BULLETIN OF THE FLORIDA STATE MUSEUM BIOLOGICAL SCIENCES Volume 10 Number 6 SPHAERODACTYLUS (GEKKONIDAE) IN THE GREATER PUERTO RICO REGION Richard Thomas and Albert Schwartz 4 Of 0 1 '...a , I I4 -i .. /853 UNIVERSITY OF FLORIDA Gainesville 1966 Numbers of'the. BULLETIN OF THE FL0RIDA STA,TE MUSEUM ara pub- lished at irregular intdvalk. Volumes 66ntain abdut 500 balds and .ate not neq- 65*arily completed, in 'any dne cabndif *4. WALTER AUFFENBERG, Mdnaging Editor OLIVER L. AUSTIN, JR., Editor Consultants for this issue. WALTER AUFFENBERG AND ERNEST E. 'WILLIAMS Communications concerning purchase or exchange of the publicat{65 and 411 manuscripts should be addressed to the Managing Editor of the Bulletin, Florida State Museum, Seagle Building, Gainesville, F16rida. 82601 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 Gunther 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. macrotepis Gunther. A new subspecies of S. beattyi G~ant 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 history of the macrolepis complex is presented. TABLE OF CONTENTS Introduction 194 Acknowledgements and Abbreviations 195 Systematic List 196 Sphaerodactylus macrolepls 196 Sphaerodactt>lus monensis 231 Sphaerodactylus foosevetti 288 Sphaerodactulus klauberi 236 Sphaerodactulus gaigeae 240 Sphaerodactylus nicho?si 242 Sphaerodactulus parthenopion 247 Sphaerodactvlus beattwi 248 Discussion 258 Literature Cited 259 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 Letocephalus cubansis 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. Sphaerodact!/tus (Gekkonidae) in the Greater Puerto Rico Region. Bull. Florida State Mus., vol. 10, no. 6, pp. 198-260. 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 parous 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 Gunther, 1859 monensis Meerwarth, 1901 grandisquamis Stejneger, 1904 roosevelti Grant, 1981 nicholsi Grant, 1931 danforthi Grant, 1981 klauberi Grant, 1981 townsendi Grant, 1981 gaigeae Grant, 1932 beatti/i Grant, 1987 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 presehee ·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. 1966 PUERTO RICAN SPHAERODACTYLUS 195 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 zised and in methods of taking counts. In general we use his style of diagnosis and de- scriplion, 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 (UP'). Felix Ifiigo of the Department of Agriculture and Commerce of the Commonwealth of Puerto Rico has repeatedly given prompt and courteous service in obtaining omcial 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 196 BULLETIN FLORIDA STATE MUSEUM Vol. 10 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 Beld: 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 Gunther Sphaerodactylus macrolepis Gunther, 1859, Ann. Mag. Nat, Hist. (3), Vol. 4, p. 215. TYPE LOCALITY: St. Croix, American Virgin Islands. DISTRIBUTION: 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. Barth61emy. 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 a]nd 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. Th6mas and its satellites north and east to Anegada; the northernmost. Leeward Islands, except Sombrero, 1966 PUERTO RICAN SPHAERODACTYLUS 197 south as far as St. Barth61emy). Dorsal scales of tail keeled, acute, imbricate, and flat-lying; ventrai 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 20-82, usually smooth, but occasionally keeled on the anteri6r part of the venter and rarely on the entire venter; scales around midbody 81 to 54; internasals 0 to.3 (mode 1); upper labials to mid-eye, 8 (some times 4); escutcheon (fig. 1la) with a broad and compact central area and extensions onto the thighs to near the underside of the knee (840 X 11-80). 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 611 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«tent 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: 608) described Sphaerodactylus grandisquamis he differentiated it from S. macrolepis Gunther 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 (1982d) he distinguished be- tween S. macrolepis and S. grandisquamis and called each a separate species. 198 BULLETIN FLORIDA STATE MUSEUM Vol. 10 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 di#icilis 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 (1982: 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. Barth61emy. 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 forrns. 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 67' ' 66· 9% /1- /4 -a -11* -- -1111~~1'j'~1~t -Villjlf#Ii/iidi# 4 .... . . . . . . . . . . . . . . . . . . . . . . . 