BULLETIN OF THE FLORIDA STATE MUSEUM BIOLOGICAL SCIENCES Volume 7 Number I SYSTEMATICS OF LESSER ANTILLEAN LIZARDS OF THE GENUS SPHAERODACTYLUS Wayne King . Mn ,1 .-isig- I UNIVERSITYOFFLORIDA Gainesville 1962 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. WILLIAM J . RIEMER, Managing Editor OLIVER L. AUSTIN, JR., Editor Consultants for this issue: Ernest E. Williams Garth Underwood Communications concerning purchase or exchange of the publication and all man- uscripts should be addressed to the Managing Editor of the Bulletin, Florida State Museum, Seagle Building, Gainesville, Florida. ~ Published 81 December 1962 ~ Price for this issue 604 SYSTEMATICS OF LESSER ANTILLEAN LIZARDS OF THE GENUS SPHAERODACTYLUS WAYNE KINGl SYNOPSIS: Poor descriptions, resulting in the misapplication of names, and the unneeded multiplicity of recognized forms has confused the taxonomy of the lizard genus Sphaerodactulus in the Lesser Antilles. Examination of specimens available from the area shows eight species to occur. Each is described and illustrated, and its known range listed. The presence of two species in the type series of S. sputator necessitates the designation and description of a lectotype. Two new races, S. macrolepis parous and S. fantasticus ligniseroutus are described from the northern Lesser Antilles. S. festus is treated as a subspecies of S. vincenti and is shown to include variation thought distinctive of S. monilifer. The sphaerodactyls of the Lesser Antilles are believed to have immigrated into the isiands from continental South America. The most probable means of dispersal available to them is ,rafting. 1The author is a John M. Prather Fellow in the Department of Zoolbgy of the University of Chicago. This paper represents, in part, a thesis prepared in partial fulfillment of the requirements for the degree of Master of Science in the Department of Biology of the University of Florida. Fieldwork for this study was supported by the Florida State Museum and National Science Foundation grant G-3896. Manuscript submitted 10 February 1962. The author was unable to examine proofs of this article before publication.-ED. King, Wayne. 1962. Systematics of Lesser Antillean lizards of the genus Sphaero- dactulus. Bull. Florida State Mus., vol. 7, no. 1, pp. 1-52. 2 BULLETIN FLORIDA STATE MUSEUM Vol. 7 INTRODUCTION In 1784 Anders Sparrman described the lizard Lacerta sputator C = Sphaerodactylus sputator) from the island of St. Eustatius. Lesser Antilles. Since then a number of species of Sphaerodactylus have been described from Other islands in the Lesser Antilles. The descriptions in most cases were based on the color pattern of one or a few speci- mens in the belief that this character varied little, and on the assump- tion that species were conGned to single islands ( Barbour, 1921:264) This resulted in, widespread misapplication of names to species with similar ccdor patterns, and ad unfortunate multiplicity of named forms. This study attempts to clarify the taxonomy of this group in the Lesser Antilles. I have examined most of the specimens available from the area and tried to determine the individual variation and distribu- tion of each of the forms. The Lesser Antilles are a discrete geographic group of islands sepa- rated from the Virgin Islands to the north by the Anegada Passage, and from Trinidad and Tobago to the south by the Tobag6 Trough. They also represent the present zone of contact between typically Antillean and typically South American species. The species of Sphaerodactylus that occurs on Trinidad and Tobago appears to be related to species in the southern Lesser Antilles. For this reason I have included these tw6 islands in the study area, even though they are not a part of that island chain. The study includes, from north to south, the islands of Sombrero, Dog Island, Anguilla, Tintamarre, St. Martin, Ile Fourche, St. Barts ( = St. Barthelemy, = St. Bartholomew), Saba, Statia ( = St. Eustatius), St. Kitts ( = St. Christo- pher), Nevis, Barbuda, Antigua, Redonda, Montserrat, Guadeloupe, D6sirade, Iles des Saintes, Marie Galante, Dominica, Martinique, St. Lucia, St. Vincent, Barbados, The Grenadines, Grenada, Tobago, and Trinidad. ACKNOWLEDGMENTS ~ I wish to thank Wm. J. Riemer and W. Auffenberg for their continued patience, interest, and criticism, and for making it possible for me to visit some of the Lesser Antilles in order to collect many of the specimens used in this study. I would like to thank M. Boeseman, C. M. Bogert, D. M. Cochran, J. Guibd, N. Hattweg, P. W. Humme- linck, R. F. Inger, A. Johnels, G. Underwood, and E. E. Williams for the loan of specimens in their care. Thanks are also due D. A. Belkin and P. C. Drummond for their enthusiasm and interest in West In- dian zoogeography. 1962 KING: ANTILLEAN SPHAERODACTYLUS 3 MATERIALS AND METHODS The characters used in this study are defined as follows: SHAPE OF DORSAL SCALEs. The dorsal scales fall into four general groups based on shape-large keeled scales, keeled granular scales, granules, and smooth scales. Only the first three of these scale types are found in the study area. The large keeled scales, granular scales, and smooth scales imbricate, while granules are juxtaposed. MIDI)ORSAL AREA OF GRANULES. An area of juxtap6sed granules in a middorsal ( vertebral ) band on the trunk. Although the aroa of gran- ules may be present in species possessing small granular scales, it is well defined only when associated with ihe larger scale types-large keeled scales and keeled granular scales. The width oE this area is expressed as the number of transverse scale rows ( granules). SCALES FROM AXILLA TO GROIN. The number of scales beginning with one lying on a line with the posterior edge of the forelimb counted caudad to one lying on a line with the anterior edge of the hindlimbs. Dorsal scales are counted in a dorsolateral position to avoid the mid. dorsal granules of some species. Ventral scales are counted along a midventral line. MICROSCOPIC STRUCTURE OF DORSAL SCALES. The presence or absence of knoblike or "hair-bearing" scale organs on the dorsal surface ( fig. 1 Al to Ad). These scale organs are also found in other groups of lizards ( Schmidt, 1920:113; Underwood, 1957:277). In Sphaerodact~- lus they occur on other scales of the body, but only the dorsal scales of the trunk are used in this study. These structures are observed by removing loose dorsal scales ( the corneal layer of the epidermis which is shed during ecdysis) from the specimen and mounfing them in air on a slide. The indices of refractitin of the scales and conven- tional mounting media are close enough to obliterate much of the detailed structure on the scale. - SHAPE OF VENTRAL SCALES. The ventrals are keeled or smooth, acute or round. SCALE ROWS AROUND MIDBODY. The number of scale rows around the middle of the trunk, including middorsal granules, if present. INTERNASALS. The number of scales lying between the supranasals, and in contact with the rostral scale. UPPER LABIALS. The number of upper labials, counted from the labial touching the rostral scale caudad to a point directly below the center of the eye. The count is made on the right,side. GULAR SCALES. These scales are smooth, keeled on the sides of the 4 BULLETIN FLORIDA STATE MUSEUM Vol. 7 A| A 2 A4 7/3 2 FIGURE 1. Types of dorsal scale organs and dorsal caudal scutellation. C Al) no scale organs; (Az) hair-bearing organs with one hair each; (AB) knoblike organs and hair-bearing organs, with one or two hairs each ( some species have more hairs); (A#) 1moblike organs; (Bl) caudal scales lie flat against the tail, or (82) stand erect from the tail. throat, keeled in a band ( collar) across the throat, or keeled across the throat and caudad to the chest and abdomen. 