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Group Title: Bulletin of the Florida State Museum
Title: Sphaerodactylus (Gekkonidae) in the Greater Puerto Rico region
CITATION THUMBNAILS PAGE IMAGE ZOOMABLE
Full Citation
STANDARD VIEW MARC VIEW
Permanent Link: http://ufdc.ufl.edu/UF00001539/00001
 Material Information
Title: Sphaerodactylus (Gekkonidae) in the Greater Puerto Rico region
Series Title: Bulletin of the Florida State Museum
Physical Description: 195-260 p. : illus., maps. ; 23 cm.
Language: English
Creator: Thomas, Richard, 1938 May 2-
Schwartz, Albert, 1923-
Publisher: University of Florida
Place of Publication: Gainesville
Publication Date: 1966
 Subjects
Subject: Sphaerodactylus   ( lcsh )
Reptiles -- Puerto Rico   ( lcsh )
Reptiles -- Virgin Islands of the United States   ( lcsh )
Genre: bibliography   ( marcgt )
government publication (state, provincial, terriorial, dependent)   ( marcgt )
non-fiction   ( marcgt )
 Notes
Bibliography: "Literature cited:" p. 259-260.
General Note: Cover title.
Statement of Responsibility: by Richard Thomas and Albert Schwartz.
 Record Information
Bibliographic ID: UF00001539
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: ltqf - AAA0848
notis - ACK4303
alephbibnum - 000443512
oclc - 05069131
lccn - a 66007538

Table of Contents
    Front Cover
        Front page 1
        Front page 2
    Table of Contents
        Front page 3
    Front Matter
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    Main
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    Back Cover
        Page 261
Full Text





BULLET


N


OF THE

FLORIDA STATE MUSEUM


BIOLOGICAL SCIENCES


Volume 10


Number 6


SPHAERODACTYLUS (GEKKONIDAE) IN THE
GREATER PUERTO RICO REGION

Richard Thomas and Albert Schwartz


UNIVERSITY


OF FLORIDA


Gainesville
1966










Numbers of the BULLETIN OF THE FLORIDA STATE MUSEUM are pub-
lished at irregular intervals. Volumes contain about 300 pages and are not nec-
essarily completed in any one calendar year.
















WALTER AUFFENBERG, Managing Editor
OLIVER L. AUSTIN, JR., Editor


Consultants for this issue:

WALTER AUFFENBERG AND ERNEST E. WILLIAMS


Communications concerning purchase or exchange of the
manuscripts should be addressed to the Managing Editor of
State Museum, Seagle Building, Gainesville, Florida. 32601


publication and all
the Bulletin, Florida


Published April 6, 1966


Price for this issue $.90












SPHAERODACTYLUS (GEKKONIDAE) IN THE
GREATER PUERTO RICO REGION

RICHARD THOMAS AND ALBERT SCHWARTZ 1



SYNOPSIS: The geckos of the genus Sphaerodactylus in Greater Puerto Rico
(those West Indian islands between Mona Passage and Anegada Passage) are dis-
cussed in detail. S. macrolepis Giinther and S. grandisquamis Stejneger are com-
bined, and seven new subspecies of the resulting S. macrolepis are described
from Puerto Rico and Isla Vieques. S. nicholsi Grant and S. townsendi Grant are
regarded as races of a single species. S. danforthi Grant is considered a synonym
of S. macrolepis Giinther. A new subspecies of S. beattyi Grant is described from
St. Croix. Variation, detailed discussions of coloration and pattern, and geo-
graphic distribution of the remaining Puerto Rican forms are given and a hypo-
thetical histGry of the macro'epis complex is presented.



TABLE OF CONTENTS


Introduction --- .. .... .....
Acknowledgements and Abbreviations ----------.


Systematic List _--.---
Sphaerodactylus.
Sphaerodactylus
Sphaerodactylus
Sphaerodactylus
Sphaerodactylus
Sphaerodactylus
Sphaerodactylus
Sphaerodactylus
Discussion ---
Literature Cited --_-


macrolep s ..---_----
monensis ------------
roosevelti ---
klauberi ------ .
gaigeae -----------
nichosi ---- ---__- -
parthenopion --....
beattyi ..........____ ...


------.- 194
_------- 195


.-----_----.---_-...... ...... ......_ 196
.- .--........-........ ........... 196
.-.--- -----_-.....- ..... ....- .. ... 231
...------------.....-----------. 233
------.----.---.-. ........ .... 236
...-.. ---.................. .... ..- 2 4 0
--... --- --....-... ......_ ........- 242
.................. . ---...... ......... 2 4 7
.------ ---.- ....... -......- -..-.. 248
.------------------ -------.- - -- .. ... 2 5 3
....----.-------------- -.. -... 2 59


1 The authors have been engaged in studies on West Indian herpetology for
12 years; a portion of their field work in Cuba from 1957 to 1961 was supported
by two National Science Foundation grants. They have collected 40,000 reptiles
and amphibians from more than 100 islands. Earlier contributions to the Bulletin
by the junior author deal with the systematics of Leiocephalus cubensis and (as
junior author with William E. Duellman) the herpetology of southern Florida.
Manuscript received 28 June, 1965.-ED.


Thomas, Richard, and Albert Schwartz. 1966. Sphaerodactylus (Gekkonidae) in
the Greater Puerto Rico Region. Bull. Florida State Mus., vol. 10, no. 6, pp.
193-260.










INTRODUCTION
The term Greater Puerto Rico was used by Schmidt (1928) to de-
note the region containing Puerto Rico and the Virgin Islands. Al-
though he used it to refer to a probable formerly unified land mass,
we use it here in recognition of the faunal similarity of the islands
involved. It includes all the islands between the Mona and Anegada
Passages, though Desecheo, as one of the few West Indian islands
from which Sphaerodactylus has not been recorded, is not pertinent
to the present problem. S. macrolepis parvus King from the north-
western Leeward Islands is also discussed for the sake of com-
pleteness.
Not until Chapman Grant's herpetological investigations was the
true richness of the genus Sphaerodactylus in this region suspected.
Grant added seven forms, all but one of which this study regards as
valid. To date no trinomens have been applied to these geckos;
though monensis was originally described as a variety of macrolepis,
it has not been subsequently recognized as such. The following
names have been applied to Greater Puerto Rican Sphaerodactylus:

macrolepis Giinther, 1859
monensis Meerwarth, 1901
grandisquamis Stejneger, 1904
roosevelti Grant, 1931
nicholsi Grant, 1931
danforthi Grant, 1931
klauberi Grant, 1931
townsendi Grant, 1931
gaigeae Grant, 1932
beattyi Grant, 1937
parthenopion Thomas, 1965
The sphaerodactyls of Greater Puerto Rico, with one possible ex-
ception, share a community of characteristics and doubtless developed
from a single radiation within this region. The dorsal scales are flat-
tened, keeled, acute, and imbricate and possess (with one exception)
hair-bearing scale organs only. There is no middorsal zone of gran-
ules or granular scales. The presence of a scapular patch and ocelli,
though not universal, appears to be basic. While some species among
the forms considered are eminently distinct, few are separable by all-
or-nothing differences in scalation, which reflects the community of
similarities mentioned above. Differences are primarily of size, shape,
and color.








PUERTO RICAN SPHAERODACTYLUS


The sequence in which the forms are treated begins with the most
widespread, and therefore the least specialized with respect to habi-
tat restriction; this form is also moderate in size and apparently basic
in coloration. Beyond this the forms are arranged in a sequence of
relationship which we hope to clarify in the discussion.
We have followed King (1962) in characters used and in methods
of taking counts. In general we use his style of diagnosis and de-
scription, although we have defined species with an inclusive descrip-
tion. Stated average size differences of a few millimeters between
taxa may seem of questionable importance until the miniscule size
of these lizards is realized. In some cases we have indicated the
size range of the adults of a population by utilizing only the upper
50% of the total size range of the specimens. We have then deter-
mined the average size of all specimens falling within this range.
In this study we have relied primarily on our own recently col-
lected material, which we know to have accurate locality data, and
which was carefully preserved and color-noted. Extensive series of
some of the forms exist in collections; we have not drawn on these,
except when they pertained to localities from which we had little or
no material or when they were the basis for questionable identifica-
tions in the literature. The usefulness of a series of over 400 old and
ill-preserved specimens of a form from one locality is dubious in this
type of study.
ACKNOWLEDGMENTS AND ABBREVIATIONS
For the loan of additional specimens we are much indebted to
the following people: Ernest E. Williams, Museum of Comparative
Zoology at Harvard (MCZ); Charles F. Walker, George Zug and
Kraig Adler, University of Michigan Museum of Zoology (UMMZ);
and Charles M. Bogert and Grace M. Tilger, American Museum of
Natural History (AMNH). ASFS designates the Albert Schwartz Field
Series. RT specimens are from the senior author's private collection.
Paratypes have also been deposited in the United States National
Museum (USNM), University of Kansas Museum of Natural History
(KU), University of Illinois Museum of Natural History (UIMNH)
and the University of Florida collections (UF).
Felix Ifiigo of the Department of Agriculture and Commerce
of the Commonwealth of Puerto Rico has repeatedly given prompt
and courteous service in obtaining official sanction for our collecting
activities; for this he merits our utmost appreciation. Likewise, our
sincere appreciation is due to Jesse E. Williams, who paved the way
for our collecting on Ramey Air Force Base, Puerto Rico and to


1966


195







BULLETIN FLORIDA STATE MUSEUM


George A. Seaman of St. Croix, who not only was of great help during
a visit to that island, but who has subsequently furnished us with in-
formation about localities there. We also wish to thank Rowan Roy
of Tortola for his aid during our visit to that and adjacent islands.
Juan A. Wirshing has been our host on three trips to the island of
Caja de Muertos; without his help visits to that island would not have
been so easy or so profitable.
We thank the following people for assistance in the field: Robert
I. Balfour, Donald W. Buden, Gerald D. Gagnon, Warren R. Faust,
Ronald F. Klinikowski, David C. Leber, and Barton L. Smith. We
would also like to thank Grady E. Lanier for use of his microscope for
examination of scale organs and Wayne King for discussion of various
aspects of this problem with us, particularly the taxonomic significance
of scale organs. The illustrations are by the senior author, the maps
by the junior author, and the escutcheons by R. F. Klinikowski and
the senior author.
SYSTEMATIC LIST
Sphaerodactylus macrolepis Giinther
Sphaerodactylus macrolepis Giinther, 1859, Ann. Mag. Nat. Hist. (3), Vol.
4, p. 215.
TYPE LOCALTY: St. Croix, American Virgin Islands.
DISTmBUTI6N: Puerto Rico, including Cayo Santiago and Isla
Pifieros; Isla Vieques, including the islets of Cayo de Afuera and Cayo
de Tierra; Culebra, including Culebrita and Cayo Luis Pefia; the
American and British Virgin Islands; the Leeward Islands from Dog
Island and Anguilla south to St. Barthelemy. On Puerto Rico occurs
almost island-wide from sea level to a known elevation of 2800 feet
north of Sabana Grande, but absent from the southwestern region
from the peninsula of Cabo Rojo eastward to near Ponce, and appar-
ently absent from the Sierra de Luquillo. On smaller islands usually
widely distributed altitudinally.
DEFINITION: A species of Sphaerodactylus with very large, acute,
strongly keeled, flattened imbricate dorsal scales, axilla to groin 16-
28; no area of middorsal granules or granular scales; dorsal body scales
with one of two structures: 1, with both knob-like and hair-bearing or-
gans, the latter, each with one hair, are located on the free posterior
edge of each scale (Puerto Rico and some of its small satellite islands;
Isla Vieques). 2, with only hair-bearing organs on the free posterior
edge of each scale (Culebra and its associated islets, St. Croix, and
the Virgin Islands from St. Thomas and its satellites north and east
to Anegada; the northernmost Leeward Islands, except Sombrero,


Vol. 10








PUERTO RICAN SPHAERODACTYLUS


south as far as St. Barthelemy). Dorsal scales of tail keeled, acute,
imbricate, and flat-lying; ventral scales of tail smooth, rounded, en-
larged midventrally; gular scales almost always keeled but occasion-
ally smooth; chest scales smooth to keeled laterally or even somewhat
keeled centrally; ventrals rounded, imbricate, scales axilla to groin
20-32, usually smooth, but occasionally keeled on the anterior part
of the venter and rarely on the entire venter; scales around midbody
31 to 54; internasals 0 to 3 (mode 1); upper labials to mid-eye, 3 (some
times 4); escutcheon (fig. 11a) with a broad and compact central area
and extensions onto the thighs to near the underside of the knee
(3-10 X 11-30).
Color pattern sexually dichromatic, except in the Leeward Island
population, and variable among the subspecies. Males generally
with a tan to brown dorsum, usually without lines, with scattered
darker scales (salt-and-pepper), or patternless; a unicolor to faintly
marked head, usually without a cephalic figure; a black or dark brown
scapular patch which may be reduced or absent, with a pair of
white to buffy ocelli which may be almost or entirely absent; throat
pattern varying from immaculate to prominently marked; and a pale
venter. Females with a unicolor tan to brown dorsum, with or with-
out a series of longitudinally parallel lines, the paramedian pair partly
fused to form a series of median blotches; an almost unicolor to boldly
marked head; a black to brown scapular patch, at times reduced, with
two white, buff, or gray ocelli, either enclosed within the patch or on
or near its periphery; throat pale to heavily marked with darker; and
a pale venter. Iris color variable, from greenish-gray or bluish to
yellow, golden, or brown. Habitus moderate; snout fairly long and
narrow. Adult size variable by subspecies from 23 mm to 35 mm
snout-vent length. S. macrolepis ranks third in size of individuals
among Greater Puerto Rican sphaerodactyls, the largest within the
species being the populations from the Cordillera Central, the small-
est the extralimital populations from the northern Leeward Islands.
REMARKS: When Stejneger (1904: 603) described Sphaerodactylus
grandisquamis he differentiated it from S. macrolepis Giinther solely
on the basis of the size of the dorsal scales. Barbour (1921: 254-55)
combined grandisquamis and S. monensis Meerwarth from Isla Mona
with macrolepis, and gave the range of the latter as five of the Virgin
Islands, Puerto Rico, and Vieques. Schmidt (1928: 72-73) followed
Barbour's action. Grant (1931) removed monensis from obscurity and
regarded it as a distinct species; later (1932d) he distinguished be-
tween S. macrolepis and S. grandisquamis and called each a separate
species.


1966








BULLETIN FLORIDA STATE MUSEUM


For reasons outlined below, the form monensis should be regarded
as a distinct species. King (1962: 18) has suggested that at least macro-
lepis and grandisquamis (along with notatus of Cuba, Isla de Pinos,
the Bahamas, southern Florida, and Little Swan Island, and difficilis
of Hispaniola) might best be considered conspecific. Examination of
their scale organs shows that macrolepis possesses only hair-bearing
structures, whereas grandisquamis has both knob-like and hair-bear-
ing types. If we consider that the type of scale organs is species con-
stant, a course already followed by King (1962: 45) and with which
we have concurred, we must regard grandisquamis as a species dis-
tinct from macrolepis. Using scale organs as the sole structural means
of differentiating between the species, the range of grandisquamis
is Puerto Rico and its satellites and Isla Vieques, whereas that of
macrolepis includes all islands and islets from Culebra east to Anegada
and the Leeward Islands south to St. Barthelemy. This diagnosis is
zoogeographically sound.
However, specimens from Vieques have the scale organs of
grandisquamis but the pattern and coloration of macrolepis. The
Vieques sphaerodactyls are thus intermediate, but not intergradient
in the orthodox usage of that word, between grandisquamis and mac-
rolepis, in that they combine undiluted the characteristics of both
forms.
As research into the characteristics of Antillean sphaerodactyls
has progressed, many characteristics that previously were supposed
to be on the specific level have been discarded one by one. Un-
fortunately the scale organs must now join the characters that must
be interpreted in the light of other evidence; we feel that although
they may still be employed as guideposts to relationships, they should
be used with discretion. To regard the Puerto Rico-Vieques geckos
as one species and the Culebra-Virgin Islands-Leeward Islands geckos
as another would obscure the very obviously close relationships of
these two forms. For those who feel that scale organs are species
constant, grandisquamis and macrolepis must be separated at the
specific level. Our choice, although without precedent, seems to us
to reflect the biological situation in a more rational nomenclatorial
manner.
It is surprising that previous workers, especially Grant, did not
detect the striking geographic variation in S. macrolepis on Puerto
Rico. This variation should not surprise anyone familiar with the
ecological diversity of Puerto Rico, which ranges from the hot and
arid southern littoral to the extremely wet, forested Cordillera Cen-
tral, and the limestone Pepino Hills in the northwest. On the basis


Vol. 10































M10 15
MI


67i b


FIGURE 1. Puerto Rico, showing the ranges of S. macrolepis (solid symbols) and S. roosevelti (semi-solid symbols). Races of
S. macrolepis as follows: grandisquamis, horizontal lines; stibarus, fine vertical lines; phoberus, medium stippling; mimetes, heavy
stippling; ateles, diagonal lines from lower left to upper right; spanius, open vertical lines; guarionex, diagonal lines from upper
left to lower right. Overlap of symbols represents intergradation between subspecies. S. roosevelti, fine stippling.







BULLETIN FLORIDA STATE MUSEUM


of the examination of 639 specimens of S. macrolepis from Puerto
Rico six subspecies are recognizable; another race is confined to
Isla Pifieros.

Sphaerodactylus macrolepis grandisquamis Stejneger
Sphaerodactylus grandisquamis Stegneger, 1904, Rept. U. S. Nat. Mus., 1902,
p. 602. Type specimen USNM 27007.
TYPE LOCALITY: Luquillo, Puerto Rico.
DISTRIBUTION: Extreme eastern Puerto Rico, from the vicinity of
Rio Piedras (except the coast from San Juan to Loiza Aldea) south
to near Punta Santiago; Cayo Santiago off Playa de Humacao (fig. 1).
DIAGNOSIS: A subspecies of S. macrolepis characterized by a com-
bination of moderate size (males to 33 mm, females to 31 mm snout-
vent length); high number of midbody scales (36 to 46, mean 41.2 -
.43 = one standard error of mean); escutcheon 5-9 X 23-30. Dorsum
of males unicolor brownish or salt-and-pepper, head brown with or
without a vague pattern, the most obvious features of which are
a pair of pale lines from the orbits extending posteriorly to the anterior
border of the scapular patch (the head may also be vaguely flecked
with darker), a fairly conspicuous dark occipital spot, and occasion-
ally a dark nuchal spot at the juncture of the two pale postorbital
lines. Black scapular patch variable from large and conspicuous with
an included pair of two transversely elongated buffy to white ocelli,
the entire patch surrounded by a broad buff to white border, to an
almost complete absence of the scapular patch, the remnants consist-
ing of the transverse ocelli each with a dark border. Dorsum of fe-
males tan to brown, usually with a series of fairly prominent dark
brown longitudinal lines, sometimes broken and fragmented and form-
ing almost a series of transverse chevronate figures; head pattern
much as in male except that the nuchal spot is more consistently
present; scapular patch always present, large, bordered by a broad
buff to white outline (fig. 2a). Iris blue to grayish-green, or yellow-
gray.
REMARKS: The color of a series of 12 specimens from the vicinity
of the type locality was noted in life as follows: Males: dorsal
ground color yellow-tan, scattered scales darker brown; tail and
head yellow; head pattern obscure and brownish to rusty-orange;
ventral ground color yellow, underside of tail orange; chin and throat
yellow to orange with some pale brown to rusty flecks; iris grayish-
green. Females: dorsal ground color tan with brown longitudinal
lines; head with a pale U set off by brown edging; ventral ground
color yellowish, throat pale yellow-orange with brownish flecks.