9 5 10 IP 6 7 ' *y 66' MI . 1 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. foosevelti, fine stippling. 200 BULLETIN FLORIDA STATE MUSEUM Vol. 10 of the examination of 689 specimens of S. macrolepis from Puerto Rico six subspecies are recognizable; another race is confined to Isla Pifieros. Sphaerodactylus macrolepis grandisquamis Stejneger Sphaerodact!/tus g,andisquamis 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 88 mm, females to 31 mm snout- vent length); high number of midbody scales (86 to 46, mean 41.2 + .48 - One standard error of mean); escutcheon 5-9 X 23-80. 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 6range; chin and throat yellow to orange with some pale brown to rusty fiecks; 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 Recks. 1966 PUERT6 RICAN SPHAERODACTYLUS 201 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 88 mm and females 81 mm. The discrepancy is doubtless due to the smaller series of females. Scale counts are: dorsal scales axilla to groin 17-28, 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; intemasals 0-3, mode 1; fourth toe lamellae 8-12, mode 12. All specimens have 8 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 sn6ut-vent length of 24 mm) agrees with S. m. grandisquamis both in pattern and scale counts. SPECIMENS EXAMINED (all from Puerto Rico): UMMZ 78587, 78589, Rio Piedras; UMMZ 78615 (8 specimens), Trujillo Alto; UMMZ 124804 (4 specimens), Campo Rico; UMMZ 78588, 78622, 10 km S Can6vanas; ASFS V4997-98, Lu- quillo (outskirts); ASFS V5001-10, V6002, V6211, 1.5 km SW Luquillo; MCZ 84519-28, Cabezas de San Juan; ASFS X7470-78, 0.5 mi. N Las Croabas; ASFS V5090-108, 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 18 February 1965 by Albert Schwartz and Richard Thomas. Original number V5080. 202 BULLETIN FLORIDA STATE MUSEUM Vol. 10 PARATYPES: AMNH 94177-78, ASFS V5020-22, V5081-84, V5044-47, KU 79887-90, UF 21271-76, UIMNH 56915-18, same data as type. DISTRIBUTION: Known only from Isla Piaeros, 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 (86 to 41, mean 88.6 + .87); 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. Dersal scales axilla t6 groin 18; ventral scales axilla to groin, -; midbody scales 88; fourth toe lamellae 11; in- ternasal 1; 3 bupralabials 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 backhas 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 80 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 86-41, mean 38.6; escutcheon 5-8 X 18-24; inter- nasals 043, mode 1; fourth toe lamellae, 9-ll, 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; 1966 PUERTO RICAN SPHAERODACTYLUS 208 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 28 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. grandisquainis 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, 88.6 in the latter) are statistically different. The breadth of the es- cutcheon in grandisquamis varies from 28 to 80 scales, that of stibarus from 18 to 24. Other features that help differentiate the two races are the less censistent 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, t:he 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 204 BULLETIN FLORIDA STATE MUSEUM Vol. 10 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 Sfandard error Subspecies N and extremes deviation of mean (x2) grandisquamis 28 41.2 (36-46) 2.29 .86 stibards 16 88,6 (86-41) 1.49 .74 phoberus 89 37.5 (38-41) 2.62 .84 mimetes 29 · 86.1 (31-40) 2.10 .78 ateles 26 89 .4 (86-42 ) 1 .78 .70 spanius 26 43.6 (41.47) 1.68 .66 guarionex 92 41.3 (86-49) 2.60 .54 inigoi 88 86.8 (38-41) 1.84 .64 macrolepis 86 48.1 (88-50) 2.46 .54 parcus 14 48.4 (44-54) 2.87 1.54 Sphaerodactylus macrolepis phoberus, new subspecies TypE: MCZ 81028, 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 Call from Puerto Rico): AMNH 94179-84, KU 79891-94, MCZ 81024-85, UF 21242-51, UIMNH 56919-24, USNM 152593-96, same locality as type, 18 June 1962, R. F. Klinikowski, D. C. Leber, A. Schwartz; ASFS X899-918, X920-86, same data as tykle; ASFS X4184.58, 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, X2029-80, X2084, X2240, X2246«47, X2264, X2829-81, X2865-66, X2898, X3135-87, X8267, X3813, X8551-52, X3860, X4019-20, X4044-46, all hatchlings from eggs taken at the type locality; ASFS X4687-48, 2.4 mi. W Loiza Aldea; UMMZ 124811, Punta Vacia Talega (see below). DISTRIBUTION: 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 Sl mm); moderate number of midbody scales (88-41, mean 37.5 + .42); males usually dark wood brown, unicolor or with scattered dark brown 1966 PUERTO RICAN SPHAERODACTYLUS 205 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 89; fourth toe lamellae 12; inter- nasal 1; 8 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 drregularly 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. 