1962 KING: ANTILLEAN SPHAERODACTYLUS 5 LAMELLAE OF FOURTH TOE. The number of lamellae under the fourth toe of the right foot, exclusive of the terminal expanded disk, counted proximally to the last large, expanded lamella ( fig. 5 A). MIDVENTRAL SCALE ROW OF TAIL. The scales of the midventral row ~ of the tail are transversely expanded into plates ( shields), enlarged (larger than adjacent rows, but not wide plates), or unexpanded ( same width as the lateral and dorsal rows). Because regeneration alters this character, only specimens with original tails were used in this count. DORSAL SCALES OF TAIL. These scales generally lie flat against the tail ( fig. 1 Bl ), but in some species they stand erect (fig. 1 82) giving , the tail a rough appearance. This is not an artifact caused by dehydra- tion in preserving fluid. Only specimens with original tails were used for this character. PRESACRAL VERTEBRAE. The number of vertebrae, counted from the atlhs caudad to, but not including, the first sacral vertebra with ex- panded diapophyses in contact with the ilium. Tbis count is made from roentgenograms. A total of 318 specimens of the species examined ( including all type specimens examined) were counted. ESCUTCHEON OF MALE. The area of differentiated glandular scales in the inguinal region of the males has been called the escutcheon (Grant, 1981: 199). The width of the escutcheon is the number of scale rows at its greatest width, including extensions along the thighs. The length is the number of scales on the midventral line counted from the anteriormost to the posteriormost. SNOUT-VENT LENGTH. The distance from the tip of the snout to the anterior margin of the cloaca. The total length is not given for most of the species as many individuals have broken· or regenerated tails. The total length corresponds roughly to twice the snout-vent length. Snout-vent length is measured with calipers and millimeter rule. Individuals are sexed by secondary sex characters, primarily the escutcheon of the male. Noble and Klingel ( 1982: 15 ) state that the presence of the escutcheon is: . . . less diagnostic of sex than another male character which we believe has not been hitherto described. The posterior lip of the cloaca is very much broader in the adult male than in the adult female. Most females... have the cloaca closed and exhibit little. or no fleshy margin to the caudal side of the cloaca. Most males ' . . . have this fieshy margin broader and exhibit a tendency for it to fold back towards the tail. Noble and Klingel themselves point out the weakness of this cloacal character in zdetermining sex, as poorly preserved male specimens do 6 BULLETIN FLORIDA STATE MUSEUM Vol. 7 not show the opened cloaca. Examination of a series of specimens shows this method -inadequate, owing to the large number of poorly preserved specimens. Grant ( 1982:405) states that: My study of a series of 2,085 specimens of nine species from the Puerto Rico Area has shown that tbe escutcheon is visible in newly hatched specimens and not confined to "adults." Examination of the specimens used in this study (including a num- ber of those Grant used in his study) fails to show the escutcheon in any individual of less than 18 mm. snout-vent length, though juve- niles of lS to 15 mm. snout-vent length are plentiful. The smallest adult size given in each species description is based on the snout- vent length of the smallest specimen with an escutcheon. The number of dorsal scales equivalent in length to the "standard length" ( snout-eye length), has previously been used t0 indicate the relative Size of the dorsal scales. Juvenile individuals, however, have relatively larger heads than do adults, and consequently have more dorsal scales contained in their "standard length." Grant ( 1959a:57) shows how this led to the description of the juveniles of S. cinereu~ as a separate species (S. elegans)-the juveniles have 24 to 25 dorsal scales in the "standard length"; the adults, 18 to 19. The measure of relative scale size that I used, the number of scales from axilla to groin, is fairly constant within a species, and does not vary ontogenetically. Individual color patterns of these animals vary as widely as any morphological character in the populations. In an eHort to include total color pattern variation of each of these forms I use only gener- alized terms in the calor descriptions. Colors are grouped into three categories-light, medium, and dark. Except where stated, all color descriptions refer to alcohol specimens. Each species description is accompanied by an illustration that shows the total observed variation in color pattern. Except in the case of the types of three species, descriptions of individuals are not given, instead reference is made to the species as a whole. In these three special cases the descriptions of the types also contain, in parentheses following each character, the variation which occurs within the species. The description and variation of the species is based. on the total observed variation in the material examined. Throughout this paper the following abbreviations are used to refer to specimens or collections: 1962 KING : ANTILLEAN SPHAER©bACTYLUS 7 AMNH-American Museum of Natural History, New York CNHM-Chicago Natural History Museum, Chicago MCZ--Muse~im of Comparative Zoology, Cambridge, Massachusetts . MNHN-Mus6um National d'Histoire Naturelle, Paris; France PWH-P. Wagenaar Hummelinck collection, Utrecht, Netherlands RMNH-Rijksmuseum van Natuurlijke Historie, Laiden, Netherlands SMNH-Naturhistoriska Riksmuseum, Stockholm, Sweden UF.-University of F.lorida Collections, Gainesville, Florida UMMZ-University of Michigan Museum of Zoology, Ann Arbor, Michigan USNM-U.S. National Museum, Washington, D.C. Comparative material in the genus Sphaerodactylus used includes: S. anthracinus anthracinus, New Providence, Bahamas S. anthracinus copei, Haiti ' S. altavelensis, Alta Vela Island, Dominican Republic S. argious, Cayman Brad S. argus, Jamaica S. bartschi, Little Gay.man Island S. beattyi, ·SE Croix, Virgin Islands S. becki, Navassa Island, Haiti S. caicosensis, Caicos Islands,. Bahamas S. cmereus, Florida S. dillicilis, Dominican Republic S. dunni, Honduras S. gaigei, Puerto Rico S. glau'Ous, Mexico . S. gonibrhynchus, Jamaica S. tewisi, Grand Cayman Island S. tineolatus, Costa Rica S. notatus, Florida S. oliceR haveri, Cuba S. oxyrrhinus, Jamaica S. pac#icus, Cocos Island, Costa Rica S. parkeri, Jhmaica S. ramsdehi, Cuba . E S. richardsoni gossei, Jamaica S. rooseuelti, Puerto Rico S. steinegeri; Haiti , S. tou)nsendi, Puerto Rico 8 BULLETIN FLORIDA STATE MUSEUM Vol. 7 SYSTEMATICS Several closely related Antillean forms are considered here as sub- species. Because they comprise insular populations separated from each other by open ocean, no intergradation can occur. Nevertheless the great similarity in color patterns and other morphological char- acters suggests strongly that if the animals were not physically sepa- rated they would probably interbreed freely. KEY TO THE SPECIES OF SPHAERODACIYLUS IN THE LESSER ANmLES 1 Ventral scales keeled on abdomen 2 Ventral scales not keeled · · · .- ... 