Vol. 10


200








PUERTO RICAN SPHAERODACTYLUS


Juveniles from this locality resembled the females in pattern and col-
oration except when very small; tiny specimens (snout-vent lengths
14 to 20 mm) are very dark uniform brown dorsally and ventrally,
with a well developed and palely outlined black scapular patch and
ocelli, and a white tail tip preceded by a black band. Coloration and
pattern in live geckos from Las Croabas and Ceiba resembled that
described above for topotypes; the iris of males from Ceiba was re-
corded as yellow-gray.
A moderately sized subspecies, S. m. grandisquamis males reach
a maximum known snout-vent length of 33 mm and females 31 mm.
The discrepancy is doubtless due to the smaller series of females.
Scale counts are: dorsal scales axilla to groin 17-23, mean 20.7; ventral
scales axilla to groin 24-32, mean 27.7; midbody scales 36-46, mean
41.2; escutcheon 5-9 X 23-30; internasals 0-3, mode 1; fourth toe
lamellae 8-12, mode 12. All specimens have 3 supralabials to mid-
eye. The usual condition is that the throat scales are keeled and
both chest and ventral scales smooth; 2 specimens have only some
of the throat scales keeled, 7 have some keeled chest scales, and
two of these have all the chest scales keeled.
A single egg from near Luquillo measured 7.0 mm X 5.2 mm.
Two hatchling S. m. grandisquamis had snout-vent lengths of 14 mm.
S. m. grandisquamis has been taken in littoral situations under
palm trash and other debris, at times rather far back from the coast
but still near it. The only major exception is the series from south of
Luquillo which was collected in an abandoned shack in rolling, rather
mesic, country. The series from near Ceiba was collected in a small
patch of lowland forest with a leaf-littered floor and much trash.
The single specimen from Cayo Santiago (MCZ 58826, a female with
a snout-vent length of 24 mm) agrees with S. m. grandisquamis both
in pattern and scale counts.

SPECIMENS EXAMINED (all from Puerto Rico): UMMZ 73587, 73589, Rio
Piedras; UMMZ 73615 (8 specimens), Trujillo Alto; UMMZ 124804 (4 specimens),
Campo Rico; UMMZ 73588, 73622, 10 km S Can6vanas; ASFS V4997-98, Lu-
quillo (outskirts); ASFS V5001-10, V6002, V6211, 1.5 km SW Luquillo; MCZ
34519-23, Cabezas de San Juan; ASFS X7470-78, 0.5 mi. N Las Croabas; ASFS
V5090-103, 1.8 km SSW Ceiba; MCZ 58824, Punta Santiago; ASFS X4059-61, 4.6
mi. E Humacao; MCZ 58826, Cayo Santiago.

Sphaerodactylus macrolepis stibarus, new subspecies
TYPE: MCZ 81022, an adult female from Isla Pifieros, Puerto
Rico, one of a series taken 13 February 1965 by Albert Schwartz and
Richard Thomas. Original number V5030.


1966








BULLETIN FLORIDA STATE MUSEUM


PARATYPES: AMNH 94177-78, ASFS V5020-22, V5031-34, V5044-47, KU
79887-90, UF 21271-76, UIMNH 56915-18, same data as type.
DISTRIBUTION: Known only from Isla Pifieros, off the eastern end
of Puerto Rico (fig. 1).
DIAGNOSIS: A subspecies of S. macrolepis characterized by a com-
bination of small size (maximum snout-vent length of both sexes 80
mm), moderate number of midbody scales (36 to 41, mean 38.6 .37);
escutcheon 5-8 X 18-24. Males with a black scapular patch and two
more or less rounded ocelli; females with or without longitudinal lines
and a fragmented dark cephalic head pattern; iris brown with golden
pupillary ring.
DESCRIPTION OF TYPE: An adult female with a snout-vent length
of 28 mm, tail 24 mm. Dorsal scales axilla to groin 18; ventral scales
axilla to groin, -; midbody scales 38; fourth toe lamellae 11; in-
ternasal 1; 3 'supralabials to mid-eye; gular scales keeled, chest and
ventral scales smooth. Dorsal body ground color uniform yellowish-
tan; ground color of head unicolor, with a fairly prominent dark brown
cephalic pattern consisting of a fragmented pair of lines on the snout,
which join an irregular and fragmented pair of postorbital lines; the
right postorbital line joins the dark nuchal spot, the left does not; be-
tween the two postorbital lines lies a very dark brown occipital spot;
a dark loreal stripe passes through the eye and continues posteriorly
onto the side of the neck. The scapular spot is irregular, not prom-
inently margined with a pale zone, and encloses two buffy ocelli. The
back has a series of three or four irregular longitudinal lines. The tail
has two pale chevrons dorsally, outlined with dark brown or black.
The throat is yellow and has some very fine inconspicuous dark stip-
pling. The ventral surface is tan.
Variation: 9 males and 9 females show a maximum size of 30
mm in each sex and the following scale counts: dorsal scales axilla
to groin 17-21, mean 19.6; ventral scales axilla to groin 21-28, mean
24.5; midbody scales 36-41, mean 38.6; escutcheon 5-8 X 18-24; inter-
nasals 0-3, mode 1; fourth toe lamellae, 9-11, mode 11. All speci-
mens have 3 supralabials to the center of the eye. Usually the gular
scales are keeled and the chest and ventral scales smooth; in two speci-
mens the keeled scales extend onto the chest, and in four the gular
scales are not completely keeled.
In live males the dorsum was tan to yellowish-tan to dull brown,
head distinctly yellow and the throat vivid yellow-orange; head pattern
an indistinct pair of rusty postorbital lines; diffuse occipital spot;
dorsum completely devoid of pattern or with scattered, unaligned
dark scales; scapular patch black, usually without a white border;


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PUERTO RICAN SPHAERODACTYLUS


and the two white ocelli more or less round and included within
the patch. The greatest reduction of the patch (ASFS V5024) is
demonstrated as a single orthodox ocellus and a smaller mate in-
cluded in a small restricted black area. The ventral surface is tan-
nish, and the bright chin and throat have vague rusty markings
in life.
The females have the same dorsal ground color as the males, but
none was so pale as the palest males. The head patterns agree with
that of the type, though the type shows the head pattern more clearly
than do most other specimens; the usual condition is a deposition of
dark pigment in the light head areas that obscures the pattern which
is not particularly distinct. The ocelli are buff in adults, white in
young, and included in the large black scapular patch; patch with a
trace of a white border, but this is seldom broad and prominent;
throat yellow in life, with vague stippled darker diagonal markings;
venter tannish.
Juveniles are similar to adult females, but the head pattern may be
even less conspicuous. The smallest juvenile (12 mm snout-vent
length) is smaller than hatchlings of S. m. grandisquamis and is brown
above and paler below. All juveniles that show any dorsal pattern
(snout-vent lengths 17 to 23 mm) have a salt-and-pepper rather than
a lined dorsum. The scapular spot in the juveniles is large and bold,
and not outlined with a pale zone, in distinct contrast to that in
young grandisquamis.
COMPARISONS: The Isla Pifieros race is differentiated from S. m.
grandisquamis primarily by three characters: 1) indistinct head pat-
tern in females; 2) smaller escutcheon in males; 3) lower number of
midbody scales (table 1). Although the range of midbody scales of
grandisquamis includes that of stibarus, the means (41.2 in the former,
38.6 in the latter) are statistically different. The breadth of the es-
cutcheon in grandisquamis varies from 23 to 80 scales, that of stibarus
from 18 to 24. Other features that help differentiate the two races
are the less consistent pale and prominent border to the scapular spot
in female stibarus, the round rather than transversely elongated ocelli
in stibarus, the modal number of 11 fourth toe lamellae (stibarus never
has counts of 12, the modal condition in grandisquamis), and tan
rather than bright yellow ventral color in stibarus.
Remarks: S. m. stibarus is obviously a derivative of and very
closely related to S. m. grandisquamis. The differences between the
two are relatively minor but nonetheless constant and significant. It
seems appropriate to recognize nomenclatorially the fact that on this
islet differentiation has definitely taken place. The species shares


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the island with S. nicholsi and S. gaigeae; that so small an island
should harbor three species of the same genus is remarkable. The
series of S. m. stibarus was collected in sea-grape (Coccoloba) leaves
along the shore and in mango leaves and the coconut trash in an
abandoned grove.

TABLE 1. COUNTS OF MIDBODY SCALES IN TEN SUBSPECIES OF S. macrolepis

Mean Standard Standard error
Subspecies N and extremes deviation of mean (x2)

grandisquamis 28 41.2 (36-46) 2.29 .86
stibarus 16 38.6 (36-41) 1.49 .74
phoberus 39 37.5 (33-41) 2.62 .84
mimetes 29 36.1 (31-40) 2.10 .78
ateles 26 39.4 (36-42) 1.78 .70
spanius 26 43.6 (41-47) 1.68 .66
guarionex 92 41.3 (36-49) 2.60 .54
inigoi 33 36.8 (33-41) 1.84 .64
macrolepis 86 43.1 (38-50) 2.46 .54
parvus 14 48.4 (44-54) 2.87 1.54


Sphaerodactylus macrolepis phoberus, new subspecies

TYPE: MCZ 81023, an adult female from Isla Verde (San Juan
International Airport), Puerto Rico, one of a series taken 15 June
1962 by Ronald F. Klinikowski and David C. Leber. Original num-
ber X919.
PARATYPES (all from Puerto Rico): AMNH 94179-84, KU 79891-94, MCZ
81024-35, UF 21242-51, UIMNH 56919-24, USNM 152593-96, same locality as
type, 13 June 1962, R. F. Klinikowski, D. C. Leber, A. Schwartz; ASFS X899-918,
X920-36, same data as type; ASFS X4134-58, same locality as type, 20 August
1962, R. I. Balfour, W. R. Faust, R. F. Klinikowski, and D. C. Leber.
ASSOCIATED SPECIMENS: ASFS X897, X944, X1355-56, X1710, X1716, X1792,
X1945, X2029-30, X2034, X2240, X2246-47, X2264, X2329-31, X2365-66,
X2898, X3135-37, X3267, X3313, X3551-52, X3860, X4019-20, X4044-46, all
hatchlings from eggs taken at the type locality; ASFS X4637-43, 2.4 mi. W Loiza
Aldea; UMMZ 124811, Punta Vacia Talega (see below).
DISTIBUTION: Known only from the type locality; intergrades
with S. g. grandisquamis to the east near Loiza Aldea (fig. 1).
DIAGNOSIS: A subspecies of S. macrolepis characterized by a
combination of small size (maximum snout-vent length of both sexes
31 mm); moderate number of midbody scales (33-41, mean 37.5 .42);
males usually dark wood brown, unicolor or with scattered dark brown


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to blackish scales; scapular patch diffuse, not outlined by a pale zone.
Females patterned like males; scapular patch usually fairly large, out-
lined by a pale zone that is fragmented and irregular in outline; head
pattern reduced or almost obliterated; iris black with bronzy pu-
pillary ring.
DESCRIPTION OF TYPE: An adult female with a snout-vent length
of 80 .mm, tail 27 mm; dorsal scales axilla to groin 20, ventral scales
axilla to groin 29; midbody scales 39; fourth toe lamellae 12; inter-
nasal 1; 3 supralabials to mid-eye; gular scales keeled, chest and ven-
tral scales smooth. Dorsal ground color dark brown with no indica-
tion of pattern; head brownish with vague indications of a pair of
pale postorbital lines and dark occipital and nuchal spots; no mark-
ings on sides of dark brown head and neck; scapular spot black and
irregularly diffuse, not outlined with a pale zone; no pale ocelli
(fig. 2b); unregenerated tail brown and practically patternless; throat
pale yellow without discrete darker markings; ventral ground color
yellowish-tan; underside of tail pale yellow.













A BC C A i _

FIGURE 2. A, S. macrolepis grandisquamis, anterodorsal view, female, ASFS
X7470, 0.5 mi. N Las Croabas, Puerto Rico. B, S. macrolepis phoberus, antero-
dorsal view, female, MCZ 81023, type, Isla Verde, Puerto Rico. C, S. macro-
lepis mimetes, anterodorsal view, male, MCZ 81036, type, 12.3 km SE Patillas,
Puerto Rico.


Variation: 20 males and 20 females have maximum snout-vent
lengths of 31 mm in each sex and the following scale counts: dorsal
scales axilla to groin 17-22, mean 19.6; ventral scales axilla to groin
23-31, mean 26.4; midbody scales 33-41, mean 37.5; escutcheon 6-10 X
24-28; internasals (data from 144 individuals) 0-3, mode 1; fourth toe


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lamellae 9-12, mode 10. All 40 specimens examined in detail have
3 supralabials to mid-eye. The usual condition of keeled gular scales
and smooth chest and ventral scales obtains in most specimens; one
has the chest scales completely keeled and two have some of the
scales on the chest keeled.
The dorsal ground color of live males varies between tannish-gray
and dark brown, with the darker hues more common; dorsum usually
unicolor, but some with scattered darker brown scales; head pattern
obscure, consisting at best of a pair of pale postorbital lines with a
diffuse darker occipital spot; nuchal spot absent except in subadult
males where it is faint. Some individuals have rather large brownish
spotting on the occiput and neck, and the edges of the postorbital lines
may be more heavily outlined in dark brown. The entire dorsal sur-
face of the head suffused with yellow; dorsum of tail commonly spot-
ted with dark brown, or with light crossbands; scapular spot very dif-
fuse, without a pale outline, nearly absent in most specimens, some-
times reduced to only the small, more or less circular creamy ocelli;
throat deep yellow; ventral ground color yellowish-tan with some fine
dark stippling.
The females resemble the type closely, and the ground color of
females shows the same variation as that of males, with the darker hues
predominating. The scapular patch varies in extent, showing a strong
tendency toward reduction of both size and intensity, and is absent
in some specimens. Even the ocelli have disappeared in some fe-
males. When fully expressed the patch is surrounded by a buff out-
line. The ocelli vary in life from white to cream. The head in life
is not yellow as in the males, and the head pattern is like that of the
type, never bold and easily discernible, and often obscured by addi-
tional brown pigment. The occipital spot is usually present, the
nuchal spot usually absent. The dorsum is either uniform and pat-
ternless or with a vague salt-and-pepper pattern, and only very
rarely shows any alignment of the dark scales to form indistinct longi-
tudinal lines. The throat is pale yellow, with occasional diffuse and
inconspicuous pale brown spotting. The ventral coloring is as that
described for males.
Hatchlings are drab brown dorsally, somewhat paler brown ven-
trally. The back is unpatterned except for the scapular patch, which
lacks a pale border upon hatching, and ocelli. The head in juveniles
lacks any pattern and is a simple brownish-tan. The tip of the tail
is white in hatchlings, preceded by a dark brown region. This color-
ation may persist into adulthood if the original tail is not lost.


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COMPARISON: S. m. phoberus differs from both grandisquamis
and stibarus in being much darker dorsally, lacking a lined pattern
in females, having round, rather than transversely elongate ocelli as
in grandisquamis, and being slightly smaller. The bold scapular patch
with its wide and conspicuous pale border and the more prominent
female head pattern in grandisquamis differentiate this race from
phoberus.
Although the midbody scales of grandisquamis and phoberus over-
lap in counts, those of phoberus are significantly lower (table 1);
these counts show no significant difference between stibarus and
phoberus. The modal fourth toe lamellar count of 10 in phoberus
differs from those of 11 in stibarus and 12 in grandisquamis.
REMARKS: A small series (ASFS X4637-43) from 2.4 mi. W Loiza
Aldea and a single individual (UMMZ 124811) from Punta Vacia
Talega between Isla Verde and the range of S. m. grandisquamis are
intermediate in color and pattern between phoberus and grandisqua-
mis, but closer to phoberus.
The type locality of S. m. phoberus is in rich, mesic woods imme-
diately adjacent to the San Juan International Airport, and lies within
an old coconut plantation. The type series of phoberus was collected
in both the plantation and the woods, where the lizards were found
under almost any sort of debris including moist piles of rotting coconut
husks, tires, rocks, concrete slabs, and palm logs. A series of ten eggs
measures 6.7-7.2 X 5.0-5.8 mm. Hatchlings measure 13 or 14 mm
in snout-vent length.
The range of S. m. phoberus is unknown. We assume that it en-
compasses the peninsula now occupied by San Juan and its suburbs,
and probably extends eastward to near the Rio Grande de Loiza.
The intergradient specimens mentioned above are from west of the
mouth of this river, so that some genetic influence of grandisquamis
is manifest at least this far to the west.
We have tried in vain to collect S. macrolepis between Mameyes
and Luquillo. This region has extensive but moderately dry decid-
uous woods near the coast that appear quite suitable for S. macro-
lepis, but intensive search has revealed no sphaerodactyls in this im-
mediate area. The same situation applies elsewhere in Puerto Rico,
bringing to mind Grant's statement (1931: 209) that this species is
"distributed in foci throughout Porto Rico except in localities of ex-
treme drought or moisture." Although we hesitate to state emphat-
ically that S. macrolepis is absent from some regions, our experience
in certain areas indicates that it is at least extremely rare.


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Sphaerodactylus macrolepis mimetes, new subspecies
TYPE: MCZ 81036, an adult male from 12.3 km SE Patillas, Pu-
erto Rico, taken 29 January 1965 by Richard Thomas. Original num-
ber V4777.
PARATYPES (all from Puerto Rico): ASFS V4778-83, V4790-95, UIMNH
56925-30, same data as type; AMNH 94186, 10 km SE Patillas, 1 February 1965,
R. Thomas; AMNH 94185, 5 mi. W Las Mareas (south side Bahia de Jobos), 30
January 1965, R. Thomas; KU 79898-99, UF 21254-55, 1.5 km S Velazquez, 9
March 1965, A. Schwartz, R. Thomas; MCZ 81087-42, AMNH 94187-89, 12.2
km WNW, 1 km S Santa Isabel, 4 March 1965, A. Schwartz, R. Thomas; ASFS
V5756-57, 2 km E Juana Diaz, 200 feet, 9 March 1965, R. Thomas; UF 21252-58,
KU 79895-96, Bafios de Coamo, 9 March 1965, A. Schwartz, R. Thomas; MCZ
34446-49, Bafios de Coamo, 1931-82, C. Grant; MCZ 84556-57, between Coamo
and Bafios de Coamo, 16 November 1930, C. Grant; KU 79897, AMNH 94190,
1 mi. SE Rio Jueyes, 9 March 1965, A. Schwartz, R. Thomas.
ASSOCIATED SPECIMENS: UMMZ 73592, 12814, Bafios de Coamo; UMMZ
73586 (3 specimens), 73620, Arroyo; MCZ 34444-45, Punta Arroyo; UMMZ 73591
(4 specimens), beach near Maunabo. S. m. mimetes X grandisquamis: MCZ
34505-10, mountains between Maunabo and Yabucoa; ASFS X4057-58, 2.5 mi.
SW Yabucoa, 800 feet.
DISTmBUTION: Southern Puerto Rico from Maunabo west to the
vicinity of Juana Diaz and Coamo (fig. 1).
DIAGNOSIS: A subspecies of S. macrolepis characterized by a com-
bination of small size (males to a maximum snout-vent length of 32
mm, females to 30 mm), low number of midbody scales (31-40, mean
36.1 .39); males yellow-brown to tan with a few scattered dark
brown scales, scapular patch small to absent, and head and throat
heavily spotted with black, females with a lineate dorsal pattern,
scapular pattern present but often much reduced and not outlined
by pale margin, head pattern bold and vivid, and throat usually heavily
spotted with black in adults. Iris dark brown.
DESCRIPTION OF TYPE: An adult male with a snout-vent length of
30 mm, tail 22 mm, regenerated. Dorsal scales axilla to groin 17;
ventral scales axilla to groin 21; midbody scales 33; fourth toe lamellae
9; internasal absent; 3 supralabials to mid-eye; escutcheon 6 X 22;
gular scales keeled, chest and ventral scales smooth. Dorsal ground
color yellow-brown with a few scattered brown scales; head ground
color tan with large black discrete spots, two of which are the usual
occipital and nuchal spots; scapular patch black, restricted, constricted
medially, enclosing two small white round ocelli, not surrounded by
a pale margin (fig. 2c); throat light gray with large discrete black
spots; venter yellowish.
VARIATION: 33 specimens of both sexes have maximum snout-
vent lengths of 32 mm in males, 30 mm in females, and the following


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scale counts: dorsal scales axilla to groin 16-21, mean 18.1; ventral
scales axilla to groin 20-29, mean 24.1; midbody scales 31-40, mean
36.1; escutcheon 4-5 X 21-24; internasals 0-2, mode 1; fourth toe
lamellae 8-12, mode 10. All specimens have 3 supralabials to mid-
eye; gular scales keeled, the chest and ventral scales smooth, though
two individuals have some chest scales keeled, and another has all
these scales keeled. Three lizards lack any keeled scales in the
gular area.
The six paratypic males are much like the type in pattern and
color. The heads are always boldly marked, and the basic female pat-
tern described below is obscured in males by additional dark pigment,
to give a darkly spotted head on a gray, tan, or straw ground color.
The scapular patch is present and small in all but one specimen
which lacks it and its ocelli completely. All males have the throat
darkly spotted, though the intensity and degree of spotting vary some-
what. The ventral color is yellowish to tan; the tails are orange; the
iris is dark brown to yellow-brown.
The 28 females were yellow-brown to tan in life, with a prom-
inent dorsal pattern of dark longitudinal lines. The head ground
color varies between gray, tan, and brownish. The bold dark brown
or black cephalic pattern is composed of a pair of lines from the
snout to the orbits, a pair of postorbital lines from the orbits onto
the neck where they join with one another and usually with the dark
nuchal spot to give a dark U-shaped postorbital figure, a bold dark
occipital spot, and a canthal line from the snout through the eye and
onto the side of the neck. Occasional specimens have additional dark
pigment in the light areas between the lines, but this does not ob-
scure the main pattern. The black scapular spot is always small and
restricted, never outlined with a pale zone, and may be completely
absent; the white ocelli are either included in the spot when it is
larger, or are peripheral if it is restricted. The throats are light gray,
with dark spots prominent in 19 specimens and faint to absent in 9,
most of which are smaller females. However, 3 adult individuals
lack spotting, and some spot-throated females are as small as some
that lack spots. In general the throat spotting in females is less ex-
tensive than in males, though some females have throats as darkly
and regularly spotted as those of some males. The ventral ground
color is yellowish to tan, and the undersides of the tails are deep
yellow or straw.
Even tiny juveniles with snout-vent lengths of 14 mm show the
female lineate pattern, as well as a scapular patch and ocelli and the
female head pattern. The throats of juveniles are unspotted.