9 1,4 't, 18 '11~/4.;;: Mlif ~~2; ' *marvj fs 16-71 )~, ),ij~~i,59;,s~i,y# C~t,f~ A k 1.-- , FIGURE 2. A, S. macrolepis grandisquamif, anterodorsal view, female, ASFS X7470, 0.5 mi. N Las Croabas, Puerto Rico. B, S. macrolepis phoberus, antero- dorsal view, female, MCZ 81628, type, Isla Verde, Puerto Rico. C, S. macro- tepis mimetes, anterodorsal view, male, MCZ 81086, type, 12.8 km SE Patillas, Puerto Rico. Variation: 20 males and 20 females have maximum snout-vent lengths of 81 mm in each sex and the following scale counts: dorsal scales axilla to groin 17-22, mean 19.6; ventral scales axilla to groin 28-81, mean 26.4; midbody scales 38-41, mean 87.5; escutcheon 6-10 X 24-28; internasals (data from 144 individuals) 0-8, mode 1; fourth toe 206 BULLETIN FLORIDA STATE MUSEUM Vol. 10 lamellae 9-12, mode 10. All 40 specimens examined in detail have 8 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 pQstorbital 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- faee 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 sh6ws 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 obscuredby 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. 1966 PUERTO RICAN SPHAERODACTYLUS 207 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 eounts, 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 X4687-48) 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 18 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- upus woods near the coast that appear quite suitable for S. macro- lepis, but intensive *earch has revealed no sphaerodactyls in this im- mediate area. The same situation applies elsewhere in Puerto Rico, bringing to mind Grant's statement (1981: 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. 208 BULLETIN FLORIDA STATE MUSEUM Vol 10 Sphaerodactylus macrolepis mimetes, new subspecies TypE: MCZ 81086, an adult male from 12.8 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 56925-80, same data as type; AMNH 94186, 10 km SE Patillas, 1 February 1965, R. Thomas; AMNH 94185, 5 mi. W Las Mareas (south side Babia de Jobos), 80 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, ZOO feet, 9 March 1965, R. Thomas; UF 21252-5S, KU 79895-96, Bafios de Coamo, 9 March 1965, A. Schwartz, R. Thomas; MCZ 34446-49, Bafios de Coamo, 1981-32, 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 78592, 12814, Bafios de Coamo; UMMZ 78586 (8 specimens), 78620, Arroyo; MCZ 84444-45, Punta Arroyo, UMMZ 78591 (4 specimens), beach near Maunabo. S. m. mimetes X grandisquamis: MCZ 84505-10, mountai~ between Maunabo and Yabucoa; ASFS X4057-58, 2.5 mi. SW Yabucoa, 800 feet. DISTRIBUTION: 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 82 mm, females to 30 mm), low number of midbody scales (81-40, mean 36.1 + .89); 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 88; 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: 88 Spe6imenS of both sexes have maximum snout- vent lengths of 82 mm in males, 80 mm in females, and the following 1966 PUERTO RICAN SPHAERODACTYLUS 209 scale counts: dorsal scales axilla to groin 16-21, mean 18.1; ventral scales axilla to groin 20-29, mean 24.1; midbody scales 81-40, mean 86.1, escutcheon 4-5 X 21-24; internasals 0-2, mode 1; fourth toe lamellae 8-12, mode 10. All specimens have 8 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 paratypie males are much like the type in pattern and color. The heads are always boldly marked, and the basic female pat- tem 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, 8 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 juyeniles are unspotted. 210 BULLETIN FLORIDA STATE MUSEUM Vol. 10 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 mountaiia, 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 84506) 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 thfoats. 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, Veldzquez, 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 81048, 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 Mayaguez, 26 February 1965, A. Schwartz, R. Thomas; AMNH 94191-94, ASFS V5515-17, V5528-28), same data as type; UF 21256, 9 mi. NW Yauco, 1000 feet, 27 Febru- ary 1965, R. Thomas; UIMNH 56981-85, 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 lan SW San Germdn, 28 February 1965, A. Schwartz. 