8 2 Caudal scales stand erect, imparting fuzzy appearance to tail; midventral tail row not expanded; no middorsal area 6f granules; 42-58 dorsal scales from axilla to groin; St. Lucia and Dominica , S microlepis Caudal scales lie flat against tail; midventral tail row expanded into trans- verse plates; middorsal area of granules present; 23-29 dorsal scales from axilla to groin; Guadeloupe S. fantastious 3 Well-defined middorsal area of granules . 4 No well-defined middorsal area of granules 7 4 Cular scales sm66th; dorsal Scales with hair-bearing scale organs only; snout-vent length 15-86 mm.; Statia, Dog Island, St. Martin, St. Barts, Ile Fourche, St. Kitts, and Nevis 0 sputator Gular scales keeled, at least on sides of throat; dorsal scales with both hair-bearing and knoblike scale organs; snout-vent length 18-29 mm __ 5 5 Presacral vertebrae 27; 88-42 dorsal scales from axilla to groin; 47-66 scale rows around midbody; Statia, St. Kitts, and Nevis - S. sabanus Presacral vertebrae 26; 23-38 dorsal scales from axilla to groin; 41-54 scale rows around midbody 6 6 Gular scales keeled on side 6f throat, or in narrow band across throat; juvenile pattern of crossbands, adult pattern uniform brown or vermiculat- ed; Antigua and Barbuda S elegantulus Gular scales strongly keeled on entire throat, keeled scales may extend caudad to the chest; juvenile and adult pattern of longitudinal stripes, or uniform brown; Guadeloupe and Montserrat - -S fantasticus 7 Dorsal scales large, keeled, and strongly imbricate, 20-29 from axilla to groin; 40-51 scale rows around midbody; Anguilla, Dog Island, St. Barts, St. Martin, and Tintamarre S macrolepis Dorsal scales weakly keeled, aild only slightly imbricate, 86-74 from axilla to groin; 58-82 scale rows around midbody 8 8 Dorsal scales with only hair-bearing scale organs; 64-74 dorsal scales from axilla to groin; Trinidad, Toboga, British Guiana, and Venezuela __-S. molei Dorsal scales with both hair-bearing and knoblike scale organs; 36-54 dorsal scales from axilla to groin; St. Vincent, St. Lucia, Dominica, and Marti- nique S uincenti 1962 KING : ANTILLEAN SPHAERODACTYLUS 9 SPECIES DESCRIPTIONS Sphaerodactylus sputator ( Sparrman) Lacerta sputator Sparrman, 1784: 164 (in part). Sphaerodactylus pictus Carman, 1887:20. Sphaerodactulus sputator Andersson, 1900:27 ( in part); Barbour, 1921:266 ( in part); Barbour,-1923:2 (in part). Anders Sparrman ( 1784:164) described Lacerta sputator ( = Sphaero- dactylus sputator) from the island of Statia on the basis of three syn- types in the Swedish Museum of Natural History, Stockholm. He described, sputator as an ash-gray lizard with brown-edged white crossbands on the dorsum ( hereafter in this discussion it will be called xariation 1-it corresponds to plate 4, fig. 1 of Sparrman's description). The smallest of the three syntypes lacked the white crossbands and had brown spots instead ( variation 2 in this discussion-plate 4, fig. 2 of Sparrman's description). Sparrman described this small specimen as a 'larval," i.e. juvenile, variation of the typical adult pattern, because the tail, which was acfually being regenerated, was not yet completely f6rmed. The island of Statia was visited so infrequently that the species was not collected there again for 188 years. This led many later authors to apply the name Sphaerodactylus sputator to crossbanded sphaero- dactyls from other islands in the Antilles. Andersson ( 1900:27), in an attempt to clarify the situation, re-examined the three types and stated that two possessed the characteristic crossbands ( variation 1); the third (Variation. 2.) was not described. Barbour ( 1914:270) pointed out some of the confusion surrounding the identity of sputator and said he believed the species was probably confined to the island of Statia. Later (1921:266) Barbour encouraged Andersson to supply the description of the types for his monograph of the genus. Anders- son described the syntypes as: . . . Two specimens . ·. . Head above indistinctly dotted with brown; between ear and vent of pairs of brown bands, the first immediately behind the ear, 2nd in front of, 8rd behind, the axilla, 4th, 5th and 6th on the body, 7th at the Vent. The basal part of the tail is provided with 6-8 bands, more or less distinctly arranged in pairs; the distal part shows no bands only small dots, in one specimen it is regenerated. The #malle* specimen ... has no bands. The head and upper surface of the neck is very distinctly and den5ely spotted with brown; the anterior part of the body indistinctly dotted. 10 BULLETIN FLORIDA STATE MUSEUM Vol. 7 In that publication the largest and smallest of the syntypes are figured ( plate 8, figs. 3 and 4), clearly showing the crossbanding of the large specimen ( variation 1) and the vermiculation of the small one ( variation 2). On re-examining Sparrman's types of the species ( fig. 2) I find two species present-two large specimens ( variation 1) are of the species now called Sphaerodactglus sputator C Sparrman ), and the small speci- men ( variation 2) is of the species now called Sphaerodactylus sabanus Cochran. The smallest specimen has 27 presacral vertebrae, keeled gular scales on the sides of the throat, and has both knoblike and hair=bearing scale organs on the surface of the dorsal scales. The two large specimens have 26 presacral vertebrae, smooth gular scales, and,only hair-bearing scale organs on the dorsal scales. King ( 1960:2) points out that Barbour ( 1928:2) similarly confused the two species in a series of specimens collected on Statia in 1922 by James L. Peters. Barbour stated that sputator was dichromatic-the females ( actually A B C I 7%· 1- 23*4109**:ff*.g f6434/ f ·>2.'51 FIGURE 2. Syntypes of Lacerta sputator Sparrman ( -Sphaerodactutus sputator). (A) SMNH 2670; (B) SMNH 2669, (C) SMNH 2671. A and B are of the species now called S. sputator, C is S. sabanus. B is the newly designated lecto- type of S. st)utator. Relative size and position of the specimens in this and the following figures is diagrammatic to facilitate comparison of color patterns. 1962 KING: ANTILLEAN SPHAERODACTYLUS , 11 sputator) large and with crossbands and blotches, the males ( sabanus) small and uniform gray brown. The largest specimen of the type series is here redescribed and designated as the lectotype of Sphaerodact!/lus sputator ( Sparrman ). _ DIAGNOSIS. A large sphaerodactyl with large, keeled, and slightly imbrieate dorsal scales, and a middorsal area of granular scales. The dorsal scales with hair-bearing scale organs.on their surface. Ventrals smooth and round. Gulars smooth. Internasals 1, upper labials 3. Dorsal scales of tail imbricate, but lie flat against tail. Presacral yer- tebrae 26. Groundcolor light brown. Pattern of dark brown-edged _ white crossbands, dark brown blotches, or dark brown longitudinal stripes. LECTOTYPE. SMNH 2669, an adult female, collected by Dr. Acrdlius in 1755 on the island of Statia ( fig. 28). DESCRIPTION OF i,ECTOTYPE ( AND SPECIES). D6rsal scales large, keeled, acute to rouild, and slightly imbricate; 84 axilla to groin ( 31-89 in Other specimens, mean 84.6). Hair-bearing scale organs, each with 1 to 2 hairs, along dorsodistal edge Of dorsal scales. Area of middorsal granular scales present, 2 scale rows wide. Ventrals smooth and round; 29 axilla to groin ('27-84 in other specimens, mean 30.8 ). Culars smooth. Scale rows around midbody 56 ( 49-62 in other speci- mens, mean 55.5 ). Internasals 1 ( 1-2, mode 