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COMPARISONS: S. m. mimetes, as the trivial name implies, mimics
S. m. macrolepis of the Virgin Islands in basic head pattern and color
(see the discussion of the nominate race for comparisons). It differs
from all other subspecies on Puerto Rico in its heavily spotted throat.
The lower number of midbody scales is statistically different from
those of both grandisquamis and stibarus, but not of phoberus (table
1). The lineate female pattern, the heavily spotted throats, and
paler coloration of mimetes easily distinguish it from phoberus. Ab-
sence of a large scapular patch surrounded by a pale border distin-
guishes mimetes from grandisquamis, and the distinct head patterns
of mimetes distinguish it from the three more northern races.
REMARKS: Eight specimens (ASFS X4057-58, MCZ 34505-10) are
from the Serrania de Panduras; this rather low (1700 feet) mountain
range lies immediately north of the range of mimetes and extends
to the coast. North of these mountains, S. m. grandisquamis occurs,
at least in the vicinity of Humacao. The three males from the Pan-
duras have plain throats and very vaguely patterned heads; of the
four females, one (MCZ 34506) has a heavily dotted throat which
the others lack. All females have the head pattern well developed
and are thus like mimetes despite their plain throats. Because of
their geographically intermediate position, the occurrence of throat
spotting in one of the females, the vaguely patterned heads of the
males, and the prominently patterned heads of the females, we regard
these lizards as intergradient between mimetes and grandisquamis.
Specimens of S. m.. mimetes were collected in a lowland cafetal
near Juana Diaz, in coastal palm groves near Santa Isabel, Velazquez,
and Las Mareas, and under logs and rocks in dry hills and plains at
Bafios de Coamo and Rio Jueyes. Only the cafetal locality can be
considered more or less mesic; all others were extremely hot and dry,
at least at the time of our visit.

Sphaerodactylus macrolepis ateles, new subspecies

TYPE: MCZ 81043, an adult female from Balneario de Boquer6n,
Puerto Rico, one of a series taken 26 February 1965 by Albert Schwartz
and Richard Thomas. Original number V5522.
PARATYPES (all from Puerto Rico): KU 79900-03, 9.6 km SSW Mayagiiez,
26 February 1965, A. Schwartz, R. Thomas; AMNH 94191-94, ASFS V5515-17,
V5523-28), same data as type; UF 21256, 9 mi. NW Yauco, 1000 feet, 27 Febru-
ary 1965, R. Thomas; UIMNH 56931-35, 8.5 mi. NW Yauco, 700 feet, 27 Feb-
ruary 1965, A. Schwartz, R. Thomas; UF 21257, 8 mi. NW Yauco, 600 feet, 27
February 1965, R. Thomas; UF 21258, 2.1 km SW San German, 28 February
1965, A. Schwartz.


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ASSOCIATED SPECIMENS: MCZ 34460-62, 5 mi. S Playa de Mayagiiez;
UMMZ 73627, Playa de Ponce; ASFS V6629-30, V6641, 4.6 km W, thence 4.6
km NW Juana Diaz.
DISTRIBUTION: Southwestern Puerto Rico from the vicinity of
Mayagiiez south to Balneario de Boquer6n and thence eastward,
north of the Valle de Lajas, to the vicinity of Ponce (fig. 1).1
DIAGNOSIS: A subspecies of S. macrolepis characterized by a com-
bination of moderate size, maximum snout-vent length of both sexes
33 mm, moderate number of midbody scales, 36-42, mean 39.4 .35.
Males dull grayish-tan, usually with many scattered dark brown scales;
head patternless and yellow-tan but with a few dark brown spots;
scapular patch very reduced or absent; ocelli very faintly discern-
ible and gray, not white; and throat unpatterned or with a few brown
dots forming a vague collar. Females tan with dark brown longitudi-
nal lines; head prominently patterned; scapular patch rather small,
without pale margins; ocelli peripheral, at times enclosing the patch
remnant between them; and throat gray, not yellow, without definite
markings. Iris grayish with a greenish tinge.
DESCRIPTION OF TYPE: An adult female with a snout-vent length
of 32 mm, tail 27 mm, regenerated. Dorsal scales axilla to groin 20;
ventral scales axilla to groin 28; midbody scales 41; fourth toe lamellae
12; internasal 1; 3 supralabials to mid-eye; gular scales keeled, chest
and ventral scales smooth. Dorsal ground color tan with four distinct
darker brown longitudinal lines; head unicolor with back and with
a dark brown pattern consisting of a faint pair of dark lines from
snout to eyes, a pair of postorbital lines which join the nuchal spot
and surround a dark occipital spot, a dark line from the lores through
the eye and onto the neck, meeting the dorsolateral longitudinal line
in the region of the small, restricted dark brown scapular patch; ocelli
peripheral rather than enclosed (fig. 3a); throat gray, unspotted; ven-
tral ground color flesh-gray; tail nearly unicolor, tan above and yellow-
ish below.
VARIATION: 14 males and 14 females including the type and para-
types have maximum snout-vent length of 33 mm in both sexes and
the following scale counts: dorsal scales axilla to groin 18-22, mean
20.1; ventral scales axilla to groin 24-29, mean 26.8; midbody scales
36-42, mean 39.4; escutcheon 4-8 X 21-27; internasals 0-3, mode 1;
fourth toe lamellae 9-13, mode 10; gular scales usually keeled; chest

1 Three specimens of S. m. ateles (ASFS V6629-30, V6641) were collected by
the senior author at a locality 4.6 km W, thence 4.6 km NW Juana Diaz. These
specimens extend the range of ateles inland from the eastermost locality of Playa
de Ponce and narrow the gap between the races ateles and mimetes in the Juana
Diaz region. The locality is not shown on the map.


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and ventral scales smooth, except for six lizards in which the gular
scales are only partially keeled and one with some keeling on the
chest scales.
Dorsal ground color of males grayish-tan to tan in life, rather
heavily spotted with dark brown; head yellow-tan, with some dark
spotting, especially on the occiput and neck; scapular patch small and
restricted to absent; ocelli inconspicuous, gray, and often not discern-
ible; throat yellow-orange, with scattered dark brown flecks form-
ing a diffuse and narrow collar in some specimens; ventral coloration
grayish to flesh-gray.











S.I

A B C_

FIGURE 3. A, S. macrolepis ateles, anterodorsal view, female, ASFS V5525, para-
type, Balneario de Boquer6n, Puerto Rico. B, S. macrolepis guarionex, antero-
dorsal view, female, MCZ 81048, type, Officers' Club Beach, Ramey Air Force
Base, Puerto Rico. C, S. macrolepis inigoi, anterodorsal view, female, MCZ
81055, type, Ensenada Sun Bay, Isla Vieques.

Females tan dorsally in life, with regular and prominent dark
brown longitudinal lines; head ground color unicolor with that of the
back, and with a bold and conspicuous dark brown head pattern.
In four specimens the occipital spot is joined to the postorbital lines,
forming a more complex figure. The scapular patch is small and
may be represented by a longitudinal stripe with the white ocelli
along its lateral margins. The throat is gray and unpatterned, and
the ventral ground color is flesh-gray.
Four juveniles are much like the females, with prominent head
patterns, restricted scapular patches, and longitudinal dark lines.
The three geckos from 5 mi. S Playa de Mayagiiez (MCZ 34460-
62) are regarded as ateles more on geographical grounds than for any
other reason. One male has a larger scapular patch with included


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ocelli than that of any other male; the female has the head pattern
faint rather than bold, and also a large black patch with included
ocelli. They may represent a variant local population within the
southwestern subspecies, but as this locality is quite close to two
recent localities for ateles, possibly the labeling of the three old speci-
mens is in error.
COMPARISONS: From its neighbor S. m. mimetes to the east,
ateles can be readily distinguished by lack of a heavily spotted throat
in either sex and the extreme reduction of the scapular patch with
the ocelli peripheral. The lower number of midbody scales (mean
36.1) of mimetes is significantly different from the higher number
(mean 39.4) of ateles. From the more northern subspecies grandi-
squamis, stibarus, and phoberus, the southwestern race differs in color,
in having females with prominent lines and with bold and vivid head
patterns. In midbody scales ateles differs significantly from phoberus
and grandisquamis but not from stibarus (table 1). Juveniles of ateles
are lined in contrast to the unicolor juveniles of the northeastern
races.
REMARKS: S. m. ateles occupies southwestern Puerto Rico north
of the Valle de Lajas; south of this valley, it is replaced by the large
S. roosevelti. The two species approach each other closely (9 km)
at Boquer6n and Pole Ojea, but have never been taken together.
Sphaerodactylus roosevelti occupies the southern coast as far east as
Punta Ventana, and the first southwestern coastal record for ateles
is from Playa de Ponce. Schmidt (1928:.71) reported "S. grandi-
squamis" from Ensenada, where we are reasonably certain that S.
macrolepis does not occur; examination of Schmidt's material shows
that he had at least S. roosevelti and S. nicholsi from this locality.
Stejneger (1904: 608) reported taking 8 specimens of "S. grandi-
squamis" in the white clay hills about 3 miles east of Ponce. His
detailed description of these specimens does not mention the heavily
spotted throat of mimetes, the race we would expect in that area.
His descriptions and figure agree very well with S. m. ateles, even to
the almost longitudinally strap-like scapular patch in the female.
Although we have not examined his material, we regard -these geckos
as S. m. ateles. The single specimen (UMMZ 73627) from Playa de
Ponce is clearly ateles and confirms this supposition.
The southwestern race was collected in palm trash along the
coast at Balneario de Boquer6n, localities SSW of Mayagiiez, and
inland under rocks on exposed hillsides on the southern slopes of the
Cordillera Central above Yauco and Susua. In this same region, sev-
eral specimens were taken in a wet and shaded ravine.


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214 BULLETIN FLORIDA STATE MUSEUM Vol. 10

Sphaerodactylus macrolepis spanius, new subspecies
TYPE: MCZ 81047, an adult female from 17.7 km NE Utuado
(= about 8 km airline), 1100 feet, Puerto Rico, taken 20 January
1965 by Donald W. Buden and Richard Thomas. Original number
V4618.
PARATYPES (all from Puerto Rico): ASFS V4617, V4619, same data as type;
USNM 15397-98, UIMNH 56936-37, UF 21259-60, KU 79904-05, same locality
as type, 1 September 1962, R. F. Klinikowski, D. C. Leber, A. Schwartz; ASFS
V4490-92, V4502, same locality as type, 17 January 1965, R. Thomas.
ASSOCIATED SPECIMENS (all from Puerto Rico): ASFS V5194, 6 km SE Las
Marias; MCZ 34492-93, MCZ 62182-83, Maricao; ASFS X4258-65, 13.8 km N
Sabana Grande, 2800 feet; ASFS V6022, 2 km NE Barranquitas, 2100 feet; ASFS
X7572, 8 km SE Las Cruces, 2080 feet, Bosque de Guavate; intergrades between
S. m. spanius and S. m. mimetes: AMNH 13035, 13037, Aibonito.
DISTRIBUTION: The interior uplands of the Cordillera Central
and the Sierra de Cayey at elevations from 1100 feet to 2800 feet.
Intergrades with S. m. mimetes at Aibonito (fig. 1).
DIAGNOSIS: A subspecies of S. macrolepis characterized by a com-
bination of large size (maximum snout-vent length of males 34 mm,
of females 35 mm), high number of midbody scales (41-47, mean
43.6 .33). Males grayish-tan to yellowish-brown with dorsal dark
scales arranged in longitudinal rows, much like females only consid-
erably fainter, Or unicolor; head ground color yellow, with a faint
residual female pattern, often with additional dark spots on the neck;
scapular patch small and restricted but with two white included
ocelli; throat gray to bright yellow, at times with a few faint brown
dots. Females light brown to tan with fairly prominent longitudinal
lines; head light brown with a bold head pattern, nuchal and scapu-
lar patches often joined; scapular patch variable in size, usually mod-
erate without a pale edge and with the ocelli included; throat gray,
often with some diffuse darker gray marbling; iris variable, yellow-
green, golden, or brown.
DESCRIPTION OF TYPE: An adult female with a snout-vent length
of 33 mm, tail 33 mm. Dorsal scales axilla to groin 23; ventral scales
axilla to groin 28; midbody scales 42; fourth toe lamellae 12; inter-
nasal 1; 3 supralabials to mid-eye; gular scales keeled, chest scales
partly keeled, ventral scales smooth. Dorsal ground color light
brown with four regular longitudinal lines with lighter brown edges;
head light brown with a dark pattern of a preocular V representing
the usual pair of snout lines, a wide pair of postorbital lines which
form a V by joining the nuchal spot and enclosing the occipital spot
between them, a dark canthal line passing through the eye and pro-








PUERTO RICAN SPHAERODACTYLUS


ceeding onto the neck and thence almost to join the scapular patch;
scapular patch moderate in size with no pale margin and including
two dull ocelli; throat gray with some darker gray stippling along
the margin of the lower jaw; ventral ground color yellowish; under-
side of tail bright orange.
VARIATION: 11 males and 16 females, including type and para-
types have maximum snout-vent lengths of 34 mm in males and 35
mm in females and the following scale counts: dorsal scales axilla
to groin 19-26, mean 22.0; ventral scales axilla to groin 24-32, mean
28.0; midbody scales 41-47, mean 43.6; escutcheon 4-7 X 19-27; inter-
nasals 0-3, mode 1; fourth toe lamellae 8-12, mode 10. Spanius shows
a higher incidence of chest-scale keeling than other races; 8 speci-
mens have some chest scales keeled and 2 have all the scales keeled.
The normal condition is to have the gular scales keeled and the chest
and ventral scales smooth.
The dorsal ground color in males varies in life from grayish-tan,
yellowish-tan, or tan, to light brown. One lizard (ASFS X4473) has
a dense salt-and-pepper pattern, two (ASFS X4258, MCZ 34492) are
unicolor, and the balance have the faint longitudinal lines typical of
females of several races of S. macrolepis, including female spanius.
The head is distinctly yellowish and shows fairly prominent remnants
of the female head pattern, especially the pale postorbital or tem-
poral lines, which may be outlined or indicated by some dark brown
to rusty spotting; nuchal region may also have some dark spotting;
scapular patch small to moderate in size, never with a pale margin,
never completely absent, and always including two pale ocelli; tail
often grossly spotted; hindlimbs conspicuously banded with buff;
throat bright yellow to gray with small grayish dotting; ventral ground
color pale creamy gray to yellowish or faintly orange; underside of
tail orange; iris variable: yellowish north of Sabana Grande, yellow
near Las Marias, yellow to brown near Utuado, golden near Barran-
quitas, or bright yellow at Guavate. The brighter iris colors are
found in the eastern portion of the range.
In life the dorsum of females is tan to light brown with prom-
inent and regular longitudinal lines; head pattern like the type; three
specimens have the nuchal spot joined with the scapular patch, and
several have these two elements approximating each other; scapular
patch variable in size and usually including two ocelli, although five
females have the ocelli peripheral and one (ASFS X4259) apparently
lacks them completely and has a much reduced scapular patch; tail
dull orange above with some indication of clearer orange crossbands,
or plain with a yellowish wash; underside of tail from pinkish to


1966


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BULLETIN FLORIDA STATE MUSEUM


bright orange; throat gray, sometimes vaguely marbled with slightly
darker gray; ventral ground color from creamy or yellowish to gray.
Three juveniles are like the females, having pale tan dorsa with
faint longitudinal lines and a scapular patch; one has the ocelli in-
cluded within the patch, and the two smaller have the ocelli periph-
eral. As usual the tail tip in the smallest is white, preceded by a
darkened band.
COMPARISONS: S. m. spanius differs from all other races in hav-
ing longitudinally lined males. This upland race is larger than any
other subspecies, being approached most closely by ateles, grandi-
squamis, and the race from the north coast. It is also noticeably
bulkier than the other forms. S. m. spanius has more midbody scales
than any other race (mean 43.6) (table 1). The absence of a dark
spotted throat separates spanius from mimetes. No other race thus
far described has the head pattern as fused with the nuchal and scap-
ular patches as does spanius.
REMARKS: Two specimens from Aibonito (AMNH 13035, 13037)
are clearly intermediate between spanius and mimetes. These lizards
have very heavily spotted throats like mimetes, and the female has
44 midbody scales (the male is uncountable), a spanius character.
The male (32 mm) is at the upper size limit of mimetes, and the
female (31 mm), is slightly larger than any female mimetes. Consid-
ering that Aibonito lies at an intermediate elevation of about 1800
feet, and is more or less geographically between Barranquitas (span-
ius) and Patillas and Coamo (mimetes), the locality seems appropri-
ate for intergradation between these two races.
The range of S. m. spanius is long, extending from Maricao in the
west to Las Cruces in the east, and we do not feel that the popula-
tions of this race are continuous. Large regions seem unsuitable for
it as, for example, the bare mountains between Casa Blanca-Orocovis-
Barranquitas, though in such areas the species may occur in residual
wooded ravines as at Barranquitas. We have also failed to find
spanius in some areas that appear suitable, as at Monte Guilarte,
Cerro de Punta, and Cerro de Dofia Juana. It is not plentiful at
Guavate. Sphaerodactylus klauberi is the dominant gecko at most
of the upland localities where spanius is rare or absent.
Sphaerodactylus m. spanius is more commonly found in forested
areas than any other subspecies. However, within such areas it is
often most common or more easily secured about clearings where
there is trash. At the type locality, Hacienda Roses, we found these
lizards under trash and leaves at the base of banana plants in a high-
canopied cafetal. At Barranquitas a single specimen was found in


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PUERTO RICAN SPHAERODACTYLUS


a similar situation, where the coffee grove was in a wooded and
moist ravine. North of Sabana Grande the series was taken about
the edges of a temporarily disused picnic area. The animals were
extremely active at dusk, when several were taken crossing open
tracts of lawn grass.