1966 PUERTO RICAN SPHAERODACTYLUS 211 ASSOCIATED SPECIMENS: MCZ 84460-62, 5 mi. S Playa de Mayaguez; UMMZ 73627, Playa de Ponce; ASFS V6629-SO, V6641, 4.6 km W, thence 4.6 km NW Juana Diaz. DISTRIBUTION: Southwestern Puerto Rico from the vicinity of Mayaguez 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 88 mm, moderate number of midbody scales, 86-42, mean 39:4 + .85. 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 82 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; 8 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. Sa); 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 83 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 86-42, mean 89.4; escutcheon 4-8 X 21-27; internasals 0-8, 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 eastennost 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. 212 BULLETIN FLORIDA STATE MUSEUM Val. 10 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 Recks form- ing a diffuse and narrow collar in sdme specimens; ventral coloration grayish to flesh-gray. 4 r e F re *V A 4/ ' 7 FIGURE 3. A, S. macrolepis ateles, anterod6rsal view, female, ASFS V5525, para- type, Balneario de Boquer6n, Puerto Rico. B, S. macrolepis guarionex, antero- dorsal view, female, MCZ 81048, type, Offcers' 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 Mayaguez (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 1966 PUERTO RICAN SPHAERODACTYLUS 218 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 m 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 86.1) of mimetes is significantly different from the higher number (mean 89.4) 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 grandisquainis but not from stibarus (table 1). Juveniles of ateles are lined in contrast to the unicolor juveniles of the northeastern races. REMARKS: S. ln. 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 78627) 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 Mayaguez, and inland under rocks on exposed hillsides on the southern slopes of the Cordillera Central above Yauco and Susda. In. this same region, sev- eral specimens were taken in a wet and shaded ravine. 214 BULLETIN FLORIDA STATE MUSEUM Vol. 10 Sphaerodactylus inacrolepis 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 15897-98, UIMNH 56986-87, 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. A55OCIATED SPECIMENS vent length of specimens larger than the median of all in the size classes of 28-28 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 187 specimens; ventral scales axilla to groin 22 to 29, mean 25.5 in 185 specimens; midbody scales 85 to 45, mean 89.3 in 103 specimens; fourth toe lamellae 8 to 11, mode 10; modal internasal count for this form is 1, range 1»8; 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-yent classes of 22-23 mm, mean 22.8 mm, as opposed to other townsendi for which this range is 28-28 mm, mean 24.8 mm. Thus in size, these specimens are 1966 PUERTO RICAN SPHAERODACTYLUS 247 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, V5888-48, Caja de Muertos; ASFS V58I2-27, 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 X1017-82, 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 Sphaerodactt/lus 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 Cordia, British Virgin Islands. DISTRIBUTION: Known only from the island of Virgin Gorda (fig. 5). DEFINITION: A species of Sphaerodactylus with small, acute, strongly keeled, Hattened, imbricate dorsal scales, axilla to groin 80- 85, mean 82.0; 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 8, 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 Hat-lying than truly erect); scales beneath tail smooth, rounded, enlarged mid- ventrally; fourth toe lamellae 8 (mode) or 9; escutcheon (fig. lig) 248 BULLETIN FLORIDA STATE MUSEUM Vol. 10 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 Bne 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. 84 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. 0 0 * 00 o 17. 009 45' 0 0 2NI 0 64'40'· FIGURE 9. St. Croix, American Virgin Islands, showing locality records for S. macrolepis macrolepis (open symbols) and S. beattvi (solid symbols). Race5 of S. beattyi: S. b. bet:ttvi, horizontal lines; S. b. seamani, vertical lines. Records of S. m. macrolepis are of specimens examined, not from the literature. 1966 PUERTO RICAN SPHAERODACTYLUS 249 ~PECIMENS EXAMINED (all from Virgin Gorda): ASFS V8658-59, AMNH 92821, MCZ 77212=14, ca. 0.5 mi. N Pond Bay; MCZ 77211 (type), AMNH 92822- 24, hillside above Pond Bay; ASFS V8664, Pond Bay, ASFS V8681, 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 beatt!/i Grant Sphaerodactylus beatt!