2). Uppef labials 8 ( 8-4, mode 3)., Lamellae of fourth. toe 10 ( 8-12, mode 10). Escuteheon of - male ( the lectotype is a female) 26 to 80 scales wide, 8 to 6 scales in length. Dersal scales· on tail weakly keeled and imbricate, and lie Hat against tail. Midventral row transversely expanded into plates. Presacral vertebrae 26. Snout-vent length 86 mm. ( snout-vent length of smallest juvenile is 15 mm.; adults 20 to 86 mm., mean 31.5). The dorsaI groundcolor is light brown. The pattern consistS of six brown-edged, dull-white crossbands between the head and the groin ('fig. 2 B). The crossbands continue onto the tail but are ill-defined and obscure. The entire animal has a slight greenish cast, as though it had been exposed to the corrosion products of zinc 'or brass. The throat has a small hole punctured between the mandibles. The tail is broken ofE close behind the vent, and is loose in a vial with the lectotype ( the tail itself has been broken into two pieces now held t6gether by a pin running. longitudinally through them). In other individuals the pattern exhibits neither sexual dichroma- tism nor. ontogenetic variation. The groundcolor is light br6wn with 5 to 8 dark br6wn-edged white crossbands between the head and groin (2-4 on, the head and neck, 3-4 on the trunk). In some indiviauals the brown edges invade the white crossbands creating a brown and 12 B U LLE T IN FLO R ID A S TATE M U S E U M Vol. 7 A C CD .· r· . 't·.·3 FIGURE 3. Variation in the niarkihgs of adult Sphaerodact!/tus sputator. (A) RMNH 10456; (B) RMNH 10456; (C) UF 10039.1; (D) MCZ 16614; (E) MCZ 16622; (F) MCZ 166I1. 1962 KING: ANTILLEAN SPHAERODACTYLUS 13 white spotted animal. In others the brown edges not 6nly invade the white crossbands, but also realign and fuse into dorsolateral stripes creating a longitudinally striped animal. All of these variations in pattern are shown in Bgure 8. RANGE. The islands of Statia, D6g Island ( west of Anguilla), St. Martin, St. Barts, Ile Fourche, St. Kitts, and Nevis. Its known range suggests that it probably occurs on Anguilla and Saba also. REMARKS. A number of females collected from Febfuary to June are gravid. Each contains one mature shelled egg. Those I collected are from beneath rocks on St. Martin and from a pile of coconut husks on St. Kitts. Its closest relationships seem to lie with S. elegantulus. S. sputator is unique in that it is the only sphaerodactyl examined during this study with a well-defined middorsal area of granules and that lacks knoblike scale organs and has only hair-bearing scale organs. MATERIAL EXAMINED. Statia: SMNH 2669 (lectotype), 2670; MCZ 16598-16688, 16635-16641 (80); RMNH 10451; UMMZ 57010. St. Kitts: MCZ 6071 (8); UF 10038. Nevis: RMNH 10449. St. Martin: PWH 474A, 606; RMNH 10486, 10487, 10489 (8), ,10440, 10441 (2), 10442, 10448, 10445 (2), 10456'(2), 10457, 10458, 10459, 10460 (4), 10462, 10464; UF 10089 (2). St. Barts: RMNH 10485, 10458 (2). Ile Fourche: RMNH 10455. Dog Island: RMNH 10446 (8). Sphaerodactylus sabanus Cochran ' Lacerta sputator Sparrman, 1784 : 164 ( in part ). Sphaerodactylus st)utator Andersson,. 1900:27 ( in part); Barbour, 1921:266 ( in part). Sphaerodactylus sabanus Cochran, 1938:148. DIAGNOSIS. A sphaer6dactyl with keeled, granular scales and a mid- dorsal area of granular scales. Dorsal scales with both knoblike and hair-bearing scale organs. Ventrals smooth and round. Gulars keeled. Internasals 1 to 8, upper labials 8. Dorsal Scales on tail imbricate, but lie flat against tail. Presacral vertebrae 27. Groundcolor brown, dark to light. Pattern of longitudinal dark stripes on head and body and light dorsolateral stripes on tail, or dark vermiculate markings on head, or uniform brown head and body. HOLOTYPE. USNM 108985, an adult male collected by S. T. Danforth On the island of Saba in January 1937. DESCRIPTION OF SPECIES. Dorsal scales keeled and granular, slightly imbricate;.88 to 42 axilla to groin, mean 38.0. Dorsal scales with knob- like scale organs and hair-bearing scale organs, each with 1 to 3 hairs. An area of middorsal scales present, 1 to 2 scale rows wide. Ventrals 14 BULLETIN FLORIDA STATE MUSEUM Vol. 7 smooth and round; 28 to 83 axilla to groin, mean 30.7. Gulars keelekl, at least on the sides of throat. Scale rows around midbody 47 to 66, mean 57.6. Internasals 1 to 8, mode 2. Upper labials 3. Lamellae of fourth toe 7 to 11, mode 9. Escutcheon of male 22 to 28 scales wide, and 5 to 6 scales long. Dorsal sdales of tail keeled, acute to round, imbricate, but lie flat against tail. Midventral row transversely ex- panded into plates. Presacral vertebrae 27. Sndut-vent length of small- est juvenile is 15 mm., adults 20 to 29 mm., mean 25.2. The pattern shows neither sexually dichromatic nor marked onto- genetic variation. The dorsal groundcolor is light to dark brown, A dark brown stripe extends from the nostril through the eye and onto the neck, where it swings toward the midline and fuses with the one from the opposite side just anterior to the scapular region. A dark brown line extends from the tip of the snout caudad be]tween the eyes, where it divides, each half passing to a point immediately pos- terior to the eye. Each then continues caudad and returns toward the midline, circles an occipital spot, and fuses with the dark lines from the opposite side just anterior to the scapular region and the point of fusion of the dark postorbital stripes ( fig. 4 A). The oecipital spot is usually a dark brown to black spot, or is a white spot with a dark edge. The pattern on the trunk consists of a dorsolateral row of light brown to white spots, which become more pronounced in the sacral S ·17/5 . A :'frim D C €619«Sh:/V EK.~Vi 4 A¥ 21 FIGURE 4. Variation in the markings of adult Sphaerodactulus sabanus.(A) UF 10045.1; (B) MCZ untagged specimen in series 54010-54015; (C)UF 10042.3; ( D) UF 10044.7. 1962 KING: ANTILLEAN SPHAERODACTYLUS 15 region and are united· to,form white stripes on the tail. In some individuals the dark head stripes break up to form a series of spots roughly suggesting the original pattern ( fig. 4 B), or they fuse with one another to form a vermiculate head pattern (fig. 4 C). Other specimens, both juveniles and adults, are uniform brown (fig. 4 D). The ventral color is light brown to white. The juvenile pattern is identical to the adult pattern except that the longitudinal stripes on the head of the juveniles seem to be slightly more pronounced, and the vermiculate head pattern of some adults is more pronounced. RANGE. The islands of Saba, Statia, St. Kitts, and Nevis. REMARKS. Females collected during May and June have mature, shelled eggs in their oviducts. I took specimens under rocks on St. Kitts and beneath coconut husks and palm_ fronds, on Nevis. As pointed out in the discussion of S. 5-putator, S. sabanus is often confused with that species because the two se6m to occupy the same microhabitat and are often collected together. King ( 1960:4, see also Barbpur, 1929:81) shows'that S. sabanus is often confused with S. elegantulus from Antigua and Barbuda because of the superficial similarity between uniform brown adults of thu two species. The)/ are undoubtedly closely related. S. sabanus differs from all other sphaerodactyls I have examined in having 27 presacral vertebrae-all other forms examined have 26. MATERIAL EXAMINED. Saba: USNM 108985 (holotyph), 103986, 108988-103993, 103995-104003; MCZ 45215-45217; RMNH 10438 (2), 10434, 10452. Statia: MCZ 54010-54015 ( 158 ); RMNH 10432 (4), 10450 (2); SMNH 2671. St. Kitts: PWH 422; UF 10041, (9), 10042 (9), 10043 (9), 10044 (9), 10045 (10); UMMZ 88817. Nevis: MCZ 88874; PWH 414; RMNH 10431, 10448; UF 10040 (3). Sphaerodactylus macrolepis Gunther Sphaerodactutus macrolepis Gunther 1859:215. DIAGNOSIS. A sphaerodactyl with large, acute, strongly keeled, im- bricate scales, and with no middorsal area of granules. Dorsal scales with hair-bearing scale organs on their dorsodistal edges. Ventrals smooth on abdomen, may be keeled on chest. Gulars keeled. Inter- nasals 1 to 2; upper labials 8. Dorsal scales on tail imbricate, but lie flat against tail. Presacral vertebrae 26. Groundcolor brown, light to medium. Pattern a black ocellus, containing ohe or two white spots, in scapular region. Trunk with 4 to 6 dark longitudinal stripes, or scattered dark scales. Suprapostorbital dark stripes encircle the dark occipital spot and may be fused together to fgan a crude V on head 16 BULLETIN FLORIDA STATE MUSEUM V.01. 