Sphaerodactylus macrolepis guarionex, new subspecies
TYPE: MCZ 81048, an adult female from Officers' Club Beach,
Ramey Air Force Base, Puerto Rico, one of a series taken 23 Febru-
ary 1965 by Albert Schwartz and Richard Thomas. Original number
V5402.
PARATYPES (all from Puerto Rico): ASFS V5400-01, V5403-06, same data
as type; ASFS V5408, V5415, AMNH 94195-98, MCZ 81049-54, KU 79906-09,
UF 21261-64, Rifle Range Beach, Ramey Air Force Base, 23 February 1965, A.
Schwartz, R. Thomas; ASFS V5477, 1 km E Ramey Air Force Base, 24 February
1965, R. Thomas.
ASSOCIATED SPECIMENS (all from Puerto Rico): ASFS V5127-35, 0.9 km S C6r-
sega; ASFS V3204-19, 1.0 mi. SW Aguada; ASFS V3225-26, ca. 1.5 mi. SW Agua-
dilla; ASFS V5104-13, 3 km NW Quebradillas; ASFS V5327-28, 7.0 km S Mora,
800 feet; ASFS V5301-04, 11 km S Mora, 800 feet; ASFS V5149, 9.6 km S, 0.8 km
E Mora, 700 feet; ASFS V3227, 2 mi. S Pueblito de Ponce; ASFS V6152, 2 km S
Pueblito de Ponce, 800 feet; ASFS V5181, 1 km S Pueblito de Ponce; ASFS
V5245, 7.2 km SE Quebradillas, 900 feet; ASFS V5880, 4 mi. W Las Llanadas,
600 feet; ASFS V5914-16, 18.8 km SW Arecibo, 800 feet; ASFS V5884-90, 8
mi. E Arecibo; ASFS X7499-500, 0.7 mi. E Palmas Altas; ASFS X7503, 1.9 mi.
W Palmas Altas; ASFS X7516-19, 2.8 mi. W Palmas Altas; ASFS V5996-97, 2.5
km SW Florida; ASFS V5854-65, 10.2 km E Dos Bocas; ASFS V5963-67, Playa
Mar Chiquita; MCZ 12242, Manati; UMMZ 73614 (2 specimens), 2 mi. W
Morovis; ASFS X7674-79, 6 mi. WNW Dorado; ASFS X7680-92, 5.8 mi. NW
Catafio; AMNH 12983, Catafio; MCZ 34511, 34513-18, Camp Buchanan; MCZ
34496-504, 1 mi. NE Bayam6n; UMMZ 73610 (2 specimens), between Caguas
and Juncos; MCZ 34490-91, Gurabo.
DISTRIBUION: Northwest and north-central Puerto Rico, from
Punta Higuero in the west, eastward as far as Gurabo, and inland to
include the Pepino Hills as far south as Florida (fig. 1).
DIAGNOSIS: A subspecies of S. macrolepis characterized by a com-
bination of moderate size (maximum snout-vent length of both sexes
33 mm), high number of midbody scales (36-49, mean 41.3 .27).
Males yellowish-brown to chocolate brown dorsally with scattered
dark brown scales; head dull yellowish, often with a pair of pale
buffy postorbital stripes .and some dark head spotting; scapular patch
large and almost always present, at times somewhat disintegrated,
and enclosing two large white ocelli; throat without extensive dark
markings, occasionally with a brown stippled "collar". Females yel-


1966


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BULLETIN FLORIDA STATE MUSEUM


lowish-tan to brown with about five dark brown and fairly regular
longitudinal lines; head pattern bold with the nuchal spot often ex-
tensive and fused either with the scapular patch or with the dark
postorbital lines and occipital spot into a large dark occipito-nuchal
figure; scapular patch large, without pale edges, and including two
large white ocelli (fig. 3b); throat with some faint tan dotting or stip-.
pling; iris variable, greenish-yellow, black suffused with yellow, gold-
en, or grayish-yellow.
DESCRIPTION OF TYPE: An adult female with a snout-vent length
of 32 mm, tail 26 mm. Dorsal scales axilla to groin 23; ventral scales
axilla to groin 29; midbody scales 43; fourth toe lamellae 11; inter-
nasals 2; 3 supralabials to mid-eye; gular scales keeled, chest and ven-
tral scales smooth. Dorsal ground color tan with four wide dark
brown longitudinal lines; head ground color straw with a prominent
pattern of a preocular reversed V on the snout, a broad pair of dark
postorbital lines fused broadly posteriorly with the occipital and
nuchal spots into a large occipito-nuchal figure, and a bold canthal
line through the eye and onto the side of the neck, fusing broadly
with the scapular patch; scapular patch large and extensive, without
pale edges, and enclosing two fairly large and bold white ocelli;
throat yellow with a few scattered dark brown flecks; ventral ground
color pale yellow, underside of tail yellow, upper side of tail yellowish
with diffuse dark brown blotching.
VARIATION: 104 adults of S. m. guarionex, including type and
paratypes have a maximum snout-vent length of 33 mm in both sexes
and the following scale counts: dorsals axilla to groin 18-26, mean
21.3; ventrals axilla to groin 22-31, mean 27.4; midbody scales 36-49,
mean 41.3; escutcheon 4-9 X 16-29; internasals 0-3, mode 1; fourth
toe lamellae 8-13, mode 11. One lizard has 4 supralabials to mid-eye
on both sides. Although most specimens have the gular scales keeled
and the chest and ventral scales smooth, 8 specimens have the gular
scales incompletely keeled, 28 have the chest scales partly keeled,
and three have the chest scales completely keeled. A single gecko
has the keeling extending onto the anterior portion of the belly
scales as well.
The dorsal ground color of males in life varies from tan or yellow-
ish-tan to brown, with scattered dark brown scales, not aligned into
longitudinal rows; head yellowish to distinctly orange; the pale (buff)
postorbital stripes are often faintly visible, and may be outlined by
irregular dark brown blotching; occipital spot diffuse and often ab-
sent; nuchal spot more often absent than present, and when present
rather vague; scapular patch fairly large, though it may be reduced


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PUERTO RICAN SPHAERODACTYLUS


or even asymmetrical, and the ocelli large, white and enclosed within
the patch; throat yellow to yellow-orange or bright orange with at
times rather dim brownish stippling or spotting, often forming a
poorly defined collar; ventral ground color gray to whitish or pink-
ish yellow to clear yellow; underside of tail yellow to bright orange,
often with large dark spots.
The females resemble the males in dorsal ground color and usually
have a series of four or five longitudinal dark brown lines; head
pattern typical of the species, except that the nuchal and occipital
spots and the postorbital lines tend to fuse into a large dark brown
occipital figure. Even when not fused, the nuchal spot is often
large and extensive, at times approaching the scapular spot; scapular
spot large, lacking any pale outline, and regularly including two large
white ocelli. Only occasional specimens (6 of 61 females) show any
reduction or obliteration of the scapular patch. The throat is yellow,
usually with at least some indication of brownish or tannish dots
which are inconspicuous after preservation. The throat markings
are never prominent, even in life. The ventral ground color is vari-
able as is that of the males, ranging from grayish-white to yellow.
The underside of the tail is yellow to orange.
The iris color is yellow to grayish-yellow near Palmas Altas, black
suffused with pale yellow and with a pale pupillary ring near Que-
bradillas, greenish-yellow to golden near Mora, brown near Dos
Bocas, or golden at Ramey AFB.
Juveniles are longitudinally lined dorsally, although the lines may
be obscured by the dark brown dorsal ground color; the scapular
patch is large and prominent, the ocelli are included within, and the
head pattern is like that of the female, although the various elements
are less bold; the ventral ground color may vary between grayish
and very pale yellow, and the underside of the tail from yellow to
orange.
COMPARISONS: In midbody scales S. m. guarionex differs signifi-
cantly from all other races except grandisquamis (fig. 6), from which it
can be distinguished by its lack of a pale outline to the scapular patch
and its round rather than transversely elongate ocelli. The consist-
ently included ocelli differentiate guarionex from ateles; the restricted
scapular patch of the latter is also noteworthy. The lined males of
spanius are quite different from those of guarionex. The heavily
spotted throats of mimetes are very different from the more diffusely
and paler spotted throats of guarionex. S. m. phoberus lacks lined
females. Persistent head patterns in male mimetes differ from the
condition in guarionex.


1966


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BULLETIN FLORIDA STATE MUSEUM


REMARKS: The trivial name guarionex is derived from the Mon-
tafias Guarionex, a portion of the Pepino Hills in which S. m. guario-
nex occurs.
The distribution of S. m. guarionex is the most extensive of any of
the Puerto Rican races of S. macrolepis. Ecologically the race occurs
from sandy beaches and coastal coconut groves to elevations of 800
feet in the forested or pseudo-forested dolines of the Pepino Hills.
Inland it was more often encountered in association with banana trash
than in natural situations, very much like spanius in the Cordillera
Central. Of the two species of geckos in the Pepino Hills, S. m.
guarionex is by far the less common, and S. klauberi outnumbered S.
macrolepis where the two were taken together. The major center of
abundance of guarionex is coastal, where it was collected in habitats
typical of those for the species in coastal localities elsewhere, coco-
nut groves being especially favored. At the type locality and at Rifle
Range Beach, the lizards were abundant in lowland coastal decidu-
ous woods where they were found under logs and trash in the heavy
leaf litter. Farther east along the coast occasional specimens were
encountered in Coccoloba leaves.
No intergradation is known between spanius or ateles and guario-
nex. The Dos Bocas specimens from a coffee plantation immediately at
the foot of the Pepino Hills escarpment and only about 5 kilometers
from the type locality of spanius, are definitely guarionex. This is
the closest these two subspecies are known to approach each other.
Presumably the valley between the escarpment and the ascending
slopes of the Cordillera Central divides the two races. The two lo-
calities where guarionex and ateles approach most closely are C6rsega
(guarionex) and south of Mayagiiez (ateles). We could find no suit-
able localities for S. macrolepis in the intervening region, a distance
of about 27 kilometers. Possibly the lowlands in the region of Afiasco
act as a barrier between these two races.

Sphaerodactylus macrolepis inigoi 1, new subspecies
TYPE: MCZ 81055, an adult female from Ensenada Sun Bay
(= Ensenada Sombe), Isla Vieques, Puerto Rico, one of a series taken
2 September 1964 by Richard Thomas. Original number V4042.
PARATYPES (all from Isla Vieques): ASFS V4040-41, V4043-46, same data
as type; ASFS V4028-30, V4037-38, UF 21265-70, UIMNH 56938-44, 7 km W
1 We are aware that the International Code of Zoological Nomenclature, 1961,
Appendix C, article 13 suggests that the Spanish 1i be represented by gn, but we
prefer to render it as n in this case in order to retain the visual approximation of
the written word as nearly as possible.


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PUERTO RICAN SPHAERODACTYLUS


Isabel Segunda, 1 September 1964, R. Thomas; ASFS V6147-51, Colonia Puerto
Real, 24 March 1965, R. Thomas; ASFS V6155, approximately 1.5 km NE Colonia
Puerto Real, 24 March 1965, R. Thomas; ASFS V6156-61, 5 km SW Isabel Se-
gunda, 24 March 1965, R. Thomas; AMNH 94201-03, bay bordered by Punta
Negra, 24 March 1965, R. Thomas; AMNH 94199-200, Puerto Mosquito, east of
Ensenada Sun Bay, 25 March 1965, R. Thomas; KU 79910-16, 3.8 km W Esperan-
zas, 25 March 1965, R. Thomas; MCZ 34414-23 + 33 untagged specimens, Isla
Vieques, 1931-32, C. Grant.
ASSOCIATED SPECIMENS: ASFS V6169-71, Cayo de Afuera, Isla Vieques;
ASFS V4068-69, V6185, Cayo de Tierra, Isla Vieques.

DISTRIBUTION: Isla Vieques and the two southern islets, Cayo
de Afuera and Cayo de Tierra (fig. 5).
DIAGNOSIS: A subspecies of S. macrolepis characterized by a com-
bination of small size (maximum snout-vent length in both sexes 30
mm), low number of midbody scales (33-41, mean 36.8 .32). Males
with dorsal ground color yellowish-brown and without pattern except
for occasional scattered darker scales; head orange to rusty and with-
out pattern except for a dull occipital spot and occasional remnants of
the nuchal spot; scapular patch small, usually medially constricted
and including two tiny white ocelli; and throat almost always un-
patterned. Females brown with a pattern of longitudinal dark lines;
ground color of head gray to yellow-brown with typical head pattern
at times somewhat fragmented; throat always heavily stippled or
blotched with dark brown to black; iris brown.
DESCRIPTION OF TYPE: An adult female with a snout-vent length
of 27 mm, tail 28 mm. Dorsal scales axilla to groin 20; ventral scales
axilla to groin 27; midbody scales 40; fourth toe lamellae 10; inter-
nasal 1; 3 supralabials to mid-eye; gular scales keeled, chest and ven-
tral scales smooth. Dorsal ground color brown with two dorsolateral
longitudinal lines and a median series of four irregular blotches which
represent the two paramedian longitudinal lines; head ground color
gray with a dark pattern consisting of a preocular reversed V, which
is the usual pair of snout lines, a pair of irregular postorbital lines
which fuse with the nuchal spot, enclose the occipital spot, and almost
meet the scapular patch, and a dark canthal line through the eye
which continues onto the neck and becomes confluent with the dor-
solateral body line; scapular patch small, without pale margins and
enclosing two small indistinct ocelli (fig. 3c); tail brown dorsally with
a series of four pale irregular crossbands, each outlined with darker;
throat gray with a dense reticulum of dark gray; ventral ground color
pale gray.
VARIATION: 22 males and 15 females, including type and para-
types and associated specimens, have a maximum snout-vent length


1966







BULLETIN FLORIDA STATE MUSEUM


of 30 mm in both sexes and the following counts: dorsals axilla to
groin 17-21, mean 18.6; ventrals axilla to groin 21-27, mean 24.6; mid-
body scales 33-41, mean 36.8; escutcheon 5-8 X 19-26; internasals 1-3,
mode 1; fourth toe lamellae 8-12, mode 10. All specimens have 3
supralabials to mid-eye; exceptions to having the gular scales keeled
and the chest and ventral scales smooth are four lizards with the gular
scales only partly keeled.
The males present little variation in color or pattern. All are
tan, gray-brown, or yellow-brown dorsally, and only occasional speci-
mens have any dorsal body markings in the form of isolated dark
scales. The head is orange or rusty without pattern except for the
vague occipital spot and rarely a remnant of the nuchal spot which
regularly is absent; scapular spot regularly present, small, and usually
constricted medially into a dumb-bell shape, and the ocelli are white
and small. Three males have the scapular patch almost or entirely
obliterated. The throat is clear gray or yellowish to orange, normally
without any pattern, although two males have a heavily reticulate
throat. The venter is light gray, and the iris brown.
The females were recorded as having a brown dorsal ground color
in life; and usually, in contrast to the type, they have four longitudinal
lines; seven females resemble the type in having a single median row
of dark blotches resulting from the fragmentation and subsequent
fusion of the two paramedian longitudinal lines. The head pattern
is like that of the type. The occipital spot may be very closely ap-
proximated to and almost fused with the postorbital lines. The dark
head elements are always irregularly scalloped and rough-edged and
are not so neat and diagrammatic as those of other races. The scapular
patch is small, often restricted, not outlined with pale, and may have
the ocelli included or on the periphery, depending upon the size of the
patch itself. The throat is pale gray, always with a heavy dark brown
to black reticulum. The venter is light gray. Unregenerated tails
are brown and show some crossbanding, the pale bands outlined with
dark brown.
Juveniles show the same variation in dorsal pattern as the fe-
males; 7 of 14 show the median series of dark dorsal blotches rather
than lines; no juvenile shows a clear pattern of four dorsal lines,
though several have this indicated. The head pattern is like that of
the females, and one juvenile has the nuchal spot fused to the scapular
spot. Tail tips of very small lizards (16 and 17 mm) are very pale
gray rather than white.
COMPARISONS: The only mainland Puerto Rican race of S. macro-
lepis with which inigoi requires comparison is mimetes. Both are the


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PUERTO RICAN SPHAERODACTYLUS


only subspecies that regularly have very heavily spotted throats in
one or both sexes. In mimetes the throats are spotted in both sexes,
and the males retain the female head pattern, accenting it with dark
dorsal head spotting; inigoi males have brightly colored heads with-
out spotting or pattern, and almost always have the throats without
pattern. Females of the two races are very much alike, as both have
head patterns and spotted throats. The latter is a much more con-
stant feature in inigoi, for all females examined possess it. The irreg-
ular median series of dorsal blotches occurs in inigoi and not in fe-
male mimetes. The two races show no scale differences, but male
mimetes reach a larger size than either sex of inigoi. Detailed com-
parisons with the other races on Puerto Rico is unnecessary; none has
the heavily spotted throat of female inigoi and the regularly unpat-
terned males with brightly colored heads. The mean number of mid-
body scales of inigoi is significantly different from those of grandi-
squamis, stibarus, ateles, spanius, and guarionex (table 1).
REMARKS: We take great pleasure in naming the Vieques race
of S. macrolepis for Sr. Felix Ifiigo of the Department of Agriculture
and Commerce of the Commonwealth of Puerto Rico, for his unfail-
ing courtesy toward us in our endeavors in his country.
Stejneger (1904: 602 et seq.) included the Vieques sphaerodactyls
in his newly described species grandisquamis. By placing grandi-
squamis in the synonymy of S. macrolepis, Schmidt (1928) put the
Vieques geckos also in that species. Grant (1932b, chart) considered
the Vieques lizards as S. danforthi Grant, a species very close to mac-
rolepis that he described from Isla Culebra to the east and character-
ized (1931: 205-06) as having a "vividly speckled throat, black and
white, stopping abruptly at the neck" and by males having either a
red-headed or speckled-headed phase. In two subsequent papers,
one dealing with the herpetofauna of Vieques (1932c) and another
(1932d) dealing with the reestablishment of S. grandisquamis as a
distinct species, he continued to regard the Vieques animals as
danforthi.
The above brief historical outline shows how the Vieques geckos
have puzzled everyone who has studied them. As we do not regard
danforthi as distinguishable from S. macrolepis even subspecifically,
Grant's assignment of the Vieques population to that form is un-
tenable. We have already commented on the fact that in scale
organs inigoi is clearly associated with mainland Puerto Rican races,
rather than with S. m. macrolepis to the north and east. These two
races are distinct not only in scale organs but in head coloration and


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224 BULLETIN FLORIDA STATE MUSEUM Vol. 10

throat pattern of males; the females of each are extremely similar to
each other.
On Cayo de Tierra inigoi was collected under rocks, primarily on
an exposed and sunny hillside. On Vieques proper these lizards were
taken in mesic ravines, in coastal Coccoloba, and in palm trash; the
sea-grape habitat yielded the most specimens.

Sphaerodactylus macrolepis macrolepis Giinther
Sphaerodactylus macrolepis Giinther, 1859, Ann. Mag. Nat. His. (3), vol. 4,
p. 215.
Sphaerodactylus danforthi Grant, 1931, Jour. Dept. Agri. Porto Rico, vol. 15,
no. 3, p. 205. Type locality Culebra Island. Type specimen MCZ 34403.
TYPE LOCALITY: St. Croix, American Virgin Islands.
DISTmIBUTION: Culebra including Culebrita and Cayo Luis Pefia;
American Virgin Islands, St. Croix including Protestant Cay; St.
Thomas and its satellites, Water, Buck, Hans Lollick, Little St. James,
Little Saba, Savana, Cockroach; St. John including Congo Key; and
British Virgin Islands, Tortola and Beef, Guana, Great Camanoe,
Peter, Virgin Gorda, Anegada (fig. 5).







i# '\
A_







A B" C
FIGURE 4. A, S. macrolepis macrolepis, anterodorsal view, male, ASFS V3306,
1.0 mi. W Cane Bay, St. Croix, American Virgin Islands. B, S. monensis, antero-
dorsal view, female, MCZ 34571, Isla Mona. C, S. roosevelti, anterodorsal view,
female, ASFS V5627, 8.5 km SE Guanica, Puerto Rico.