/i 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 Beld data). DISTRIBUTION: 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 c6nceivable 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 (axilla to groin 84-44). 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 8; dorsal scales of tail small, acute to rounded, keeled, slightly swollen and slightly erected; scales beneath tail smooth, rounded, enlarged 250 BULLETIN FLORIDA STATE MUSEUM Vol. 10 midventrally; fourth toe lamellae 7 to 11, mode 10; escutcheon (fig. 11h) compact with no extension onto thighs, 4-6 X 8-12. Coloring a uniform brown or yellowish brown (juveniles) er 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 26-80 mm, average 27.2 mm. REMARKS: 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 beauui beattui Grant, new combination Sphaerodact{/lus 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. DISTRIBUTION: 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 beatti/i 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 86 to 44, mean 41.9; ventral 5cales axilla to groin 29 to 35, 1966 PUERTO RICAN SPHAERODACTYLUS 251 mean 82.1; 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 6f 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 80788) contains both heavily patterned and unpatterned specimens. These may represent intergrades or they maybe the result of pooling of localities. SPECIMENS EXAMINED: St, Croix: ASFS V8600-14, 1.0 mi. NE Lavallee; ASFS V8246-49, 5 mi. E Christiansted; ASFS V3251-56, 7 mi. E Christiansted; ASFS V8568-91, 1.0 mi. W of eastern tip of island; ASFS V8561-65, ca. 0.75 mi. W of eastern tip of island; UMMZ 80788 (17 specimens, intergrades?), east end of St. Croix; ASFS V3827-51, Milord Point (west side of Great Pond Bay); UMMZ 80567 (holotype), ?"Good Hope." P I . - jr - 5 111 3 1 4 11 4 r, 15 1# d- 7 r *4 +9) , 4 il' 1 , , -t414'lo "- 1 917£ '''-1 P .6 ,/ I.'' , jkll'' ' ' f 'I--/ 24-* 11 - 1 11 J1 1 1 lili - :1 4 S A B FIGuRE 10, A, S. beattyi beattyi, anterodorsal view„ female, ASFS V8564, 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. 252 BULLETIN FLORIDA STATE MUSEUM Vol. 10 Sphaerodactylus beattyi seamani new subspecies TypE: MCZ 81056, an adult male from ca. 0.5 mi, ea'st 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-SOL V8540-48, KU 79917-28, MCZ 81057-62, UF 21277-82, UIMNH 56945-50, USNM 152599-600, same data as type; ASFS V8592-99,2 mi. W Grapetree Bay, St. Croix, 8 August 1965, R. Thomas. ASSOCIATED spECIMENS: UMMZ 80568-71 (paratypes of S. beatt!/i), 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. beatt!/i 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. 1Ob). DESCRIPHON 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 82; scalek around midbody 68; 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 5icles; 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 Recked and mottled with dark brown; ground color of throat in life blue-gray; venter light gray; underside of tail yellow to orange; iris golden. VARIATION: In dorsal scale counts axilla to groin the range in 22 specimens is 87 to 44, mean 89.8; ventral scales axilla to groin 26 to 85, mean 80.9, midbody scales 57 to 64, mean 61.5. Qther 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 1966 PUERTO RICAN SPHAERODACTYLUS 253 20 percent. The dorsa of the paratypes vary from almost unicolor like the type to a pattern of bold brbwn mottling or reticulations. The cephalic pattern fs 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 specimens have a dark stippling on the venter. FIGURE 11. Escutcheons of Greater Puerto Rican Sphaerodactylus, as follows: A, S. macrolepis phoberus, ASFS X909; B, S. monensis, MCZ 84568; C, S. roose- uetti, ASFS X4288; D, S. klauberi, ASFS V4507; E, S. gaigeae, ASFS X4050; F, S. nicholsi nicholsi, ASFS X4281; C, S. parthenopion, ASFS V3681; H, S. beattyi beattyi, ASES 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: 254 BULLETIN FLORIDA STATE MUSEUM Vol. 10 1. Scalation: Dor5al scales acute, keeled, flattened, no zone Of middorsal granules, scales of tail rRat-lying. 2. Scale organs: Hair-bearing organs are similar in kind and disposition. 8. 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, rooseuelti) certain elements keem 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 1966 PUERTO RICAN SPHAERODACTYLUS 255 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 kecond 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 rooseuelti, klauberi, gaigeae, nitholsi, 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 Sphae,o- 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. Tooseuetti as an allopatric isolate in the southwest, prob- ably on a peninsula or chain of offshore keys. b. Derivation of the nicholsi-pal'thenopion progenitor as an ecologically specialized form in the arid littoral. c. Derivation of th6 klauberi-gaigeae progenitor as an eco- logically specialized form in mesic forest. d. Isolation of the monensis progenitor on Mona either by 256 BULLETIN FLORIDA STATE MUSEUM V61. 