7 and neck, or may be fused with scapular ocellus, or may converge but remain slightly apart. KEY TO THE SUBSPECIES. Ventral scales keeled on chest or anterior abdomen; adult snout-vent length 18-24 mm.; scapular ocellus reduced or absent; dark scales on dorsum scattered at random; St. Martin, St., Barts, Anguilla, and Dog Island. --«_ Sphaprndactylus m. parous, new subspecies Ventral stales smooth, only gulars keeled; adult snout-vent length 18-28 mm.; scapular oodles usually large, may be reduced or absent; dark scales on dorsum arranged in, 4-6 longitudinal rows or absent; Virgin Islands. _ _trhnerndact!/lus m, macroteps ( As this subspecies does not occur in the Lesser Aritilles, .it is not dis- cussed further.) Sphaerodactylus macrolepis parous, new subspecies. HonoTypE. UF 10034.1, an adult male collected by Wayne King, on the island of St. Martin, 2-1/2 miles west, 1/4 mile north of Phil- ipsburg, 16 July 1958 ( fig. 5). DESCRIPTION OF HOLOTYPE ( AND OF SUBSPECIES 1, Dorsal scales large, acute, strongly keeled, and imbricate; 27 axilla to groin ( 28-29 in - li"C FIGURE 5. Holotype of Sphaerodact!/lus macrolepis pan>us, new subspecies-UF 10034.1. (A) subdigital lamellae of fourth toe, right pes; (B) scutellation of head; (C) dorsal markings. 1962 KINC : ANTILLEAN SPHAERODACTYLUS 17 other individuals, mean 25,9 ). 'Dorsal scales with hair-bearing scale organs, with one hair each, along dorsodistal edge. No middorsal area of granules. Ventrals smooth and_ round ( keeled on sides of abdomen in some specimens ); 29 axilla to groid (26-29 in other specimens, mean 27.5). Scale rows around midbody 50 ( 45-51 in other speci- mens, mean 48.4. Internasals 1 ( 1-2, mode 1). ,Upper labials 8. Lamellae of fourth toe 9 ( 9-11, mode 10). Escutcheon 19 scales wide ( 19-25 in other specimens )„ 5 scales long ( 4-7 in other specimens). Dorsal Scales of tail keeled, imbricate, and acute to round, but lie flat against tail. Midyentral row transversely expanded into plates. Presacral vertebrae 26. Snout-vent length 22 mm. ( smallest juvenile is 18 mm.; adults 18-24 min., mean 21.7) The pattern shows neither sexual dichromatic nor ontogenetic vari- ation. The dorsal groundcolor is a light to medium brown. A dark brown stripe extends from the nostril through the eye into the side of the neck where it fades. A pair of dark stripes extend caudad from the tip of the snout to a point between the eyes where they diverge from the midline and fuse with a suprapostorbital dark stripe immedi- ately posterior to the eye. These stripes then continue caudad and circle a dark occipital spot and nearly meet. The caudal ends of these stripes are separated by three scale rows ( in a few individuals the ends fuse ). A black ocellus in the scapular region contains two white spots in its center and has a light brown caudal b6rder ( in some ABC .-D ' .42'fz .94*{ 44* . .'.Ag .* 02'le. 4 9.'ma -' @r. Ficuit 6. Variation in the markings of adult Sphaerodactulus m. parous. (A) UF 10035; (B) PWH 474A; (C) UF 10036.2; (D) UF 16036.1. 18 BULLETIN FLORIDA STATE MUSEUM Vol. 7 individuals the two white spots fuse to form spectacles [fig. 6 A and, B], or the ocellus contains only one white spot [fig. 6 C], or thu ocellus is obhcure or absent [fig. 6 D]). The venter is a uniform dull white; the chin is speckled with brown. RANGE. The islands of Anguilla, Dog Island, St. Barts, St. Martin, and Tintamarre. REMARKS. Single mature eggs occur in the oviducts of several females collected during June and July. I collected the species beneath rocks in a cave on Anguilla, and in the leaf litter under rocks in a cave, and under the rocks from a stone wall on St. Martin. One egg was present with the geckos from the cave on St. Martin; it meas- ured 6 mm. in length and 4.5 mm. in diameter. S. m. parous and S. m. macrolepis are closely related to S. notatus, S. grandisquamis, and S. di#icilis, which probably represent races of the same species. MATERIAL EXAMINED. S. macrolepis parous ( paratypes) -St. Martin: MCZ 89741; PWH 474A (5); RMNH 10488 (8), 10444, 10461 (8), 10463; UF 10034 ( 10), 10035. Anguilla: RMNH 10447; UF 10086 (6), 10037 (2). Dog Island: RMNH 10466. Tintamarre: RMNH 10465. St. Barts: RMNH 10454. S. 1nacrolepis macrolepts-St. Croix: AMNH 39270-39278; MCZ 10725-10782; PWH 614 (5), 617, one unnumbered specimen; RMNH 6672 (4), 6675 (8), 6676 (6),_6679 (8); UMMZ 46750-46752 (17), 46759, 56016 (4), 78628 (5), 80550 (11), 80558- 80554 (18), 80556 (4), 80557 (9), 80558. St. Thomas: CNHM 42081- 42087; PWH 621, 628; RMNH 6674, 6681 (8), 6682 (2), 9174; UMMZ 73608-73609, 80547 (8), 80549 (8), 80559 (2), 80561 (8), 80562 (2), 80568, 80566 (8), 80629. St. John: MCZ 34587-84545 (14); PWH 618, 618A, 619; RMNH 6678 (3); UMMZ 78683 (10), 80551 (7). Tortola: CNHM 11288 (9); RMNH 6673 (20), 6677 (4); UMMZ 78607 (20), 73907 (.20), 80548 (19), 80564, 80565 (2). Virgin Gorda: MCZ 12256- 12259; RMNH 6671 (8); UMMZ 80550 (20). Anegada: MCZ 12260- 12268; UMMZ 80784 ( 20). Sphaerodactylus elegantulus Barbour Sphaerodactylus fantasticus Boulenger, 1885:223 (in part ). Sphaerodactghts degantubs Barbour, 1917b:163; Barbour, 1921: 269. DIAGNOSIS. A sphaerodactyl with large, acute, strongly keeldd, im- bricate scales, and an area of middorsal granular scales. Dorsal scales with knoblike and hair-bearing scale organs. Ventrals smooth and round. Gulats keeled on sides of throat. Internasals 1 to 8, upper labials 8. Dorsal scales on tail imbricate, but lie fiat against tail. Pre- sacral vertebrae 26. Pattern and color undergo an ontogenetic change 1962 KING: ANTILLEAN SPHAERODACTYLUS 19 -juveniles have light brown groundcolor and pattern of white cross- bands on neck and trunk; adults have a uniform brown color or darker brown speckling or vermiculations on trunk and head. HOLOTYPE. MCZ 12084, a juvenile collected by D. W. Griswold on the island of Antigua in 1917. DESCRIPTION OF THE SPECIES. Dorsal scales large acute, strongly keeled, and imbricate; 24 to 31 axilla to groin, mean 27.5. Dorsal scales with knoblike scale organs and hair-bearing scale organs, each with 2 hairs. Middorsal area of granular scales present, 1 to 8 rows wide. Ventrals smooth and round,25 to 32 axilla tb groin, mean 27.9. Gulars keeled on sides of throat, in a few individuals keeled gulars extend across throat in a narrow band. Number of scale rows around midbody 41 to 49, mean 45.2. Internasals lto 3, mode 2. Upper labials 2 to 3, mode 8. Lamellae of fourth toe 7 to 10, mode 10. Escutcheon of male 22 to 25 scales wide, 3 to 6 scales long. Dorsal scales on tail keeled and slightly imbricate, but lie flat against tail. Midventral row transversely expanded into plates. Presacral vertebrae 26. Snout-vent length of smallest juvenile 18 mm.; adults 18 to 29 mm., mean 25.1. The color pattern shows no sexual dichromatism, but it undergoes a ·marked ontogenetic change from juvenile to adult (fg. 7). The juvenile has a light brown dorsal groundcolor. The pattern consists A B C D zr t FIGURE 7. Variation in th6 markings of Sphaerodactulus elegantulus. (A) UF 10047.8; (B) MCZ 55610; (C) UF 10049.5; (D) UF 10051.3. A is a juvenile, the others are adult. 