DIAGNOSIS: A subspecies of S. macrolepis characterized by a
combination of small size (males to 80 mm snout-vent length, largest
females 31 mm), dorsal scales with only hair-bearing organs, high
number of midbody scales (88 to 50, mean 43.1 .27). Dorsum of
males gray or tan to brown, usually without any darker dorsal scales








PUERTO RICAN SPHAERODACTYLUS


but occasionally with a few scattered dark scales; head contrastingly
colored either with dark brown or black vermiculations on a pale gray
to pinkish- or purplish-gray ground color, or brightly colored in ochre,
orange, or rich reddish-brown, and virtually patternless except for a
few diffuse dark nuchal dots and an indistinct occipital spot; scapular
patch and ocelli always very small, often both absent; throat heavily
marbled black on a yellow to yellow-orange ground color. Females
yellowish-tan to dark brown with a pair of dorsolateral and a pair of
lateral brown stripes and a median series of about four or five brown
blotches which may be much fragmented and salt-and-pepper in
effect; dark head pattern composed of a preorbital reversed V, a pair
of postorbital lines, usually constricted at the level of the occipital
spot but continuing to join the nuchal spot posteriorly, all the head
lines irregular and scalloped in outline on a gray ground color; scap-
ular patch fairly large but variable in extent, rarely reduced or ab-
sent, and including two pale cream ocelli; throat variable but usually
with some dark markings, less extensive than those of males, on a
whitish or yellowish ground. Iris yellow to pale yellow.
REMARKs: The nominate subspecies of S. macrolepis occupies a
wide range throughout the islands between Puerto Rico and the
Anegada Passage. Although it has been reported from relatively
few of these islands, it is presumed to be far more widespread than
the reports indicate.
The coloration of a series from St. Croix in life follows: Males-
dorsal ground color gray to yellowish-brown with a few dark scales
sprinkled over the back, or with merely a few dark scale edges or
no dark scales whatsoever; head usually grayish with heavy dark
vermiculations which represent the typical female head pattern much
obliterated by additional black pigment; the nuchal spot is absent,
and the dark head vermiculations end abruptly on the neck; ocelli
white, connected by a rudimentary V-shaped scapular black patch;
both scapular patch and ocelli sometimes completely absent (fig. 4a);
throat yellow with faint dark marbling or white with heavy dark
marbling, the dark pigmentation stopping abruptly at the same
nuchal level as the dorsal head markings; venters white; underside
of tail pale yellow. A very few males from St. Croix have the head
unmarked dorsally and more or less unicolor with the yellowish-
brown body. Females-dorsal ground color gray with a pair of
lateral stripes on the lower sides just above the limbs, a pair of dorso-
lateral stripes, enclosing between them a series of four or five median
dark blotches, which in turn may be much fragmented giving a rather
irregularly patterned median dorsal area; scapular patch fairly large


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BULLETIN FLORIDA STATE MUSEUM


and dark-edged enclosing a pair of cream, ocelli, but patch occasion-
ally absent; tail yellowish dorsally with light crossbars proximally;
throat whitish with some faint, usually lateral, dark markings; head
gray with a bold but irregular dark brown pattern consisting of a
preocular reversed V which additional dark pigment on the snout may
make very irregular, an, irregular and much scalloped pair of post-
orbital lines constricted at the level of the occipital spot which ap-
proach and sometimes join the nuchal spot; a few specimens have
some dark pigment in the gray areas on the head. A dark canthal
line passes through the eye and along the neck, where it usually joins
the lateral line. The ventral ground color is gray to pinkish, the
underside of tail orange to yellowish.
We have seen living specimens of S. m. macrolepis from 13 islands
(other than St. Croix) from Culebra to Anegada and find that, except
for the head color and pattern of males, little variation is evident in
dorsal patterns and color. The major intrapopulation differences are
the frequencies of brightly-colored headed males.
Grant (1931: 205-07) diagnosed S. danforthi as follows: "Pro-
nounced sexual dichromatism. Adult averages 50 mm. Squamation
as in S. macrolepis. Males less marked and females more than in
S. macrolepis. The outstanding character is the vividly speckled
throat, black and white, stopping abruptly at the neck." He then
proceeded to discuss two categories of males, "males, red-head phase"
and "males, speckled-head phase". Later (1932c: 37-38) be recorded
the species from Vieques and described the population as "a two-
color-phase male form". Grant found no intermediates between the
two phases on Culebra, but on Vieques "all intergrades were found".
Whether he meant by this that the entire population was intergradient
between danforthi and macrolepis or whether the entire gamut of
intergradient possibilities was encountered is unknown. Still later
Grant (1937: 520) reported S. danforthi fromvAnegada, where "some
males had lemon-yellow heads, others- tlhe ical orange-red heads."
He made no mention of males with patterned heads. Finally Grant
and Beatty (1944) recorded S. danforthi from St Croix on the basis
of four specimens from four localities. They also noted the wide-
spread occurrence of S. macrolepis on St. Croix, with 330 specimens
from 35 localities.
The type of S. danforthi (MCZ 34403) is a male with a patterned
head. So far as we can determine, the pattern of Culebra males
does not differ from that of males from St. Cfoix, or for that matter
from specimens from elsewhere within the raif' pf S. m. macrolepis.
Of 36 males from Culebra, 8, including all tfie largest males, have
i '4


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PUERTO RICAN SPHAEROBACTYLUS


plain (brightly colored in life) heads, and the balance have patterned
heads. Grant (1931: 206) noted that 22 percent of his sample of 67
males were in the plain-headed phase, whereas the balance were
patterned. Obviously, from the data presented by Grant and Beatty
above, the bright-headed males are uncommon on St. Croix, which
our recently collections there substantiate. In a series of 28 males
from Virgin Gorda, 11 had yellow and patternless heads and throats,
and the others had some yellow on the heads plus patterning. Most
of the males collected at Lloyd's Pond on Tortola had the heads yellow-
orange or rich brown, without pattern; only one of the series had the
head gray with black vermiculations as in typical macrolepis. Another
male had the head ground color yellow, but retained the black head
markings; this specimen would be "intermediate" between danforthi
and macrolepis. On Beef Island most males had ochre heads and
yellow to yellow-orange throats. Only a very few had the patterned
macrdlepis type head or showed the transitional stages between this
and the ochre-headed condition.
We regard the bright-headed males as a stage in the ontogeny of
male S. m. macrolepis; as females and young males have patterned
heads it seems logical to consider the bright-headed males as "super-
males"-possibly breeding adults or males approaching old age. The
varying incidence of this condition in populations from various islands
is of interest, but cannot be construed as showing the occurrence of
two species. The fallacy of such interpretation is that one is left with
a certain, albeit often small, percentage of "intergrades," "intermedi-
ates" or "hybrids" in most samples large enough to have various onto-
genetic stages. As there is no other difference between danforthi with
patterned heads and macrolepis, we consequently regard S. danforthi
as a synonym of S. m. macrolepis.
VARIATION: 90 specimens of S. m. macrolepis from several islands
throughout the range (Culebra, St. Croix, St homas, Virgin Gorda,
and Anegada) have a maximum snout-ventftligth of 30 mm in males,
31 mm in females and show the following scale counts: dorsal'scales
axilla to groin 18-26, mean 22.5; ventral scales axilla to groin 21-31,
Mean 25.2; midbody scales 38-50, mean 43.14 escutcheon 3-8 X 11-28;
int6rnasals 0-3, mode 1; fourth toe lamellae 9-12, mode 9. There
are regularly 3 supralabials to mid-eye. Usually the gular scales are
keeled and the chest and ventral scales are smooth. However, some
variation occurs; in a series of 18 specimens from Culebra, 7 have no
keels on the gular, chest, or ventral scales, and 4 Culebra lizards have
only some of the gular scales keeled. In 20 from St. Croix, only 3
have some of these scales keeled, while 5 of 20 from St. Thomas
i


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BULLETIN FLORIDA STAT MUSEUM


and 6 of 20 from Virgin Gorda have some of the scales keeled. One
SVirgin Gorda macrolepis has some of the chest scales keeled as well,
and in 2 of 13 Anegada specimens only some of the gular scales are
keeled, and 3 have the keels on some of the chest scales.
The absence of internasals is apparently unique to Virgin Gorda
S. m. macrolepis; 5 of 84 Virgin Gorda lizards have this condition,
which occurs in none of the 258 specimens examined from other
islands. The mean numbers of dorsal scales in axilla to groin distance
likewise vary between islands, ranging from a low of 21.7 on Culebra
to a high of 23.5 on Virgin Gorda. The mean of 22.2 on Anegada,
the easternmost island inhabited by S. m. macrolepis, is intermediate
between the two extremes. The same situation applies to midbody
scales, where the means range from a low of 41.0 on Culebra to a
high of 44.8 on Virgin Gorda; once again Anegada, with a mean of
42.1, is intermediate between the two. The subspecies reaches a
maximum snout-vent length of 30 mm in males (St. Croix and Virgin
Gorda), and of 31 mm in females (St. Croix). Maximal sizes for
specimens from other islands are somewhat lower, the smallest being
those of Anegadan lizards where the largest male has a snout-vent
length of 22 mm and the largest female 26 mm; however, the series
is small (7 males, 3 females). St. Thomas and Culebra macrolepis
are only slightly larger, males having a snout-vent length of 27 mm,
females 26 mm (43 examined). The apparent smallness of Anegada
lizards suggests an approach toward the smaller S. m. parvus from
the northern Leeward Islands, but Anegada specimens show no
parvus scalation or pattern characteristics. The slight variations that
S. m. macrolepis shows between islands in male head color and in
scalation and size are random and present a crazy-quilt configuration
that does not lend itself to nomenclatorial treatment on any single
character or combination of characters.
A single hatchling from Tortola has a spoubtvent length of 12 mm
and lacks a white or gray tail tip. In -~edi and pattern juveniles
resemble females and show the same variation. Noteworthy is the
rather small size at which the.male dorsal pattern and patterned head
is.assumed, males with snout-vent lengths of as little as 18 mm being
readily identified as such on the basis of tle unicolor dorsum.
S. m. macrolepis occurs from sea level to a known elevation of
750 feet at Rose Lodge, Tortola, and probably reaches \even higher
levels. It is a common inhabitant of wooded situations, both highland
and low, especially where leaf-litter covers the ground, and is un-
usually plentiful in the ruins of Danish plantati houses and wooded
estates on St. John. We encountered this gkd ..rather rarely on


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PUERTO RICAN SPHAERODAWTYLUS


Tortola except on the eastern end where we took numbers in palm
trash piles in a coastal coconut grove. On Beef Island an abandoned
coconut grove yielded a large series, and a few were seen in the more
open scrub. Seagrape stands were a favored habitat on Buck Island.
On Virgin Gorda S. m. macrolepis is widespread, although most read-
ily collected in coastal coconut trash. Grant and Beatty (1944) indi-
cated that macrolepis on St. Croix has a general distribution except
for the "eastern extremity"; we secured no specimens there east of
Milord Point on the west side of Great Pond Bay. The species occurs
on St. Croix from coastal coconut groves and dry maritime woods to
upland woods.
S. m. macrolepis requires comparison only with the races mimetes
from Puerto Rico and inigoi from Vieques: macrolepis is distinguish-
able at a glance from all other subspecies by virture of the usually
blotched dorsum of females and the densely patterned heads and
throats of males, or by the bright-colored heads of males if they are
without pattern. Male macrolepis differ from male mimetes in lacking
remnants of a head pattern and the bold dark head spotting. The
bold head pattern without constrictions and scalloping and the lineate
dorsal pattern of female mimetes separate the females of the two
races. The difference in the mean numbers of midbody scales,
mimetes 36.1 and macrolepis 43.1, are statistically significant (table
1). The absence of knob-like scale organs on the dorsal scales of
macrolepis also separates this race from mimetes.
The scale organs likewise differentiate macrolepis from inigoi-
the latter has both knob-like and hair-bearing organs, the former only
hair-bearing organs. As previously pointed out, inigoi resembles
macrolepis in many details of color and pattern. Male inigoi always
have brightly colored, unpatterned heads and uppatterned throats;
macrolepis males have patterned heads and throats until the bright-
colored head condition is reached. Even when the upper surface
of the head is virtually patternless, as in-ini~as well, the throat is
still patterned with dark, an extremely rare condition in inigoi. The
difference in the mean's of midbody scales, inigoi 36.8, macrolepis
'43.1, are statistically significant. The modal number of fourth toe
lamellae is 9 in macrolepis, 10 in inigoi. The dorsal scales between
axilla and groin are 18 to 26, mean 22.5 in macrolepis, 17 to 21, mean
18.6 in inigoi.
SPECIMENS EXAMINED: St. Croix: ASFS V3268-82, 4 mi. S Lavallee; ASFS
V3615-24, ca. 0.4 mi. E Lavallee; ASFS V3284-300, Cany 6Bay; ASFS V3304-16,
1.0 mi. W Cane Bay; ASFS V3414-17, 2 mi. S Fredericksd ;/ASFS V3421, An-
naly; ASFS V3318-20, 2 mi. NW Annaly; ASFS V3426-32, ir ni. E Annaly; ASFS


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BULLETIN FLORIDA STATE MUSEUM


V3433-36, 2 mi. E Annaly; ASFS V3341-43,' Fountain; ASFS V3552-59, 2 mi.
NW Orange Grove; ASFS V3625-28, Anguilla' ASFS V3317-26, Milord Point
(west side of Great Pond Bay); Culebra: ASFS V4133-37, Dewey; ASFS V4138-
39, ca. 1 km W Dewey; MCZ 34403 (type of S. danforthi), 34404-12 (total 57
specimens), no other locality; Cayo Luis Peiia: MCZ 34424-33 (13 specimens);
Culebrita: MCZ 42370; St. Thomas: ASFS V7270-77, 0.5 mi. SW Mandal;
ASFS V7278-88, 1 mi. SW Mandal; ASFS V7318-28, Dorothea; ASFS V8022-33,
2.7 mi. NW Charlotte Amalie; Water Island: ASFS V7419-22, east side; Buck
Island: ASFS V7466-68; Hans Lollick Island: ASFS V7453, east side; St. John:
ASFS V7488-89, Calabash Boom; ASFS V7491-503, Windberg ruins; ASFS V7565-
83, Frederickdal ruins; Tortola: ASFS 7679-80, east side Sea Cow Bay; ASFS
V7692-94, V7905-11, Pasea Hall Estate; ASFS V7751, Rose Lodge, 750 feet;
ASFS V7873-94, V7937, Jackass Gut; ASFS V7961-75, V8073, Lloyd's Pond; Beef
Island: ASFS V7747-50, V7761-805, western end; Guana Island: ASFS V3947-
68, White Bay; Great Camanoe Island: ASFS V3983-V4001, between Lee Bay
and Cam Bay; Peter Island: ASFS V7846-69, Little Harbour; Virgin Gorda:
ASFS V3660-63, Pond Bay; ASFS V3653-57, V3691-92, ca. 0.5 mi. N Pond Bay;
ASFS V3682, between Little Dix Bay and Savana Bay; ASFS V3718-19, south-
eastern part of island, inland from Copper Mine Bay; ASFS V3723-29, just north
of Garden Rock; ASFS V3747-48, Mahoe Bay; ASFS V3756-83, V3817-50, inland
margin of Salt Pond back of St. Thomas Bay; Anegada: ASFS V3925-33, vicinity
of The Settlement; MCZ 12260-63, no other locality.

Sphaerodactylus macrolepis parvus King
Sphaerodactylus macrolepis parvus King, 1962, Bull. Fla. State Mus., Biol.
Sci., Vol. 7, no. 1, p. 16. Type specimen UF 10034.1.
TYPE LOCALITY: 2.5 mi. W, 0.25 mi. N Phillipsburg, St. Martin.
DISTrIBUTIoN: The Lesser Antillean islands of Anguilla, Dog,
St. Martin, St. Barthelemy, and Tintamarre (fig. 5).
DIAGNOSIS: A subspecies of S. macrolepis characterized by a com-
bination of very small size (maximum snout-vent length 23 mm in
each sex; King, 1962: 16, recorded 24 mm), dorsal scales with hair-
bearing organs only, high number of midbody scales (44-54, mean
48.4 .77); not markedly sexually dichromatic, dorsal ground color
varying between pale pink, tan and browai>th scattered dark brown
scales giving a vague salt-and-pepper effect; black scapular patch
moderate in size to small or absent, with a pale buff border and en-
olosing two buff to cream ocelli, the patch slightly more prominent
in life in females than males; heads washed with yellow; throat yel-
low speckled with brown in males, yellow to orange on base of throat
in females with chin usually cream to very pale yellow and somewhat
speckled with brown; tails pale yellow to dull orange below; ventral
ground color creamy white; iris golden to brdnzy orange. Juveniles
have underside of tail coral; head pattern dfijse and much frag-
mented and often very obscure, but consistig 'of a dark preocu-
1 .14


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PUERTO RICAN SPHAERODACTYLUS


lar V, a pair of postorbital dark lines which approach each other
and form an incomplete dark U behind the, occipital spot which may
be obscure or may have a median anteriad extension toward the inter-
orbital area.
REMARKS: We have 16 specimens of S. m. parvus from the islands
of Anguilla, St. Martin, and St. Barthelemy. These agree well with
King's description and diagnosis in size, color, and scalation. In ad-
dition to the scale counts given above, the escutcheon is composed
of 4-6 X 12-17 scales (King's data for escutcheon size are 4-7 X 19-25)
and there are from 0-3, mode 1, internasals. Two tiny juveniles (snout-
vent lengths 12 and 13) are colored like adults except for the coral
underside of the tail.
King (1962: 18) took specimens under rocks in a cave on Anguilla,
and on St. Martin in leaf litter under rocks in a cave and under rocks
from a stone wall. We found ours on St. Martin in leaves on scrubby
hillsides near the ocean, where we also saw others we did not collect
in rock piles inhabited also by S. sputator Sparrman, on Anguilla
under rocky talus at the base of trees in littoral scrub, and on St.
Barthelemy in piles of dead Opuntia in a pasture adjacent to man-
groves. An egg from St. Martin measured 6.0 mm X 4.5 mm (King,
1962).
S. m. parvus hardly requires comparison with any other subspecies.
From the Puerto Rican and Vieques races, it differs in having only
hair-bearing organs on the dorsal scales, in being much smaller, and
in having a higher mean of midbody scales (48.4) than that of any
other race tableel. The races stibarus, phoberus, mimetes, ateles,
and inigoi show no overlap of midbody scales with parvus. Differ-
ences in pattern need not be repeated; the lack of marked sexual
dichromatism in parvus is of itself definitive.
Compared with its neighbor to the west, S. m. macrolepis, the only
other subspecies which also has only hair-bearing organs on the dor-
sals, parvus is much smaller and less bulkyt The head pattern in
macrolepis is much more conspicuous than mi parvus females, and no
parvus males show either the monochromatic or. heavily patterned
heads of male macrolepis. Female macrolepis have the scapular patch
mIch larger and more prominent than female parvus. The throats
of both sexes of macrolepis are more heavily pigmented than those
of parvus. Midbody scales in macrolepis vary between 38 and 50,
mean 43.1, whereas in parvus the midbody scales range from 44
to 54, mean 48.4. The slightly higher means of parvus in counts of
dorsals and ventrals, and the 10 versus 9 modal number of fourth toe
lamellae are also suggestive. A,


1966






BULLETIN FLORIDA STATE MUSEUM


Specimens examined: Anguilla: ASFS X132, X195, Rendezvous Bay; St.
Martin: ASFS 19928, Mullet Pond; St. Barthelemy: ASFS X265-77, Baie de
SSt. Jean.
Sphaerodactylus monensis Meerwarth
Sphaerodactylus macrolepis var. monensis Meerwarth, 1901, Mitt. Naturh.
Mus. Hamburg, vol. 18, p. 20.
DEFINITION: A species of Sphaerodactylus with very large, acute,
strongly keeled, flattened, imbricate dorsal scales axilla to groin 23-
32; no area of middorsal granules or granular scales; dorsal scales
with hair-bearing organs, each with one hair along the free posterior
edge of each scale; throat scales varying from smooth (rarely) to
keeled, chest scales smooth to completely keeled, ventral scales
smooth (usually) to keeled anteriorly, rounded and imbricate axillaa
to groin 22-32). Scales around midbody 42 to 51. Dorsal scales of
tail keeled, acute, imbricate and flat-lying; scales beneath tail smooth,
rounded, enlarged midventrally; internasals 1 to 3 (mode 1); 3 upper
labials to mid-eye; escutcheon (fig. 11b) short and compact medially
with extensions onto the underside of the thighs but never reaching
the knees (4-7 X 9-18). Color pattern not sexually dichromatic, choco-
late to gray-brown above and gray to yellowish below, with dark
brown dorsal markings varying from isolated dark scales to almost a
dorsal dark reticulum composed of individual darker scales; a black
scapular patch, prominently outlined by very pale gray and including
an irregular and often transversely elongate pair of very pale gray
ocelli; a prominent dark sacral U, at times followed by a paler and con-
comitant U; head pattern not bold, consisting of a pair of dark brown
preocular lines on the snout (which are usually joined to.form a re-
versed V), a pair of brown postorbital lines which proceed posteriorly
and become lost in some vague nuchal blotching but which may join
and enclose a grayish-brown occipital diamond which is usually in-
cised medially by a vague dull brown occipital spot; nuchal spot
absent (fig. 4b); throat immaculate pinkish-gray; iris color brown to
gray-brown. Habitus moderate; snout short and rather broad. Max-
imum snout-vent length 30 mm in both sexes.
TYPE LOCALjTY: Isla Mona.
DIsTRmBUrTIN: Isla Mona; not as yet, known from Isla Monito
(see Rolle et al, 1964: 322).
REMARKS: We have outlined the nomenclatorial history of S.
monensis in the discussion of S. macrolepis.
The amount of dorsal dotting is variable, some specimens being
almost without any dorsal pattern (ASFS V6435) and others having
it rather conspicuous (ASFS V6437). The scaplar patch is variable


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PUERTO RICAN SPHAERODACTYLUS


in extent, often invaded anteriorly by the grayish-brown or chocolate
ground color, and the ocelli are likewise variable, in some cases the
two ocelli being joined to form a single enclosed transverse dash. The
scapular patch is invariably outlined by a prominent pale gray border.
The head pattern is generally very subdued and difficult to determine,
as the posterior head and neck are variously marbled and mottled
with brownish on the lighter ground, and the postorbital lines be-
come lost in this irregularly mottled area. In other specimens of both
sexes, the postorbital lines meet behind the vague occipital spot and
form a pale occipital diamond which may include the slightly darker
spot. The spot may be joined to the dark posterior edge of the dia-
mond, thus incising the pale area. The sacral U is present and prom-
inent in both sexes and juveniles; it is often outlined posteriorly by a
pale clear concomitant tan to brownish U. The tail may be,unicolor,
have a pair of dorsal longitudinal lines, or may even be spotted with
darker.
By virtue of its lacking any sexual dichromatism, S. monensis is
quite distinct from S. macrolepis. The head pattern, such as it is, is
easily derivable from that of S. macrolepis, although it is much fainter
and more diffuse. The head seems shorter and broader than that of
S. macrolepis as well. In having a broadly pale-bordered scapular
patch, S. monensis resembles the eastern subspecies S. m. grandi-
squamis. Without adequate material of S. macrolepis from Puerto
Rico, it is not surprising that various authors considered monensis
a synonym of macrolepis.
Grant (1931: 211) reported taking specimens under trash on the
ground, and Schmidt (1928: 74) noted that he had collected four (of
which three specimens were collected at night) among limestone boul-
ders; two were taken beneath pieces of coral on the flat southern ter-
race. Heatwole (1961: 33) found S. monensis under leaf litter at two
localities, and under debris near the lighthouse. The senior author
found S. monensis extremely abundant; most af pjfr recent material was
collected on the southwestern terrace. TlE7ituations where these
lizards were encountered varied from moderately. mesic woods to
xeric scrub; many were seen in palm trash and even in open Casuarina
plantings among the "needles". S. monensis 4s also moderately abun-
dant in the scrub and scrubby woods on the limestone platform.
SPECIMENS EXAMINED: MCZ 13480, MCZ 34558-75, Isla Mona; ASFS
V6435-62, ASFS V6502-04, ASFS V6568-78, Anclaje Sardinero; ASFS V6489-92,
Uvero Desembarcadero; ASFS V6542-47, between Anclaje Sardinero and Uvero
Desembarcadero.


