17.2-19.3) (14.0-17.2) (44.7-49.3)
3.7 3.4 13.0 13.1 11.7 47.3
( 3.3- 4.6) ( 2.7- 3.8) (10.7-13.3) (11.8-14.1) (10.2-12.5) (43.4-50.2)
4.7 4.4 15.5 16.3 14.3 45.3
( 4.1- 5.2) ( 3.5- 5.0) (13.4-16.9) (14.5-17.3) (12.3-15.5) (43.0-48.0)
3.7 3.4 12.2 12.4 11.0 51.1
( 3.0- 4.1) ( 3.0- 3.8) (11.5-13.0) (11.5-13.2) ( 9.6-11.7) (48.4-53.6)
5.1 5.1 17.8 17.0 15.9 48.9
(4.3- 6.0) ( 4.0- 6.1) (14.9-20.3) (16.5-22.2) (13.5-18.0) (45.8-52.7)


1976








BULLETIN FLORIDA STATE MUSEUM Vol. 21, No. 1


APPENDIX 3.-TEN SPECIES OF RELATED Eleutherodactylus SHOWING OCCURRENCE OF
STRUCTURAL FEATURES (GLANDULAR AREAS AND DIGITAL DISCS) AND
PRESENCE OF VOCAL ACTivrry.

Supra-axillary
and inguinal Postfemoral Throat Digital
Species glands glands glands discs Voicel


+
(inguinal only,
and rarely)
+

+
(supra-axillary
only,
and rarely)


Poorly
developed
Poorly
developed
Well
developed
Well
developed
+ Poorly
developed
+ Poorly
(males) developed
Well
developed
+ Well
(rarely) developed

Well
developed
Well
developed


- Moderately
developed


1"?" in this column suggests species may be mute (see text for comments).


pictissimus +


weinlandi


alcoae


probolaeus

lentus

richmondi

bresslerae

ricordi


thomasi

monensis



warren


?

Vocal

Vocal

Vocal

?

Vocal

?

?


?

Vocal














Contributions to the BULLETIN OF THE FLORIDA STATE MUSEUM, BI-
OLOGICAL SCIENCES SERIES, may be in any field of biology. Manuscripts deal-
ing with natural history of systematic problems involving the southeastern United
States or the New World tropics are solicited especially. Manuscripts should be of
medium length-circa 35 to 200 pages (10,500-16,000 words). Examination for
suitability is made by an Editorial Board.

The BULLETIN is distributed worldwide through institutional subscriptions and
exchanges. It is considered the responsibility of the author to distribute his paper
to all interested individuals. To aid in this the authors) receives(s) 50 copies free,
and he may purchase additional separates at cost if ordered when page proof is re-
turned. The author is also responsible for any charges incurred for alterations made
by him on galley or page proofs. The Museum will send an invoice to the author
for this amount upon completion of publication.


PREPARATION OF MANUSCRIPT

Contributors should consult recent numbers of the BULLETIN for preferred style
and format. Highly recommended as a guide is the CBE Style Manual, 3rd Edition,
1972 (Washington, D.C., Amer. Inst. Biol. Sci.).
MSS must be submitted in duplicate (please no onionskin) and satisfy the following
minimal requirements: They should be typewritten, double-spaced (especially tables,
figure captions, and "Literature Cited"), on one side of numbered sheets of standard
(8-1/2x11 in.) bond paper, with at least one-inch margins all around. Tables
(which should be unruled) and figure legends should be typed on separate sheets.
All illustrations are referred to as figures. They must comply with the following
standards: Photographs should be sharp, with good contrast, and printed on glossy
paper. Drawings should be made with dense black waterproof ink on quality paper
or illustration board. All illustrations should have a cover sheet. All lettering will
be medium weight, sans-serif type (e.g. Futura Medium, News Gothic) in cutout,
dry transfer or lettering guide letters. Make allowance so that after reduction no
lowercase letter will be less than 1 mm high (2 mm is preferred) nor any capital
letter greater than 5 mm high. The maximum size for illustrations is 8-5/8x14 in.
(twice typepage size); illustrations should not be less than typepage width (4-5/16
in.). Designate the top of each illustration and identify on the back with soft
pencil by author's name, MS title, and figure number.



Manuscripts and all editorial matters should be addressed to:
Managing Editor of the BULLETIN
Florida State Museum
Museum Road
University of Florida
Gainesville FL 32611







University of Florida Home Page
© 2004 - 2010 University of Florida George A. Smathers Libraries.
All rights reserved.

Acceptable Use, Copyright, and Disclaimer Statement
Last updated October 10, 2010 - - mvs