20 BULLETIN FLORIDA STATE MUSEUM Vol. 7 of dark brown-edged .white crossbands on the neck, trunk, and tail. There are 4 to 6 of these crossbands between the axilla and groin, mode 5-UF 10047.8 ( fig. 7 A) has 5 on the left side and 6 on the right; MCZ 55598 has 6 on the ieft and 5 on the right; MCZ 55618 has 4 on the left and 6 on the right. The chin and venter of the juvenile is white. As the lizard grows older the dorsal crossbands fade until tliey are indistinguishable from the groundcolor. The age at which the crossband pattern is lost seems to vary with the individual-some juveniles 13 to 14 mm. in snout-vent length are uniform brown ( MCZ 15544, 15547, and 55599), and several adults of 28 to 24 mm. snout- vent length still retain the crossbands ( UF 10049.5; MCZ 13487 and 55610). N6r does loss of the crossbanding seem correlated with sex. The dorsal color of the adult varies from light to dark brown with a white to light brown chin and venter. The dorsum may be uniform in color or may be speckled with dark scales. In some individuals these dark scales are numerous and form vermiculations, especially on the head. The occipital spot may be dark brown or may be light brown with a dark edge. RANGE. The islands of Antigua and Barbuda., (Eor comment on its reported occurrence on St. Lucia, see under S. oincenti festus) REMARKS, A large series collected on Barbuda during June and July includes a number of gravid females, each with one mature egg in its oviduct. Eggs occurred with this series of individuals in the moist debris under palm fronds and rocks in Darby Cave, a large sinkhole., The eggs measured 7.0 mm. by 5.5 mm. Boulenger ( 1885:224 ) lists S. fantasticus from Antigua. He describes fantasticus as a sphaerodactyl. that is light brown above and has a medium brown head vermiculated with white lines. This description, which Bts elegantulus better than fantasticus, is probably based on the former. The crossbanded juvenile of elegantulus has been confused with the crossbanded juvenile of S. cinereus of Cuba and Hispaniola. Two species from Barbuda in the U. S. National Museum ( USNM 187828- 187829, an adult and a juvenile), are catalogued as S. elegans (a duo- nym of S. cinereus based on its crossbanded juvenile). S. elegantulus appears to be closely related to S. fantasticus and S. sabanus, but is easily distinguished by its color pattern. MATERIAL EXA*INED. Antigua: MCZ 12084 ( holotype), 18487- 18492, 18494-18495, 15588-15539, 15541, 15548-15544, 28741, 55594- 55618; AMNH 24708; PWH 591, 594 (5), 595 (5); UMMZ 76096 (7); USNM 61067. Barbuda: MCZ 15546-15547, 57770-57774; PWH 600 1962 KING : ANTILLEAN SPHAERODACTYLUS 21 (7), 608 (15); UF 10046 (9), 10047 (8), 10048 (4), 10049 (9), 10050 (9), 10051 (9), 10052 (5); USNM 187828-137829. Sphaerodactglus fantasticus Dumdril and Bibron Sphaeriodactulus fantasticus Dum6ril and Bibron, 1886:406, Dum6ril, 1855:469. Sphaerodactylus fantasticus Barbour, 1915:72, 1921:262. Sphaerodactylus sputator Andersson, 1900:28. Barbour ( 1915:73) points out that the type locality of this species was originally listed as Martinique i Dum6ril and Bibron, 1836:407). The syntypes were collected by Mr. P16e, notorious for tire many incorrect localities on his "Martinique" specimens. Stejneger ( 1904: 622) states that Plde's trip through the Antilles began in Puerto Rico and ended in Martinique, from where he sent all his, specimens to the Mus6um National d'Hist6ire Naturelle, Pari_s. Apparently all of the specimens were assumed to have been collected on Martinique. Bar- bour ( 1915:78) compared Dum6ril and Bibron's description of fan- tasticus with a series of sphaerodactyls from Guadeloupe, and con- cluded that Guadeloupe was its correct type locality, not Martinique. Recently, however, three sphaerodactyls-closely related to Bar- bour's fantasticus-were collected on Monserrat. The spe6imens Bt Dum6ril and Bibron's description almost as easily as the Guadeloupe specimens do, yet scutellation differences exist between the Montserrat and Guadeloupe populations. Examinations of the syntypes of Sphae- rodactylus fantasticus Dum6ril and Bibron ( MNHN 1772) shows that the number of dorsal scales from axilla to groin-29 in all of the syn- types-is included within the variation of the Guadeloupe population (28-29), and not within the variation of the Montserrat population (85-38). The number of scale rows around midbody is 45 to 49 in the syntypes-also within the range of variation in the Guadeloupe (41-50), but not the Montserrat ( 52-54), population. Differences also exist in color pattern between the two populations, but these afe slight. In addition to these two islands, S. fantasticus has been collected on D6sirade. In color D6sirade specimens are identical to ones from Guadeloupe. In scutellation the two are similar save for one charac- ter-the number of dorsal scales from axilla to groin is 80 to 32 in the D6sirade specimens. A larger series of specimens will probably pro- duce a range of yariation in this character overlapping that of the Guadeloupe population. On the basis of the observed variation in color and scutellation 22 BULLETIN FLORIDA STATE MUSEUM Vol. 7 between the three populations-D6sirade, Guadeloupe, and Mont- serrat-Barbour's belief that the syntypes of Sphaerodactylus fan- tasticus Dum6ril and Bibron were collected on Guadeloupe seems justified. DIAGNOSIS. A sphaerodactyl with large, keeled, acute to round and slightly imbricate dorsal scales, and middorsal area of granular scales. Dorsal scales with both knoblike and hair-bearing scale organs. Ven- trals keeled or smooth, and acute to round. Gulars keeled. Internasals 1 to 8, upper labials 3, Dorsal scales on tail imbricate, but lie flat against tail. Presacral vertebrae 26. Groundcolor brown, medium to light. Pattern sexually dichromatic-males with uniform brown heads, or light brown heads with dark brown speckling or dark brown post- orbital stripes; females with light to medium brown heads with dark brown postorbital stripes extending caudad from eyes along neck and fading on trunk. KEY To THE SUBSPECIES. Ventral scales keeled or smooth; 28-34 dorsal scales from axilla to groin; 41-50 scale rows around midbody; Guadeloupe, D6sirade , Sphne:rndactulus f· fdntasticus Ventral scales smooth; 85-38 dorsal scales from axilla to groin; 52-54 scale rows around midbody; M6ntserrat. Sphaerodactulus f. ligniserut,Zus, new subspecies Sphaerodactylus fantasticus fantasticus Dum6ril and Bibron Sphaeriodactylus fantasticus Dumdril and Bibron, 1836:406; Dum6ril, 1855:469. Sphaerodactylus fantasticus Barbour, 1915:72, 1921:262. Sphaerodactylus sputator Andersson, 1900: 28. SYNTYPES, Three adult males, MNHN 1772, collected by M. Pl6e on the island of "Martinique" ( = Guadeloupe ), ( fig. 8 E). DESCRIPTION OF THE SPECIES. Dorsal scales large, keeled, acute to round, and slightly imbricate; 28 to 84 axilla to groin, mean 25.7. Dorsal scales with knoblike scale organs and hair-bearing scale organs, each with one hair. Area of middorsal granular scales present, 2 to 4 rows wide. Ventrals lightly keeled or smooth ( keeled dorsal scales extend down onto lateral edges of venter, and in some individuals continue across stomach), and acute to round; 27 to 32 axilla to groin, mean 29.3. Gulars strongly keeled, and extend caudad onto chest in all specimens. Scale rows around midbody 41 to 50, mean 44.8. Inter- nasals 1 to 3, mode 2. Upper labials 8. Lamellae of fourth toe 8 to 12, mode 9 or 10. Escutcheon of male 28 to 29 scales wide, and 8 to 5 scales long. Dorsal scales of tail keeled, acute, and imbricate, but lie fiat· against tail. Midventral row transversely expanded into plates. 