* 1966


233






BULLETIN FLORIDA STATE MUSEUM


Sphaerodactylus roosevelti Grant
Sphaerodactylus roosevelti Grant, 1931, Joui; Dept. Agri. Porto Rico, vol. 15,
no. 3, p. 203, figs. 4-5.
TYPE LOCALITY: Near Parguera, Puerto Rico.
DISTRIBUTnoN: The southwestern coast from Cabo Rojo to Punta
Ventana and on Caja de Muertos (figs. 1 and 5); probably the island
of Vieques (Grant, 1932c). This is the first record for Caja de Mu-
ertos. The species is most abundant in the littoral zone and is found
sparingly in the xeric hills about three kilometers inland east of
GuAnica. It may extend slightly farther east than Punta Ventana,
but it does not appear to reach Ponce. The Vieques record is prob-
ably correct; Grant based it on a hatchling of uncertain identification.
A juvenile specimen (MCZ 36666), whose "Virgin Island" locality
may well be mis-copied for Vieques Island, is probably the specimen
Grant recorded. It is unquestionably a roosevelti. The bright yellow
coloration Grant noted agrees with the color we recorded for a single
hatchling (ASFS V9005). The egg size Grant mentioned, 9 X 7 mm,
agrees with our data for two roosevelti eggs (8.2 X 6.3-6.6 mm).
DEFINITION: A species of Sphaerodactylus with large, acute,
strongly keeled, flattened, imbricate dorsal scales axillaa to groin 19-
25, mean 21.0); no area of middorsal granules or granular scales.
Hair-bearing scale organs (one hair) only present on distal edge of
scale. Ventrals smooth, including gulars with one exception and
chest scales, rounded and imbricate axillaa to groin 19-27, mean 23.6).
Scales around midbody 35-43, mean 37.7. Internasals 0-3, mode 1;
upper labials to mid-eye 3, 2 or 4 abnormally. Dorsal scales of tail
keeled, acute, imbricate and flat-lying; scales beneath tail smooth,
rounded, enlarged midventrally. Fourth toe lamellae 9 to 12, mode
11. Escutcheon (fig. 11c) compact with virtually no extension onto
thighs (3-6 X 8-17). Color pattern of female a series of six brown
longitudinal lines on body, the medianmAwf of which extend onto
head as postocular stripes; ground coloiYght gray to tan; scapular
ocelli prominent, white and black edged, confined to paramedian
stripes; no scapular patch (fig. 4c). Pattern of males much faded,
usually only faintly evident; coloration page purplish tan to fleshy tan,
venter white. Iris golden to golden brown. Habitus stout; snout
broad and short. This is the largest of Puerto Rican Sphaerodactylus;
of 93 specimens the 52 largest vary from 33 to 39 mm snout-vent
(males 33-39, mean 36.0; females 33-38, mean 35.2).
REMARUs: This species inhabits the extreme xeric part of Puerto
Rico. It is most abundant in littoral situation~ljch as in piles of palm


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FIGURE 5./ Distribution of Sphaerodactylus. S. macrolepis on Puerto Rico (no details of subspecies shown), vertical lines; S. mac-
rolepis inigoi, crosshatching; S. macrolepis macrolepis, fine stippling; S. macrolepis parvus, coarse stippling; S. nicholsi townsendi,
open circles (questionably' recorded from Culebra); S. roosevelti, semi-solid circle; S. parthenopion, P; S. beattyi, B.







BULLETIN FLORIDA STATE MUSEUM


trash in Cocos groves, or among the dead leaves in the Coccoloba
fringe. It also occurs in scrub or xeric woods a short distance inland,
but less commonly than in the littoral situations. It was not en-
countered in the Sierra Vermeja or the xeric country west of Yauco
in the foothills of the Cordillera Central. Caja de Muertos is quite
dry and is more similar to the Guanica region than to the immediately
adjacent mainland. Therefore the distribution of S. roosevelti ap-
pears to be primarily that area south of the Valle de Lajas which
at one time was doubtless a peninsula or chain of offshore keys
(Thomas, MS).
S. roosevelti and S. macrolepis are allopatric; the record of macro-
lepis for Ensenada (Schmidt, 1928) appears to have been based on
specimens referable to roosevelti (AMNH 13811) and nicholsi (AMNH
13813). S. roosevelti does occur with nicholsi in this region, although
the latter is of wider distribution in Puerto Rico.
SPECIMENS EXAMINED: ASFS V5551, 0.5 km N Cabo Rojo lighthouse; ASFS
X4238-45, 1 mi. N Cabo Rojo lighthouse; ASFS V5562-63, 2 mi. S Pole Ojea;
ASFS X4228-30, X4251-52, 3 mi. SW Ensenada; AMNH 13811, Ensenada; ASFS
V4684-87, 6.7 km NE Guanica; ASFS V5641-42, 7.3 km E Guanica, 600 feet;
V5615-32, V5729-52, V9005, 8.5 km SE GuAnica; ASFS X4266-79, X4679-80,
9 km SE Guanica; ASFS X4288-92, 10 km SE Guanica; ASFS V5583-86, Punta
Ventana; MCZ 66451, ASFS V4752, Caja de Muertos; MCZ 36666, "Virgin
Island" (= Vieques Island?).

Sphaerodactylus klauberi Grant
Sphaerodactylus klauberi Grant, 1931, Jour. Dept. Agri. Porto Rico, vol. 15,
no. 3, p. 207, fig. 13.
TYPE LOCALITY: El Yunque, Puerto Rico.
DISTRIBUTION: Broadly speaking, the higher elevations of Puerto
Rico (fig. 7); actually about all that can be said is that it avoids the
coastal lowlands. It is an inhabitant of mesic regions from elevations
of 300 feet near Rosario to 4,000 feet and probably higher. It occurs
in the mesic parts of the Pepino Hills; the specimen from Camp
Buchanan (MCZ 34512), if the locality is correct, may be a low ele-
vation record. The species has not been taken in the Sierra de Pandu-
ras where the habitat is appropriate but where the closely related
S. gaigeae occurs.
DIAGNOSIS: A species of Sphaerodactylus having moderate to
small, acute to rounded, strongly keeled, flattened, imbricate dorsal
scales axillaa to groin 23-32); no area of middorsal granules or granular
scales. Hair-bearing organs (one hair) present on distal edge of scale.
Ventrals including throat and chest scales partly to entirely keeled,


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Vol. 10










S


*
*


0


* .


0


4"


FIGURE 6. Puerto Rico, showing ranges of S. klauberi (solid symbols), S. gaigeae (semi-solid symbols), and S. nicholsi (hollow
symbols). Range of S. klauberi stippled, shown as four discrete areas as discussed in text. S. gaigeae from south of Can6vanas
is questioned. Races of S. nicholsi; S. n. nicholsi, diagonal lines from upper left to lower right; S. n. townsendi, diagonal lines
from lower left to upper right; overlap of symbols in south-central Puerto Rico indicates sample of presumed intergrades.








BULLETIN FLORIDA STATE MUSEUM


acute, and imbricate axillaa to groin 24-34); scales around midbody
42-67; internasals 1-6, mode varies with population; upper labials to
mid-eye 3, 2 abnormally. Dorsal scales of tail acute, imbricate, and
flat-lying; scales beneath tail smooth, rounded, enlarged midventrally;
fourth toe lamellae 9-13; escutcheon (fig. 11d) large with extensions
well onto thighs (3-7 X 14-27). Color generally dark brown; body
with irregular darker and lighter spots and mottling; scapular pattern
present in most specimens; patch black, well developed or diffuse and
broken; paired ocelli white to light gray. Head pattern complex;
light brown postocular stripes extend onto neck, frequently joined by
transverse stripe which may be scalloped in center; a dark transverse
occipital spot joins laterally with dark stripes which pass posteriorly
over eyelids, leaving a light, bilobed cephalic figure; other light and
dark vermiculations may be present (fig. 7a); sacral pattern U-shaped;
throat black and gray mottled or uniform light gray; venter and un-
derside of tail typically orange or reddish pink or yellow, but may be
gray; iris orange. Habitus moderately stout; snout broad and short.
Size varies geographically (see below); the largest specimen is 37
mm snout-vent.




















A B
FIGURE 7. A, S. klauberi, anterodorsal view, combination of ASFS V4558, 30
km N, 3.1 km E Ponce, and V4946, ca. 20 km SE Cidra, Puerto Rico. B,
S. gaigeae, anterodorsal view, male, ASFS X4050, 2.5 mi. SW Yabucoa, 800 feet,
Puerto Rico.


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PUERTO RICAN SPHAERODACTYLUS


REMARKS: Rather striking geographical variation in three char-
acters, size, number of internasals, and size of dorsal scales, is dis-
cussed with reference to four geographical samples (tables 2, 8).
The lizards of the northern limestone region and the northern and
western parts of the Cordillera Central (sample I) have from 2 to 6
internasals with a mode of 3; size small to moderate; dorsal and mid-
ventral scale counts comparatively low (table 3); less than 50 percent
of the specimens have entire keeling of the ventral scales. The speci-
mens from the southern part of the Cordillera Central (sample II),
which includes some of the higher elevations on the island, are strik-
ingly larger lizards; internasals from 1 (mode) to 3; dorsal scales axilla
to groin 27 to 32; midbody scales 50 to 67; their ventral scales are all
entirely keeled. Were it not for the presence of two other samples
from farther east, sample III from Guavate, and sample IV from El
Yunque, there would be little question of the nameworthiness of the
two populations. However, the other two samples, III and IV, are in-
termediate between samples I and II in size and dorsal scale counts
(table 3); their internasals are more like those of Sample II (table 2).
No color or pattern differences are manifest between the populations.

TABLE 2. INTERNASALS IN SAMPLES OF S. klauberi

Internasals

Samples N 1 2 3 4 5 6

I 159 32 97 19 10 1
II 24 14 3 7
III 11 3 4 4
IV 19 17 2


SPECIMENS EXAMINED: I. ASFS V5306-26, 7.0 km S Mora, 800 feet; ASFS
V5220-44, 7.2 km SE Quebradillas, 900 feet; ASFS V5182-84, 1 km. S Pueblito
de Ponce; ASFS V5166-78, 1.5 km S Pueblito de Ponce; ASFS V5158-61, 2 km
S Pueblito de Ponce, 800 feet; ASFS V3228-38, 2 mi. S Pueblito de Ponce, Bosque
de Guajataca; ASFS V4530-41, 3 mi. N Lares; ASFS V5150, 9.6 km S, 1.6 km
E Mora, 800 feet; ASFS V5913, 18.8 km SW Arecibo, 800 feet; ASFS V5866,
3 mi. W Las Llanadas, 600 feet; ASFS V5985-95, 2.5 km SW Florida; ASFS
V5286-87, 1.5 km SW Rosario, 300 feet; ASFS X4253-57, 13.8 km N Sabana
Grande, 2900 feet; UMMZ 73594, Maricao; ASFS V5917, 7 km E Lares, 1200
feet; ASFS X4458-72, X4495-507, V4503-18, 17.7 km (= approx. 8 km. straight
line distance) NE Utuado, 1100 feet; ASFS V6203-26, 2 km NE Barranquitas,
2100 feet; AMNH 12988-90, 13034, 13036, 13142, 13192-94, 13196, 13198,
Aibonito.


"1966


239







BULLETIN FLORIDA STATE MUSEUM


II. ASFS V5974-76, 4.1 km NE Villa P6rez, 2200 feet; ASFS V5982, 9.1
km S Villa P6rez, 2900 feet; ASFS V6192, 10.5 km SSE Villa P6rez, 2900 feet;
ASFS V6192, 10.5 km SSE Villa P6rez, 2900 feet; ASFS V4633, 10.6 km SSE Villa
Perez, Monte Guilarte, 3400 feet; ASFS V6198-204, 1.8 km S Adjuntas; ASFS
V4557-60, 30 km N, 3.1 km E Ponce, 3500 feet 3800 feet.
III. ASFS X7573-75, Bosque de Guavate, 8 km SE Las Cruces, 2080 feet;
ASFS V4827-28, 18.8 km NNE Guayama, 2000 feet; ASFS V4839, 18.6 km NNE
Guayama, 2000 feet; ASFS V4943-46, ca. 20 km. SE Cidra; UMMZ 124805,
Bosque Guavate, nr. Prison Camp.
IV. MCZ 34473 (type), 34474-80 (paratypes), UMMZ 73595 (4 paratypes),
El Yunque; UMMZ 73596 (2), Ranger's Cabin, El Yunque; UMMZ 73598 (3),
Ranger's Cabin Luquillo Mt. (= El Yunque); ASFS X1189-90, 12 km S Palmer,
2500 feet.

TABLE 3. RANGES AND MEANS FOR THREE CHARACTERS OF
FouR GEOGRAPHICAL SAMPLES OF S. klauberi

Samples Dorsal scales Midbody scales Adult size*

N=93 N-=94 N=99
I
25.6 (23-32) 53.3 (42-55) 26.9 (26-29)
N=22 N=22 N==15
II
29.0 (27-32) 56.0 (50-67) 34.3 (33-37)
N=9 N=8 N=7
III
27.6 (23-30) 53.0 (48-57) 31.9 (31-34)
N=16 N=16 N=11
IV
26.2 (23-29) 50.4 (46-55) 31.5 (31-35)

* The upper half of the size classes (one millimeter) of the sample.

Sphaerodactylus gaigeae Grant
Sphaerodactylus gaigeae Grant, 1932, Jour. Dept. Agri. Puerto Rico, vol. 21,
no. 4, p. 508, fig. 5.
TYPE LOCALITY: Mountains near Yabucoa, Puerto Rico.
DISTIBUTION: Grant (1932a, 1932b) indicated the range of this
species as being only the "mountains between Maunabo and Yabucoa."
Since, it has been recorded from Cayo Santiago (Heatwole, Sade and
Hildreth, 1963), and we obtained a specimen on Isla Pifieros. It
appears, however, that the situation is not so simple. There is a
series of specimens (MCZ 34436, 34438-40, 34443) from 10 km S
Can6vanas, which should be on the western slopes of the Sierra de
Luquillo. This is a considerable range extension from the vicinity
of Yabucoa. According to Grant's own fieldbook he collected only
one gaigeae at 10 km S Can6vanas (in litt., Norman Hartweg to Ernest


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PUERTO RICAN SPHAERODACTYLUS


Williams). Because of the frequent and irresolvable confusion that
surrounds the Grant collections, we prefer to regard this locality for
gaigeae as questionable. The reliable localities for this species then
are the Sierra de Panduras, Cayo Santiago, and Isla Pifieros (fig. 6).
DENITION: A species of Sphaerodactylus with large, acute,
strongly keeled, flattened, imbricate dorsal scales axilla to groin 20-30,
mean 23.0; no area of middorsal granules or granular scales; hair-
bearing scale organs (one hair) present on distal edge of scale; gular
scales keeled, chest scales variably keeled, ventrals unkeeled in all
but one specimen examined, which had incomplete ventral keeling;
ventral scales acute to rounded and imbricate, axilla to groin 23-28,
mean 24.8; scales around midbody 42-52, mean 45.6; dorsal scales of
tail keeled, acute, imbricate and flat-lying; scales beneath tail smooth,
rounded, enlarged midventrally; internasals 1-4, mode 3; upper labials
to mid-eye 3; fourth toe lamellae 9-11, mode 10; escutcheon (fig. lie)
of moderate size with extensions onto thighs to behind knee (3-5 X
16-21). No sexual dichromatism; general coloring dark brown; dor-
sal pattern of darker scales or groups of scales on brown ground
color or two rows of dark-edged light (buff) ocelli which continue onto
tail as a dark stripe scalloped by light areas; scapular pattern faint
or absent, consisting of slightly darker brown patch with two small
ocelli; pattern on head and neck consists of two anteriorly pointed
light chevrons. The anterior one is derived from the light area be-
tween the dark supraciliary stripes and the nuchal dark spot of the
pattern basic to the group and the posterior one from the light anterior
margin to the scapular patch (fig. 7b). In a few specimens the chev-
rons are much expanded and continuous with the light postocular
stripe. Venter light but tends to be much invaded by dark pigment;
throat nearly uniform pale yellow or with dark brown streaking;
undersides of tails dull orange; iris golden. Habitus moderate, snout
short. Size moderate; the 15 largest specimens of 31 vary from 22
to 25 mm snout-vent, mean 22.9.
REMARKS: S. gaigeae is so closely allied to S. klauberi the two
may in reality be conspecific. S. gaigeae occupies a habitat in the
Sierra de Panduras (where klauberi does not occur) very similar to
that klauberi typically occupies elsewhere; however it also occurs in
low elevation and even xeric situations on Cayo Santiago and Isla
Pifieros. The colorations of the two are similar and are doubtless
derivative. If the 10 km S Can6vanas locality for gaigeae is correct
the two are almost certainly not conspecific, as this locality is possibly
within the range of klauberi. The relationships of the two forms must
be determined by future collecting.


1966






BULLETIN FLORIDA STATE MUSEUM


Our specimens were collected under trash, leaves, fallen palm
fronds, and tree trunks on a hillside in a moist coffee grove in the
Sierra de Panduras and under a rock on a xeric, wooded hillside on
Isla Pifieros.
SPECIMENS EXAMINED: MCZ 34436, 34438-40, 34443, 10 km S. Can6vanas;
ASFS V5053, Isla Pifieros, MCZ 62181, Cayo Santiago; ASFS X4047-56, 2.5
mi. SW Yabucoa, 800 feet; ASFS V4968, RT 1238, 6 km SW Yabucoa, 700 feet;
UMMZ 73605-06 (12 specimens), mountains between Maunabo and Yabucoa.