10 geological separation of Mona from Puerto Rico or by dispersal across the water gap. 8. Continued evolution towards the present day macrolepis. 4. Invasion of the northern Leeward Islands by macrolepis and subsequent differentiation. 5. Invasi6n over water to St. Croix by macrolepis. KLAUBERI MONENSIS GAIGEAE PROTO-MACROLEPIS r MACROLEPIS %1CHOLS\ ROOSEVELTI PARTHENOPION FIGURE 12. Diagram of hypothetical evolutionary relationships of Greater Puerto Rican Sphaerodact!/lus, with the exception of S. beatt!/i. 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 prdisumably 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 differdnt 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 1966 PUERTO RICAN SPHAERODACTYLUS 257 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 clistribution 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 southeru 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 upiand 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 56utheastern 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 8) phoberus- grandisquamis-stibarus in northeastern and eastern Puerto Rico and on Isla Pifieros. Of the Puerto Rican races ody 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 Piheros, 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, 258 BULLETIN FLORIDA STATE MUSEUM Vol. 10 From the above, it is easily observed that the major companion of S. inacrolepis 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- renee 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* 1 E E € 5 uodoum pied augBIEB0 0 Cd IsIo oru6 23 macr61epis - roosevelti - klauberi - - m I pi s 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 difBcult 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 f6r 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, 1966 PUERTO RICAN SPHAERODACTYLUS 259 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 ha's 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 P6rto Rico, Culebra and Mona islands. Jour. Dept. Agri. Porto Rico, vol. 15, no. 8, pp 199-218, 5 pls. 1932a. A new sphaerodactyl from Porto Rico. Jour. Dept. Agn. 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, pA 88-85, t' chart. 1932c. The herpetology of Vieques Island. Jour. Dept. Agri. Puerto Rico, vol. 16, no. 1, PA 87-39, 1982d. Sphaerodactylus grandisquamis, a valid species. Jour. Dept. Agn. Puerto Rico, vol. 16, no. 1, pp. 43-45. 1987. 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. 110-118, 1 fig. ' Heatwole, Harold 1961. Institute of Caribbean Studies Field Excursion to Isla Mona. Herpe- tology. Caribbean Jour. Sci.,,vol. 1, no. 1, pp. 32-88. - Donald S. Sade, and Richard Hildreth 1963. Herpetogeography of Puerto Rico. I. Herpetofauna of Cayo Santiago and Cayo Batata. Caribbean Jour. Sci,, vol. 8, no. 1, pp. 1-5, 2 Egs. King, Wayne 1962. Systematics of Lesser Antillean lizards of the genus Sphaerodactulus. 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. 260 BULLETIN FLORIDA STATE MUSEUM Vol. 10 Rivas, Luis Rene 1964. A reinterpretation of the concepts "sympatric" and -allopatric" with proposal of th6 additional terms "syntopic" and "allotopic". Systematic Zool,, vol, 13, pt. 1, pp. 42-43, tiolle: 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. 821-822. 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 1964. The herpetology of Porto Rico. Rept. U. S. Nat. Mus,, 1902, pp. 549- 724, 1 pl., 197 figs. Thomas, Richard MS. Additional notes on the amphisbaenids of Greater Puerto Rico. Breviora, in press. Contributions to the BULLEZIN OF THE FLORIDA STAfE MUSEUM may be in any field of biol6gy. 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In it list all new organisms described and give their ranges; indicate all taxonomic changes pro- posed. The synopsis, written in full sentences, should be concise, but completely intelligible in itself without reference5 to the paper, thereby enabling the busy reader to decide more surely than he can from the title alone whether the paper merits his reading. The synopsis will be published with the paper. It does not replace the usual conclusians or summary sections. It may als6 serve as copy for the abstracting services. Manuscripts and all editorial matters should be addressed to: Managing Editor of the BULLETIN Florida State Museum Seagle Building Gainesville. Florida