1962 KING: ANTILLEAN SPHAERODACTYLUS 23 Presacral vertebrae 26. Snout-vent length of smallest juvenile is 18 mm.; adults 22 to 29 mm., mean 26.1. The pattern is sexually dichromatic. Both sexes have a brown dorsal groundcolor. The female has a dark brown stripe from the nostril to the eye, which continues as a postorbital dark stripe onto the neck and converges slightly in the scapular region with one from the Oppo- $ite side. In some individuals a dark stripe, beginning as a subpost- orbital stripe, parallels the postorbital stripe on the neck and in the scapular region, and is separated from it by a light strip of ground- color. Both fade into the groundcolor on the trunk. A dark stripe extends from the snout to a point between the eyes where it divides into two, each branch continuing to a point immediately behind the orbit. They then continue caudad and toward the midline, where + 2 0 ;50;U D ; sn1/}popoignydS }Inpu jo sB u ill 044 U I U O IJB LIE A '8 gR A P 21 AS ,p. ..~ -/. , -/1/_/4 ././-.. .-- I. ' A LLI I. C !9890I (a ) 4990I Z O K ) 56990I ZO IN ) !6p90I ZO W (V ) 918 fi pU B 'a 'o :sa le ,11 3 ; 9 IE plle V '( sedt'4Uks aait[} jO 000 ) E LLI N H =<,BLEEJE*~Tls Ir ,1. 'l™7,•~]rf-ler -Li. 9*'s=R, ,·r-m:~3 - 6 *1 6 ~m 1 -- - - 12-1 1 -1 11- 1 " U ~ir \-3. -1, 13/~ Mv I M. 2 7 - 1 '2- ---4 -1 .- .saIBUI 24 BULLETIN FLORIDA STATE MUSEUM Vol. 7 anterior to the scapular region they unite to form a dark vertebral stripe, exactly the width of the area of middorsal granules, that continues to the base of the tail. The occipital spot is white to light brown and has a dark br6wn Outline, and may be connected by dark lines to the head stripes. In some individuals this complex of markings is obscure or ill-defined (fig. 8 B), in others sharp and distinct (Bg. 8 A). Males have a light brown head which may lack all markings, or have only a hint of the female pattern ( Bg. 8 C). Other males have the dark stripes broken into small dark brown spots scattered evenly over the sides and upper surface ( fig. 8 D). In still others the dark brown spots coalesce to produce a dark head with light stripes or spots ( fig. 8 E). Trunk markings of both sexes are the same. Both have the dark vertebral stripe and a scattering of dark scales which form rough crossbands or vermiculations. The groundcolor of the tail becomes lighter than the body color, at its base, changing to a dull white or light brown toward its tip. Dark scales on the tail form broken longi- tudinal lines, crossbands, or spots, or may enclose a few white scales to form ocelli. The chin and throat are uniform white or may have a series of dark brown scales arranged in longitudinal rows. The juvenile pattern is similar to that of the adult female except that the pattern is usually sharp and clear. RANGE. The islands of. Guadeloupe and D6sirade, and probably Iles des Saintes and Marie Galante. REMARKs. Dumdril and Bibron ( 1886: 406 ) described S. fantasticus, on the basis of the three types ( MNHN 1772), as having a black head with white vermiculations. They also (1836:407) described a variety of fantasticus that was identical in scutellati6n, but differed in the coloration of the head. These three specimens ( MNHN 1770) have longitudinal stripes on their heads. One of the syntypes of the species was figured in that publication ( plate 82, fig. 2). Later Dum6ril ( 1855:469) redescribed and figured ( plate 17, fig. 8 ) both forms. Examination of the three syntypes· of S. fantasticus ( MNHN 1772) reveals that they are all males ( with dark heads). The three syntypes of the variety ( MNHN 1770) are two females ( with longi- tudinally striped heads) and a single male ( with a uniform brown head). Dumdril and Bibron's variety of S. fantasticus is based on the sexually dichromatic pattern of the species. A roentgenogram reveals that this species ( USNM 108984) ingests snails as part of its diet. The soft parts of these animals are digested and the shells probably passed in the fecal material. Barbour ( 1921:263) reports that Noble collected Single oblong eggs 1962 KING: ANTILLEAN SPHAERODACTYLUS 25 of this species in rotten wood. The eggs measured,6.5 mm. by 4.75 mm. The series of individuals that Noble collected during the summer of 1914 contains a number of gravid females. Each contains one mature, shelled egg. MATERIAL EXAMINED. Guadeloupe: MNHN 1772 (3) syntypes, 1,770 (3); AMNH 24714; MCZ 10681-10682, 10684-10680 (81); RMNH 2681 (2), 2688 (2); USNM 108984. Ddsirade: MCZ 66419-66420. Sphaerodactylus fantasticus Zigniservulus, new subspecies. HOLOTYPE. MCZ 66968, an adult male collected by P6re Pinchon, at Plymouth, Montserrat (fig. 9 C). DESCRIPTION OF HOLOTYPE ( AND OF SUBSPECIES). Dorsal scales keeled, acute to round, and slightly imbricate; 88 axilla to groin (85 to 38, mean 37.0). Dorsal scales with knoblike and hair-bearing scale organs, each with 1 to 2 hairs. Area of middorsal granular scales pres- ent, 8 rows wide. Ventrals smooth, and acute to round; 84 axilla to groin ( 82 to 84, mean 32.6). Gulars keeled across throat. Scale rows around midbody 54 (52 to 55, mean 53.6). Internasals 2 (1 to 2, mode 2). Upper labials 8. Lamellae of the fourth toe 10 ( 10 to 11, mode 11 ). Escutcheon 28 scales wide ( 27 to 28) , and 6 scales long. Dorsal scales of tail keeled, acute, and imbricate, but lie flat against tail. Midventral row transversely expanded into plates. Presacral vertebrae 26. Snout- vent length 21.5 mm. ( 21.5 to 22.5 mm., mean 22.6). The pattern is sexually dichromatic. Both sexes have a light brown dorsal groundcolor. The holotype, a male, ha5 an obscure medium brown stripe from the nostril to the eye. It continues caudad from the eye as a postorbital dark stripe and extends onto the neck where it · ' swings toward the midlihe to become a paravertebral stripe. It con- tinues onto the trunk where it fades into the groundcolor. A faint dark stripe extends from the tip of the snout to a point between the eyes where it divides into two. Each branch continues caudad, swings slightly laterad to 'encircle an occipital spot, and returns to the mid- line where it'fuses with the one from the opposite side. It immedi- ately fades into the groundcolor in the scapular region. The occipital spot is a light gray-brown triangle with dark edges. The trunk is a uniform light brown. The area of middorsal granular scales is slightly darker than the surrounding scales. The tail is light brown, with scattered white scales arranged in irregular transverse series. The chin, throat, and venter are white. The pattern of the femaie is like that of the male, but much more vivid. The stripe from the nostril to the eye is dark brown. The post- 26 BULLETIN FLORIDA STATE MUSEUM Vol. 7 A 8 C , FIGURE 9. Variation in the markings of adult Sphaerodactjlus fantasticus ligni- seroutus.