Sphaerodactylus nicholsi Grant
Sphaerodactylus nicholsi Grant, 1931, Jour. Dept. Agri. Porto Rico, vol. 15,
no. 3, p. 204, fig. 11. Type specimen MCZ 34578.
Sphaerodactylus townsendi Grant, 1931, Jour. Dept. Agri. Porto Rico, vol. 15,
no. 3, p. 208.
TYPE LOCALITY: Near Ensenada, Puerto Rico (3 mi. W Ensenada,
Grant field data).
DISTIBUrTIN: Peripheral Puerto Rico from Cabezas de San Juan
south and around the island and east along the north coast at least to
Playa Mar Chiquita (fig. 6). The occurrence of this form is mainly
coastal but it extends inland, especially in the dry southwest as far
as the Sierra Vermeja and Laguna Cartagena and to the area between
Yauco and Sabana Grande.
DEFINITION: A species of Sphaerodactylus with moderately sized,
acute, strongly keeled, flattened, imbricate dorsal scales axillaa to
groin 19-27); no area of middorsal granules or granular scales; hair-
bearing scale organs (one hair) only, present along free distal edge of
scales; gular scales keeled or unkeeled; chest scales unkeeled; ventrals
large, rounded, imbricate and smooth, axilla to groin 20-29; midbody
scales 36-45; internasals 1-3; upper labials to mid-eye 3; dorsal scales
of tail acute, imbricate, and flat-lying or only slightly erected; scales
beneath tail smooth, enlarged midventrally; fourth toe lamellae 8-11;
escutcheon (fig. 11f) moderate sized with extension onto thighs (4-7 X
9-22). No sexual dichromatism; coloring brown to tan with darker
brown spotting or dark-edged ocelli; light postocular stripe usually
extends along flanks onto tail. Head pattern of diffuse mottling, or a
light crescent which may meet postocular'stripes; scapular pattern re-
duced but may involve dark patch of varying intensity with two ocelli,
or two ocelli alone, or several small irregular ocelli; sacral pattern
U-shaped and dark with light edge; caudal pattern of irregular dark
mottling or ocelli; venter pale; throat pale yellow with or without
dark streaking or spotting; underside of tail yellow to pale orange.
Iris brown to golden orange. Habitus moderate, snout broad and


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242







PUERTO RICAN SPHAERODACTYLUS


short. Size small to moderate; the largest specimen measures 28
mm snout-vent, the smallest, a juvenile, 9 mm. This species includes
the smallest of Puerto Rican, but not Greater Puerto Rican, Sphaero-
dactylus.
REMARKS: This species prefers a dry habitat. In the eastern,
western, and northern portions of its range it is absent from sections
of coast where more mesic conditions prevail. The mesic areas have
only S. macrolepis, usually in some abundance; in the dryer and more
exposed regions nicholsi is abundant and macrolepis less so. Along
the southwestern littoral the two are not sympatric.
Sphaerodactylus nicholsi as here defined includes both nicholsi
and townsendi, which have heretofore been considered separate spe-
cies. The evidence for their conspecificity is not entirely conclusive;
one series may be intergradient, but others seem to indicate some
overlap. Our reasons for considering them as one species are: 1) they
show no morphological differences except in size and this is not abso-
lute, 2) their colorations are very similar, and what differences exist
are not absolute, 3) they have the same or very similar ecological pref-
erences, 4) their ranges are similar in being the peripheral parts of
the island. They are so similar in morphology and color that some
specimens of nicholsi can not be distinguished from some townsendi.
Grant (1931, 1932b) used several characters to distinguish between
nicholsi and townsendi as separate species: 1) ventral keeling (used
variously), 2) keeling and shape of snout scale's, 3) sacral pattern of
parallel lines versus Y-shaped sacral pattern, 4) scapular pattern. We
have found none of these differences consistent enough for even sub-
specific recognition. As already noted, the presence or absence of
a cephalic crescent and differences in size are the only satisfactory
characters.

Sphaerodactylus nicholsi nicholsi Grant, new combination
Sphaerodactylus nicholsi Grant, 1931, Jour. Dept. Agri. Porto Rico, vol. 15,
no. 3, p. 204. Type specimen MCZ 34578.
TYPE LOCALITY: Near Ensenada, Puerto Rico (3 mi. W Ensenada,
field data).
DISTRIBUTION: From Playa Mar Chiquita, east of Arecibo, west
and south along the coast, then east along the southern marginal re-
gion as far as Juana Diaz; range discontinuous because of interrup-
tion of habitat (fig. 6).
DIAGNOSIS: A subspecies of S. nicholsi characterized by the pres-
ence of a crescentic cephalic figure in 95 percent of the specimens


1966


243







BULLETIN FLORIDA STATE MUSEUM


(fig. 8a) and a well-defined scapular patch and paired ocelli in about
46 percent, and by small size, the snout-vent length of specimens
larger than the median fall in the size classes of 20-25 mm, aver-
age 21.7 mm.
REMARKS: The axilla to groin dorsal scale counts for this form
range from 19 to 28 with a mean of 23.7 in 109 specimens; ventral
scales axilla to groin 20 to 28, mean 24.6 in 107 specimens; midbody
scales 35 to 45, mean 39.3 in 103 specimens; fourth toe lamellae 8 to
11, mode 9. The internasals show some geographic variation; speci-
mens from the north and northwest have a modal internasal count of
2 and a high proportion with 3 (6 out of 41); those from the south and
southwest have a modal count of 1, a few with 2 (15 out of 112) and
only one with 3.















FIGURE 8. A, S. nicholsi nicholsi, anterodorsal view, female, ASFS X4286, 10 km
SE Guinica, Puerto Rico. B, S. nicholsi townsendi, anterodorsal view, female,
ASFS V5825, Esperanza, 2 mi. S Mercedita, Puerto Rico. C, S. parthenopion,
anterodorsal view, female, MCZ 77211, type, hillside above Pond Bay, Virgin
Gorda, British Virgin Islands.

Live specimens are brown dorsally with tan dorsolateral stripes and
dark brown spotting. The crescentic cephalic figure, which may
appear oval, is tan and present in all but 5 percent of the specimens,
some of which are juveniles with no pattern of any kind; scapular
ocelli, when present, white; black scapular patch single or broken
into two parts; tails brown to orange brown with some spotting or
ocellation; ventral ground color gray or flesh; throat plain gray to pale
yellow or with spots or streaks of varying intensity; underside of tail
yellow to pale orange or orange brown; iris golden orange to brown.
The specimens from 6.5 km S of Juana Diaz (ASFS V5699-712)


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PUERTO RICAN SPHAERODACTYLUS


we regard as intergrades; 5 of the 14 have no cephalic crescent, a
much higher proportion than in any sample of the subspecies nicholsi;
in the other 9 specimens the figures vary from well-developed to ob-
scure. This series could be interpreted as containing representatives
of two species, and this may be eventually found to be the case. The
specimens from Esperanza are pure townsendi in both size (they are
the largest specimens of this form) and pattern, while those from south
of VelAzquez are equally good nicholsi on both counts. These locali-
ties indicate an overlap of the two forms, yet they did not occur to-
gether at either locality. The distribution of these lizards in that cru-
cial area needs clarification by more detailed collecting. While we
regard nicholsi and townsendi as conspecific, we realize that the last
word has not been said about this problem.

SPECIMENS EXAMINED (all from Puerto Rico): ASFS V5968-69, Playa Mar
Chiquita; ASFS X7497-98, 0.7 mi. E Palmas Altas; ASFS X7520-23, 2.8 mi. W
Palmas Altas; ASFS V5891-99, 8 mi. E Arecibo; ASFS V5882-83, 4 mi. E Arecibo;
ASFS V5455-76, 1 km E Ramey Air Force Base; ASFS V5500, 9.6 km SSW
Mayagiiez; ASFS V5504-06, 10.6 km SSW Mayagiiez; ASFS V5529-32, Balneario
de Boquer6n; ASFS X4246-49, 1 mi. N Cabo Rojo lighthouse; ASFS V5552, 0.5
km N Cabo Rojo lighthouse; ASFS V5553-61, 2 mi. S Pole Ojea (on Cabo Rojo);
ASFS X4362, Laguna Cartagena, west end; ASFS V5589-93, Sierra Vermeja,
3 km SW Maguayo; ASFS X4213-27, X4250, 3 mi. SW Ensenada; AMNH 13813,
Ensenada; ASFS V5564-65, 9 mi. NW Yauco, 1000 feet; ASFS V5574-76, 8 mi.
NW Yauco, 600 feet; ASFS X4206-12, Playa Cafia Gorda; ASFS V4692-712,
6 km SE Guanica; ASFS V4688-89, 6.7 km SE Guinica; ASFS X4199-205, 8.4
km SE Guanica; ASFS V5633-39, 8.5 km SE GuAnica; ASFS X4280-84, V4683,
9 km SE GuAnica; ASFS X4286-87, 10 km SE Guanica; ASFS V5582, Punta
Ventana; ASFS V5699-712, 6.5 km S Juana Diaz (nicholsi X townsendi); ASFS
V5675-83, 12.2 km WNW, 1 km S Santa Isabel.

Sphaerodactylus nicholsi townsendi Grant, new combination

Sphaerodactyius townsendi Grant, 1931, Jour. Dept. Agri. Porto Rico, vol.
15, no. 3, p. 208. Type specimen MCZ 34613.
TYPE LOCALITY: Cape San Juan, Puerto Rico (NE corner Cape
San Juan, field data).
DIsTRIBUroN: From Cabezas de San Juan (= Cape San Juan)
south along the east coast, then west along the south coast to just
east of Ponce, localities not continuous because of incomplete collect-
ing and interruption of habitat; also Cayo Icacos, Cayo Lobos, Isla
Pifieros, Isla Vieques including Cayo de Tierra and Cayo de Afuera,
Caja de Muertos, and questionably Isla Culebra (figs. 5 and 6). Grant
never recorded townsendi from Culebra, but among a series of para-
types of S. danforthi (= S. macrolepis), many of which are untagged,


1966


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BULLETIN FLORIDA STATE MUSEUM


we found a single untagged specimen of S. n. townsendi. Whether
this specimen is truly from Culebra and had passed unnoticed these
many years, or whether it was inadvertently dropped among the Cule-
bra specimens from some other untagged series, can never be deter-
mined.
DIAGNOSIS: A subspecies of S. nicholsi characterized by the ab-
sence of a crescentic cephalic figure and a well-defined scapular patch
and paired ocelli (fig. 8b), and by large size, the snout-vent length of
specimens larger than the median of all in the size classes of 23-28
mm, average 24.6 mm.
REMARKS: The axilla to groin dorsal scale counts for this form
range from 20 to 28, with a mean of 24.4 in 137 specimens; ventral
scales axilla to groin 22 to 29, mean 25.5 in 135 specimens; midbody
scales 35 to 45, mean 39.3 in 103 specimens; fourth toe lamellae 8 to
11, mode 10; modal internasal count for this form is 1, range 1-3; 2
internasals occur with moderate frequency and 3 in only 4 of 166
specimens; no geographic variation in this count is evident.
The dorsal color of this form ranges from sandy gray to dull gray-
ish brown or yellow-brown. Irregular darker brown mottling and
spotting is present, sometimes as light centered ocelli. Light dorso-
lateral lines extending from eye to sacrum usually paler (buff) than
the ground color but are sometimes indicated more by a lack of spot-
ting and a massing of pigment along their edges than by a lighter
color; they are present in most but not all specimens, much as in
nicholsi. The head is patterned with fine vermiculations of darker
brown on the lighter ground color. Scapular ocelli, when present,
faint, irregular and asymmetrical; a dark patch may be indicated but
is usually much reduced. Only a very few specimens approach the
well-defined but faint paired ocelli and patch seen in some nicholsi;
sacral pattern evident as either a light-edged pair of black parallel
lines or as a light-edged black U; it is sometimes weak or absent.
The caudal pattern consists of diffuse darker mottlings much as in
nicholsi. The ventral coloring is typically uniformly dirty white to
gray; throat heavily marked with black streaking and mottling to un-
marked; underside of tail yellow to orange or pale orange. Iris color
was recorded as golden at Cayo Icacos or brown in intergrades south
of Juana Diaz.
The Caja de Muertos specimens present an interesting problem.
They are like townsendi in color but are very small; the larger one-
half of 27 specimens is included in the snout-vent classes of 22-23
mm, mean 22.3 mm, as opposed to other townsendi for which this
range is 23-28 mm, mean 24.3 mm. Thus in size, these specimens are


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PUERTO RICAN SPHAERODACTYLUS


closer to nicholsi. Caja de Muertos is opposite that portion of the
mainland where nicholsi and townsendi meet and presumably inter-
grade. Faunistically Caja de Muertos is more closely related to the
southwestern portion of Puerto Rico. We prefer to regard the Caja
de Muertos population as a dwarf local population of S. n. townsendi
rather than as a separate subspecies.
Sphaerodactylus n. townsendi generally inhabits somewhat less
arid situations than nicholsi, whose range encompasses the most arid
parts of Puerto Rico. It has been found in coastal palm groves, in
sea-grape litter, and in leaf litter of xeric maritime woods. On Cayo
de Tierra off Vieques specimens were collected most abundantly un-
der rocks on the exposed, almost vegetation-free, crown of the island;
the few S. macrolepis collected, and a few townsendi, were found in
rocks and leaf litter beneath a large, shady tree.
SPECIMENS EXAMINED, all from Puerto Rico except as indicated: ASFS
12507, V4743-51, V4771-76, V5838-48, Caja de Muertos; ASFS V5812-27, Es-
peranza, 2 mi. S Mercedita; ASFS V5772-89, 1.5 km S Velazquez; ASFS V4813-
25, 5 mi. W Las Mareas (south side of Bahia de Jobos); ASFS V5048-52, Isla
Pifieros; ASFS X7435-69, X7945, X8236, V35, 0.5 mi. N Las Croabas; ASFS
X1121-30, 0.6 mi. N Las Croabas; ASFS X1017-32, Cayo Icacos; MCZ 84488,
ASFS V4047-67, V6186, Isla Vieques (Cayo de Tierra); MCZ (untagged, part of
MCZ 34404-12: 57 specimens), Isla Culebra.

Sphaerodactylus parthenopion Thomas
Sphaerodactylus parthenopion Thomas, 1965, Quart. Jour. Florida Acad. Sci.,
vol. 28, no. 1, p. 118. Type specimen MCZ 77211.
TYPE LOCALITY: Hillside above Pond Bay, Virgin Gorda, British
Virgin Islands.
DISTmRBUrioN: Known only from the island of Virgin Gorda
(fig. 5).
DEFINITION: A species of Sphaerodactylus with small, acute,
strongly keeled, flattened, imbricate dorsal scales, axilla to groin 30-
35, mean 32.0; no area of middorsal granules or granular scales; dorsal
scales with hair-bearing (single hair) organs only; throat and pectoral
scales keeled in all but one specimen, ventral scales smooth, rounded,
and imbricate, axilla to groin 26-29, mean- 27.9; scales around mid-
body 50-55, mean 51.4; internasals 1 to 3, mode 2; upper labials 3 to
mid-eye, 2 unilaterally in 4 specimens; dorsal scales of tail keeled,
acute, imbricate, and flat-lying (although noted in the original descrip-
tion as standing somewhat erect from the tail, they are more flat-lying
than truly erect); scales beneath tail smooth, rounded, enlarged mid-
ventrally; fourth toe lamellae 8 (mode) or 9; escutcheon (fig. 11g)


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BULLETIN FLORIDA STATE MUSEUM


relatively small and with only slight extension onto thighs, 3-5 X
11-13. No sexual dichromatism. Coloring deep brown with a pat-
tern of scattered darker scales or a fine reticulum of darker pigment.
A narrow, dark-edged postocular stripe extends over each temple and
fades out at base of head; a dark-edged more or less oval, transverse
light (yellow-brown) bar is present on the head across the occipital
region; its ends may or may not reach the postocular stripes laterally;
a faint preocular light bar crosses the base of the snout (fig. 8c).
Ground color of tail yellowish-brown with a pattern of irregular, short,
linear or clustered dark elements; ventral color grayish to cream with
varying amounts of invasion of dark pigment; gular pattern of faint
to bold linear markings. Habitus moderate, snout moderately long.
This is the smallest species of Sphaerodactylus, the snout-vent length
of specimens larger than the median fall in the size classes of 17-18
mm, average 17.4 mm.
REMARKS: As pointed out in the original description, this species
appears most closely related to S. n. nicholsi although the differences
between the two are rather striking. The geographical interposition
of S. n. townsendi between the ranges of nicholsi and parthenopion
suggests that nicholsi and parthenopion are more primitive peripheral
members of this group of three forms and that townsendi has evolved
since the geographic continuity of nicholsi and parthenopion was
severed.
S. parthenopion inhabits rocky hillsides covered with scrubby xeric
woods and is more secretive than S. macrolepis which occurs with it.
It appears to be absent from the purely littoral localities where
macrolepis is abundant.






S45
0

0

64* 40'.
FIGURE 9. St. Croix, American Virgin Islands, showing locality records for
S. macrolepis macrolepis (open symbols) and S. beattyi (solid symbols). Races of
S. beattyi: S. b. beattyi, horizontal lines; S. b. seamani, vertical lines. Records
of S. m. macrolepis are of specimens examined, not from the literature.


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PUERTO RICAN SPHAERODACTYLUS


SPECIMENS EXAMINED (all from Virgin Gorda): ASFS V3658-59, AMNH
92821, MCZ 77212-14, ca. 0.5 mi. N Pond Bay; MCZ 77211 (type), AMNH 92822-
24, hillside above Pond Bay; ASFS V3664, Pond Bay; ASFS V3681, KU 79852-
53, between Little Dix Bay and Savana Bay; KU 79854, SW slope of Gorda
Peak, ca. 500 feet. Other than the type, all specimens listed are paratypes.

Sphaerodactylus beattyi Grant
Sphaerodactylus beattyi Grant, 1937, Jour. Dept. Agri. Puerto Rico, vol. 21,
no. 4, p. 508. Type specimen UMMZ 80567.
TYPE LOCALITY: ?Good Hope, St. Croix, American Virgin Islands
(Grant field data).
DISTRBrrTIoN: The original description of S. beattyi gives no
type locality, but states the distribution of this species as "eastern
end of St. Croix." Subsequently Grant and Beatty (1944) iterated
that beattyi occurs only east of Christiansted. Peters (1952) recorded
the type locality as "Gode Hope," St. Croix, which is correctly Good
Hope and is so recorded in Grant's field book for the type specimen of
beattyi (George Zug, in litt.). Good Hope, however, is in the south-
western corner of St. Croix. Because of Grant's repeated statements
that beattyi occurs only in the eastern part of the island, which our
own approximately similar collecting experience confirms, we think
the Good Hope locality probably is the result of a mistake in cata-
loguing. The field tags are no longer on the specimens, so it is im-
possible to determine what specimens actually pertain to what locali-
ties. The southwestern part of St. Croix is ecologically similar to the
dry and low eastern portion, so it is conceivable that beattyi does
occur there, though the evidence at present is against it. Unfortu-
nately we were not aware of this situation during our visit and made
no especial attempt to find beattyi in the southwest, though we did
collect there. The credible locality records indicate that the range
of beattyi is the eastern portion of the island, west only as far as
Rustoptwist (1.0 mi. NE Lavallee) to the northwest of Christiansted,
and on Green Cay and Buck Island.
DEFINITION: A species of Sphaerodactylus with small, acute to
rounded, weakly keeled, slightly swollen, slightly imbricate dorsal
scales axillaa to groin 34-44). A few large, hair-bearing organs (three
hairs) present on the posterior face of each dorsal scale. Ventrals
smooth to completely keeled, rounded, and imbricate, axilla to groin
26-36; gulars always keeled; pectorals smooth to keeled; scales around
midbody 57-64; internasals 0 to 3, mode 1; upper labials to mid-eye 3;
dorsal scales of tail small, acute to rounded, keeled, slightly swollen
and slightly erected; scales beneath tail smooth, rounded, enlarged


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BULLETIN FLORIDA STATE MUSEUM


midventrally; fourth toe lamellae 7 to 11, mode 10; escutcheon (fig.
llh) compact with no extension onto thighs, 4-6 X 8-12. Coloring a
uniform brown or yellowish brown (juveniles) or with a more or less
uniform reticulum of darker brown; head buff to cream with faint
middorsal and supraocular stripes or with a bold dark trilineate pat-
tern; sacral pattern a light Y with an elongate extension onto tail;
venters grayish to yellowish gray or white; throats yellow, orange,
or whitish; throats unpatterned or with light to moderate mottlings
and striation; iris golden. Habitus moderate to stout; snout moderate
in length. Size large, the snout-vent length of specimens larger than
the median fall in the size classes of 26-30 mm, average 27.2 mm.
REm~ARS: This is the most distinct species of the Greater Puerto
Rican region, which no doubt bespeaks long isolation on St. Croix.
Its restricted distribution on the island is due principally to its limi-
tation to the more arid and desert-like areas, though it may be due
in part to competition with S. macrolepis, if indeed beattyi does not
inhabit the southwestern part of St. Croix.
Grant noted in his original description that "immature, but full
grown specimens of both sexes have an elaborate pattern." Our col-
lections show the elaborate pattern is geographically restricted to
a section of the southern portion of the eastern part of the island, as
it is found both in young and adults. It is not an ontogenetic trait.
The heavily patterned specimens represent a distinct race; the un-
patterned race is the one to which the type specimen pertains and
therefore is the nominate one.

Sphaerodactylus beattui beattyi Grant, new combination
Sphaerodactylus beattyi Grant, 1937, Jour. Dept. Agri. Puerto Rico, vol. 21,
no. 4, p. 508. Type specimen UMMZ 80567.
TYPE LOCALITY: ?Good Hope, St. Croix, American Virgin Islands.
DisTRIBUTroN: The eastern part of St. Croix east of Christiansted,
except for a segment of the southeastern coast; northwest of Christian-
sted at least as far as Rustoptwist; questionably to the southwest as
far as Good Hope; also Green Cay and Buck Island (fig. 9).
DIAGNOSIs: A subspecies of Sphaerodactylus beattyi characterized
by a very faint head pattern of indistinct middorsal and supraciliary
stripes which appear almost unicolor, and a generally subdued dorsal
body pattern (fig. 10a).
REMARKS: The two races show no apparent scalation differences.
The axilla to groin dorsal scale counts for 22 specimens of this form
range from 36 to 44, mean 41.9; ventral scales axilla to groin 29 to 35,


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PUERTO RICAN SPHAERODACTYLUS


mean 32.1; midbody scales 58 to 65, mean 61.1; 54 percent of the speci-
mens of beattyi have keeled ventral scales against only 20 percent in
the other race. No geographic variation is apparent in this species
except in color. The color of this race is relatively drab. The ceph-
alic pattern is present but so slightly darker than the ground color
that the head appears almost unicolor. A dorsal body pattern of a
brown reticulum is present in most specimens and varies from mod-
erately prominent to faint. Throat pattern absent (mode) or very
faint. A series of 17 specimens from "east end of St. Croix" (UMMZ
80783) contains both heavily patterned and unpatterned specimens.
These may represent intergrades or they may be the result of pooling
of localities.
SPECIMENS EXAMINED: St. Croix: ASFS V3600-14, 1.0 mi. NE Lavallee;
ASFS V3246-49, 5 mi. E Christiansted; ASFS V3251-56, 7 mi. E Christiansted;
ASFS V3568-91, 1.0 mi. W of eastern tip of island; ASFS V3561-65, ca. 0.75 mi.
W of eastern tip of island; UMMZ 80783 (17 specimens, intergrades?), east end
of St. Croix; ASFS V3327-51, Milord Point (west side of Great Pond Bay); UMMZ
80567 holotypee), ?"Good Hope."






i.:








A B



FIGURE 10. A, S. beattyi beattyi, anterodorsal view, female, ASFS V3564, ca.
0.75 mi. W eastern tip of island, St. Croix, American Virgin Islands. B, S.
beattyi seamani, anterodorsal view, male, MCZ 81056, type, ca. 0.5 mi. E Mt.
Fancy, St. Croix, American Virgin Islands.


1966







BULLETIN FLORIDA STATE MUSEUM


Sphaerodactylus beattyi seamani new subspecies

TYPE: MCZ 81056, an adult male from ca. 0.5 mi. east of Mt.
Fancy, St. Croix, one of a series collected on 7 August 1965 by David
C. Leber and Richard Thomas. Original number V3502.
PARATYPES: AMNH 94204-09, ASFS V3498-501, V3540-43, KU 79917-23,
MCZ 81057-62, UF 21277-82, UIMNH 56945-50, USNM 152599-600, same data
as type; ASFS V3592-99, 2 mi. W Grapetree Bay, St. Croix, 8 August 1965, R.
Thomas.
ASSOCIATED SPECIMENS: UMMZ 80568-71 (paratypes of S. beattyi), east
end of St. Croix.
DISTRIBUTION: Known from ca. 0.5 mi. E of Mt. Fancy to 2 mi.
W of Grapetree Bay along the southeastern coast of St. Croix. In-
tergradation apparently takes place at the eastern portion of the
island, possibly through the discontinuity in the hills between Grape-
tree Bay and Tague Bay (fig. 9). Intergradation to the west must
occur between Mt. Fancy and Milord Point where a large series of
S. b. beattyi was collected.
DIAGNOSIS: A subspecies of Sphaerodactylus beattyi characterized
by a bold, dark head pattern and by somewhat more prominent dor-
sal body and throat patterns (fig. 10b).
DESCRIPTION OF TYPE: An adult male with a snout-vent length
of 28 mm, incomplete tail 23 mm. Dorsal scales between axilla and
groin 40, ventral scales axilla to groin 32; scales around midbody 63;
fourth toe lamellae 9; internasal 1; supralabials to mid-eye 3; gular
scales keeled; pectoral scales partly keeled; ventrals smooth; escutch-
eon compact, not extending onto thighs, size 5 X 8; dorsal body color
uniform tan with a very faint darker reticulum on the lower sides;
ground color of head light gray; three heavy brown stripes begin on
snout and proceed posteriorly onto top of head and fade out on neck;
heavy brown mottling on sides of head forms a pattern of irregular
lines radiating from eye; heavy but faint markings on neck fade out an-
terior to scapular region, laterally they are continuous with reticulum
on lower sides; sacral pattern a pale Y with stem extending the length
of the tail; throat flecked and mottled with dark brown; ground color
of throat in life blue-gray; venter light gray; underside of tail yellow
to orange; iris golden.
VAIATION: In dorsal scale counts axilla to groin the range in
22 specimens is 37 to 44, mean 39.3; ventral scales axilla to groin 26
to 35, mean 30.9, midbody scales 57 to 64, mean 61.5. Other characters
vary much as recorded for the species. As noted under the nominate
race, the incidence of ventral scale keeling in seamani is only about


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PUERTO RICAN SPHAERODACTYLUS


20 percent. The dorsa of the paratypes vary from almost unicolor
like the type to a pattern of bold brown mottling or reticulations. The
cephalic pattern is very much like that described for the type in all
specimens except two from 2 mi. W of Grapetree Bay that resemble
the nominate race and may indicate a trend toward intergradation
to the east. In boldly marked specimens the tail is also marked and
the stem of the sacral Y is heavily outlined with dark pigment. The
throats are much more heavily marked in most specimens of seamani
than in beattyi, particularly in adults; ventral coloring is much like


that described for the type except that a few
stippling on the venter.


D E


specimens have a dark


F


0AK

G

H I


FIGURE 11. Escutcheons of Greater Puerto Rican Sphaerodactylus, as follows:
A, S. macrolepis phoberus, ASFS X909; B, S. monensis, MCZ 34568; C, S. roose-
velti, ASFS X4288; D, S. klauberi, ASFS V4507; E, S. gaigeae, ASFS X4050; F,
S. nicholsi nicholsi, ASFS X4281; G, S. parthenopion, ASFS V3681; H, S. beattyi
beattyi, ASFS V3574.
DISCUSSION
We feel that, with the probable exception of S. beattyi, the sphae-
rodactyls of Greater Puerto Rico represent a single radiation. Evi-
dence in support of this is the similarity of all forms concerned in:


A


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B C







BULLETIN FLORIDA STATE MUSEUM


1. Scalation: Dorsal scales acute, keeled, flattened, no zone of
middorsal granules, scales of tail flat-lying.
2. Scale organs: Hair-bearing organs are similar in kind and
disposition.
3. Scapular pattern: The presence of a scapular patch and ocelli,
though variously modified.
4. Head pattern: The cephalic patterns of the Puerto Rican
Sphaerodactylus appear to be derived from a basic pattern composed
of the following elements: a) supraocular stripes that join on the
snout, pass over the eyelids, and extend posteriorly to encircle or fuse
with the nuchal spot, b) a dark nuchal spot which may be alone or
included as a darker center to a larger dark area, c) postocular light
stripes between the posterior portion of the supraocular stripe and
the postocular stripe; d) postocular dark stripe, which may be con-
tinuous with the dorsolateral body stripes.
This basic pattern is variously modified. In some forms (nicholsi,
parthenopion, roosevelti) certain elements seem to have been lost,
while in others (klauberi, gaigeae) the anterior light edge of the scap-
ular pattern has been added to produce an even more complex ceph-
alic pattern. In some forms the dark areas of the pattern appear to
have coalesced leaving as a cephalic pattern only parts of the original
light ground color (cf. nicholsi, parthenopion, klauberi, gaigeae).
Sphaerodactylus m. macrolepis of the Virgin Islands and the Puerto
Rican subspecies of macrolepis taken together are widespread geo-
graphically. They are less restricted in habitat and their color pat-
terns are less modified than any other Sphaerodactylus in this area.
For these reasons we consider these populations as being the most
primitive of the living geckos of this genus in Greater Puerto Rico.
The presence of knob-like scale organs both in the Puerto Rican
subspecies of macrolepis and macrolepis inigoi, which is in essence
also Puerto Rican, is perhaps a key to the evolutionary history of this
genus in the area. As we are convinced of the relationship of the
Greater Puerto Rican forms, this can be interpreted in two ways:
1. The presence of knob-like organs is primitive. Subsequent
radiation in this region involved loss of these organs in almost every
case. On this basis it is assumed that on the prehistoric continuous
land mass of Great Puerto Rico, differentiation between m. macrolepis
and the Puerto Rican subspecies of macrolepis began with the sepa-
ration of the Virgin Islands from the larger land mass. This would
account for the intermediate nature of the Vieques race inigoi, had it


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PUERTO RICAN SPHAERODACTYLUS


originally been intergradient but closer to the Puerto Rican races.
After the separation of Vieques from Puerto Rico inigoi diverged
even more from its parent stock. In the occurrence of S. n. townsendi
and S. roosevelti, Vieques is faunistically closer to Puerto Rico.
2. The presence of knob-like scale organs is advanced. They
were developed as a racial character of the Puerto Rican-Vieques
populations either prior to, or after, the separation of the Virgin
Islands from the Puerto Rican landmass; again the Vieques popula-
tion would have been intergradient before the separation. Mona
would also have to have been colonized by Sphaerodactylus (or sep-
arated from Puerto Rico) prior to the acquisition of knob-like organs
by the Puerto Rico-Vieques population.
As a number of forms scattered throughout the West Indies have
knob-like scale organs, the 'second alternative is less satisfying to us
because it would mean that knob-like organs had arisen on separate
occasions. It is easier to imagine that these organs have been lost
by the derivative Greater Puerto Rican forms than that they developed
independently in different forms.
Yet another alternative is that the Greater Puerto Rican sphaero-
dactyls are the result of two separate invasions, an older one that
resulted in the species roosevelti, klauberi, gaigeae, nicholsi, partheno-
pion, and monensis without knob-like organs, and a relatively re-
cent one resulting in the widespread, relatively poorly differenti-
ated S. macrolepis, in which the knob-like organs have only been lost
once in the Virgin Islands form, unless monensis be included in
this invasion. The overall similarity of these species leads us to re-
ject this possibility.
Based on alternative number one above, we may summarize the
hypothetical invasion and radiation of Greater Puerto Rican Sphaero-
dactylus (fig. 12):
1. Invasion of the more or less continuous Greater Puerto Rican
area by a proto-macrolepis species of Sphaerodactylus.
2. Diversification of this species into the several derivative forms:
a. S. roosevelti as an allopatric isolate, in the southwest, prob-
ably on a peninsula or chain of offshore keys.
b. Derivation of the nicholsi-parthenopion progenitor as an
ecologically specialized form in the arid littoral.
c. Derivation of the klauberi-gaigeae progenitor as an eco-
logically specialized form in mesic forest.
d. Isolation of the monensis progenitor on Mona either by


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255







BULLETIN FLORIDA STATE MUSEUM


geological separation of Mona from Puerto Rico or by dispersal across
the water gap.
3. Continued evolution towards the present day macrolepis.


4. Invasion of the northern Leeward
subsequent differentiation.


Islands by macrolepis and


5. Invasion over water to St. Croix by macrolepis.


MONENSIS





PROTO-





ROOSEVELTI


FIGuRE 12. Diagram of hypothetical evolutionary relationships
Rican Sphaerodactylus, with the exception of S. beattyi.


KLAUBERI

GAIGEAE









/ NICHOLSI

SPARTHENOPION
of Greater Puerto


The above events are not intended to be in exact sequence; some
of them would have to have taken place more or less synchronously.
We presume that S. roosevelti arose allopatrically by differentiation
in the southwestern region, which may then have been a peninsula or
chain of offshore islands. The fact that roosevelti occurs on Caja de
Muertos and presumably Vieques suggests that its range was prob-
ably more extensive. The Vieques population might have arisen
from waif dispersal; Caja de Muertos has other southwestern en-
demics and was certainly part of this region.
S. klauberi and its relative gaigeae present a different picture. We
cannot imagine that they arose by allopatric isolation in the usual
sense of the word, for they are associated with no easily defined
area. We postulate that they differentiated altitudinally and later,
possibly correlated with the reduction of extremes in elevation, be-
came more widespread as forest species and thereby partially sym-
patric with macrolepis. In support of this it may be noted that


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PUERTO RICAN SPHAERODACTYLUS


klauberi today reaches elevations some 1000 feet higher than are
known for macrolepis, so the two are at least in part allopatric now.
The present distribution of nicholsi and parthenopion indicates
that their progenitor was rather widespread when the Virgins became
separated from Puerto Rico. There is little to indicate what might
have been the isolating factor in the evolution of this group, at least
geographically; perhaps it was an endemic of a southern marginal
xeric region.
On Puerto Rico itself the evolution of S. macrolepis has resulted
in the formation of six subspecies; stibarus from Isla Pifieros need not
concern us further as it is obviously derived from grandisquamis of
the adjacent mainland. Of the six races ateles and guarionex seem
most closely related, and with them should also be placed the upland
spanius. Ateles inhabits the xeric southwest exclusive of that region
occupied by S. roosevelti, and guarionex generally the more mesic
north coast and adjacent Pepino Hills. Ateles and guarionex are not
known to intergrade, although such intergradation may well take
place. Spanius is presumably a derivative of the ateles-guarionex
pair, occupying the higher interior of the island. In the east grandi-
squamis intergrades with phoberus, and the latter apparently is a
local derivative of the former, presumably restricted to the peninsula
upon which the city of San Juan presently stands. Grandisquamis
intergrades with mimetes in the southeast and mimetes is known to
intergrade with spanius at Aibonito. Doubtless intergradation be-
tween all or most adjacent races of S. macrolepis on Puerto Rico will
ultimately be demonstrated, so that present absence of intergradation
should not be weighted unduly.
In summary there appear to have been three major areas of
evolution of the proto-macrolepis stock in Puerto Rico and Vieques:
1) inigoi-mimetes on Vieques and southeastern Puerto Rico; 2) ateles-
guarionex-spanius in the southwest, on the north coast east to the
vicinity of Gurabo, and in the interior uplands; and 3) phoberus-
grandisquamis-stibarus in northeastern and eastern Puerto Rico and
on Isla Pifieros.
Of the Puerto Rican races only phoberus is not known to occur
with any other sphaerodactyls. Of the remaining races, grandi-
squamis, mimetes, and inigoi occur sympatrically both grossly and
in detail with S. n. townsendi. The races spanius and guarionex
occur with klauberi, and the latter as well with S. n. nicholsi. S.
gaigeae occurs with stibarus on Isla Pifieros, with mimetes X grandi-
squamis intergrades in the Sierra de Panduras, and with grandi-
squamis on Cayo Santiago. Finally, ateles occurs with S. n. nicholsi.


1966


257







BULLETIN FLORIDA STATE MUSEUM


From the above, it is easily observed that the major companion of
S. macrolepis in Puerto Rico is S. nicholsi, except for the races guar-
ionex and spanius that occur in uplands or forested situations, which
are occupied by S. klauberi. In those localities where races of S.
macrolepis occur with S. klauberi, the latter is invariably the more
common of the two. Syntopic (see Rivas, 1964, for definition) occur-
rence of the Greater Puerto Rican species of Sphaerodactylus is dia-
grammed in table 4; positive syntopy does not necessarily indicate
extensive sympatry.

TABLE 4. CHART SHOWING SYNTOPIC OCCURRENCE OF THE SPHAERODACTYLS
OF THE GREATER PUERTO RICAN REGION*

0





macrolepis
roosevelti -
klauberi + -
gaigeae + ?
nicholsi' + + +
parthenopion + -
beattyi + -

*Positive syntopy does not necessarily indicate extensive sympatry.

S. beattyi, like the other St. Croix endemics, is not closely re-
lated to other Greater Puerto Rican forms. It is difficult to de-
termine whether this distinctness is due to long isolation, St. Croix
having been longer isolated than the other Virgin Islands, or to col-
onization from another direction, or both. S. beattyi is peculiar in
possessing large scale organs bearing three hairs, very small but keeled
and imbricate dorsal scales, and a compact escutcheon. It does not
appear to be related to any of the Greater Puerto Rican species, nor
for that matter to those of the Lesser Antilles. King (1962) regarded
numerous hairs in large scale organs as primitive. Three hairs ap-
pear to be the maximum for West Indian species (King, 1932; per-
sonal observation). Some of the other characters of S. beattyi also
appear to be primitive. We feel that beattyi is a relict species long
isolated on St. Croix and not closely related to surrounding species.
S. macrolepis is doubtless of relatively recent advent on St. Croix,


258


Vol. 10







PUERTO RICAN SPHAERODACTYLUS


though there is no reason to suppose that it has been introduced,
as Grant and Beatty (1944) postulated. The suggestion by these
authors that macrolepis might be crowding beattyi out is a distinct
possibility. The two are in part allopatric on St. Croix, and if beattyi
has occupied the island a much longer time, there is no evident reason
why it should not have occupied all of the island when it was alone
there. It may be that with the occurrence of two species each has
fallen heir to the habitat which is most suitable, although, according
to the data of Grant and Beatty, macrolepis has invaded more of
beattyi's domain than the reverse.

LITERATURE CITED
Barbour, Thomas
1921. Sphaerodactylus. Mem. Mus. Comp. Zool., vol. 47, pp. 217-278, 26 pls.
Grant, Chapman
1931. The sphaerodactyls of Porto Rico, Culebra and Mona islands. Jour.
Dept. Agri. Porto Rico, vol. 15, no. 3, pp. 199-213, 5 pls.
1932a. A new sphaerodactyl from Porto Rico. Jour. Dept. Agri. Puerto Rico,
vol. 16, no. 1, p. 31.
1932b. Chart for determining the sphaerodactyls of the Puerto Rico region.
Jour. Dept. Agri. Puerto Rico, vol. 16, no. 1, pp. 33-35, + chart.
1932c. The herpetology of Vieques Island. Jour. Dept. Agri. Puerto Rico, vol.
16, no. 1, pp. 37-39.
1932d. Sphaerodactylus grandisquamis, a valid species. Jour. Dept. Agri.
Puerto Rico, vol. 16, no. 1, pp. 43-45.
1937. Herpetological notes with new species from the American and British
Virgin Islands, 1936. Jour. Dept. Agri. Puerto Rico, vol. 21, no. 4, pp.
503-522, 4 pls.
and Harry A. Beatty
1944. Herpetological notes on St. Croix, Virgin Islands. Herpetologica, vol.
2, pp. 110-113, 1 fig.
Heatwole, Harold
1961. Institute of Caribbean Studies Field Excursion to Isla Mona. Herpe-
tology. Caribbean Jour. Sci., vol. 1, no. 1, pp. 32-33.
,Donald S. Sade, and Richard Hildreth
1963. Herpetogeography of Puerto Rico. I. Herpetofauna of Cayo Santiago
and Cayo Batata. Caribbean Jour. Sci., vol. 3, no. 1, pp. 1-5, 2 figs.
King, Wayne
1,989. Sy~JiAsAr a Ldxee- AnmY All-,rr I,'-arrcuk ofr t~e genus SpAaeroa'act'y'us.
Bull. Florida State Mus., Biol. Sci., vol. 7, no. 1, pp. 1-52, 17 figs.
Peters, James A.
1952. Catalogue of type specimens in the herpetological collections of the
University of Michigan Museum of Zoology. Occ. Papers Mus. Zool.
Univ. Mich., pp. 1-55.


1966


259








260 BULLETIN FLORIDA STATE MUSEUM Vol. 10

Rivas, Luis Rene
1964. A reinterpretation of the concepts "sympatric" and "allopatric" with
proposal of the additional terms "syntopic" and "allotopic". Systematic
Zool., vol. 13, pt. 1, pp. 42-43.
Rolle, Francis J., Harold Heatwole, Richard Levins, and Frank Torres
1964. Faunal notes on Monito Island, Puerto Rico. Caribbean Jour. Sci., vol.
4, no. 1, pp. 321-322.
Schmidt, Karl P.
1928. Amphibians and land reptiles of Porto Rico, with a list of those reported
from the Virgin Islands. Sci. Surv. Porto Rico and the Virgin Islands,
vol. 10, pt. 1, pp. 1-160, 4 pls., 52 figs.
Stejneger, Leonhard
1904. The herpetology of Porto Rico. Rept. U. S. Nat. Mus., 1902, pp. 549-
724, 1 p1., 197 figs.
Thomas, Richard
MS. Additional notes on the amphisbaenids of Greater Puerto Rico. Breviora,
in press.





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