(A) MCZ 66969; (B) MCZ 66967; (C ) MCZ 66968. Aisa female, B and C, males. C is the holotype. orbital stripe is dark brown to black, and is bordered above and below by a light brown to white stripe of equal width. It continues as a paravertebral stripe along the trunk and onto the base of the -tail. The stripe from the tip of the snout, which divides and encircles the occipital spot, is dark brown. It continues onto the trunk as a vertebral dark stripe exactly the width of the middorsal area of gran- ular scales. The occipital spot is white with dark brown to black edges. The chin and throat are white with dark brown to black bars and vermiculations. The venter is white, the edge of each scale sparsely stippled with black. These variations in pattern are illustrated in figure 9. RANGE. The island of Montserrat. REMARKS. S. f. ligniserculus appears to be almost intermediate be- tween S. f. fantasticus of Guadeloupe and S. sabanus of nearby Nevis, St. Kitts, Statia, and Saba. The color patterns of all three are similar, and it is easy to imagine how the sabanus pattern could be derived from the fantasticus pattern. S. f. fantasticus has large dorsal scales ( 23-24 axilla to groin) and S. sabanus has smaller ones ( 83-42). S. f. ligniseroulus is intermediate ( 85-88 ). It is also intermediate in the Other scutellation differences that exist between S. f. fantasticus and S. sabanus. However, no fantasticus possesses as many presacral ver- tebrae as S. sabanus. 1962 KING: ANTILLEAN SPHAERODACTYLUS 27 The dark-headed males, seen in both S. f. fantasticus ( fig. 8 E) and S. sabanus (fig. 4 C), have n6t yet been observed in ligniservulus. MATERIAL EXAMINED. Montserrat: MCZ 66968 (holotype), 66967, 66969 ( paratypes). Sphaerodactylus.microlepis Reinhardt and Lutken Sphaerodactutus microtepis Reinhardt and Lutken, 1862:278; Barbour, 1921:267. DIAGNOSIS. A large sphaerodactyl with medium-sized, keeled, acute, and slightly imbricate dorsal scales, and a middorsal area of smaller keeled scales ( no well-defined area of granules). Dorsal scales have hair-bearing scale organs on their dorsodistal edge: Ventrals strongly keeled and acute. Gulars strongly keeled. Internasals 2 to 4; upper labials 8. Dorsal scales of tail imbricate, and stand erect from tail. Presacral vertebrae 26. Color and pattern of dorsum light brown, which may have dark brewn crossbands. Venter white. Head with - straw brown grounde61or. Dark brown postorbital stripes form a V on neck. Dark brown.or black collar often present. Chin and throat with dark brown and white longitudinal stripes. HOLOTYPE. A specimen ( museum number unknown) in the Kobe- havns Universitet, Zoologisk Museum, Copenhagen, collected by Mr. Knudsen on the island of St. Lucia. DESCRIFTION OF THE SPECIES. Dorsal scales keeled, acute, and slightly imbricate; 42 to 58 axilla to groin, mean 47.4. Dorsal scales with hair- bearing scale organs, bearing one hair each, on dorsodistal edge. No well-defined middorsal area of granules, but middorsal scales reduced in size. Ventrals strongly keeled ( more strongly than dorsals), ind acute; 82 to 40 axilla to groin, mean 34.5. Gulars strongly keeled. Num- ber of midbody scale rows 62.to 74, mean 68.7. Internasals 2 to 4, mode 3.. Upper labials 3. Lamellae of fourth toe 9 to 12, mode 16 and 11. Escutcheon of male 22 to 81 scales wide, and 7 to 10 scales long. Dorsal scales on tail keeled, acute, and imbricate-almost conical ( triangular in cross-section ) - and stand erect giving a fuzzy appear- ance to tail. Midventral row acute, keeled, and not transversely expanded. Presacral vertebrae 26 ( one specimen in series UMMZ 71990 has 25). Snout-vent length of smallest juvenile 24 mm.; adults 28 to 38 mm., mean 81.2. The color pattern of adults shows a wide range of variation. No sexual dichromatism is evident, but there seems to be a slight onto- genetic change from juvenile to adult. The juveniles and some adults have a light brown dorsal groubdcolor with a series of 6 to 7 dark brown crossbarids between the axilla and groin. In some adults these 28 BULLETIN FLORIDA STATE MUSEUM Vol. 7 crossbands break up or fade until they are indistinguishable from the groundcolor. On the neck just anterior to the shoulders is a brown or black collar with a white or light brown posterior border. A black to dark brown postorbital stripe extends caudad and toward the mid- line, and joins the one from the opposite side on the neck. These stripes are faded or absent in some specimens (fig. 10 D ) and in some they may fail to join. They are frequently connected to the dark collar band ( fig. 10 A and C). On each side of the midline on the neck, in the angle formed by the postorbital stripes and the dark collar, is a white spot (fig. 10 A and C). Some individuals with a faded collar and postorbital stripes show a trace of these spots. The 1¢f' -'.0 34+ -t- 0 .% -1.:2ikrE,»2: 1-1: ,#4~-~2~0 0 C V ) 'stc 01010?lu snl/lioopoievllds qlnpe p sBur)[reu! atp u! uor}ejm A '0,I E,Ens '*,99Ze ZO K (a ) !6,695£ ZO IN ('0 ) 1£1,96£ ZO IN (i[) 1£995£ ZODN 2:r 1 starting a. new population. The eggs of Sphaerodactylus, and of geckos in general, ha-ve hard calcareous shells whith are nearly impervious to salt water. Brown and Alcala ( 1957:89) demonstrate that the eggs of three species of Eacific geckos ( Cosymbotus platyurus, Gehyra mutilata, and Hemi- dact!/lus frenatus ) can be exposed to sea water for at least 11 days and remain *liable. The eggs, ,laid in communal nests and relatively unharmed by ·sea water, eliminate the need for a gravid female lizard or male' and 40 BULLETIN FLORIDA STATE MUSEUM Vol.,7, female lizards to be transported to another island in order to start a new population-a single nest log is sufficient. Logs and similar vegetative flotsam suitable for rafts are swept northward into the Lesser Antilles when the Orinoco River floods each summer. The North Equatorial Current, which picks up these ' rafts, sweeps northwest past the Orinoco and up into the Caribbean at the rate of 20 to 80 miles per 24-hour period ( Sverd«rup, Johnson, and Fleming, 1942:642). The average open water distance between adjacent islands from Trinidad to Anguilla is 24 miles. If the rate of drift of a raft were equal to the velocity of' the current, it would take an average of one or two days to raft from one island to the next. The wind in this same area blows from an easterly or northeasterly direction. This would not hinder the westward progress of a raft, but would greatly slow its northward drift. Each year probably many animals, especially reptiles, are swept toward the Lesser Antilles on rafts. Guilding ( 1828) cites a boa constrictor ( Constrictor constrictor) that drifted ashore on St. Vincent on the trunk of a cedar tree. Feilden ( 1889: 298) states that an alli- gator ( Crocodylus intermedius) rafted to Barbados on the trunk of a tree in 1886 . Another Crocodylus intermedius came ashore on Grenada in 1910, with no apparent raft ( Barbour, 1914:222). Another example of the general direction of current drift in this island chain was supplied by_the_eryption Qf -Mount Pelee on Mar. tinique in 1902. Pumice and dugout canoes from this island drifted ashore in the Virgin Islands far to the north ( Barbour, 1917a:97). With adaptations for communal nesting and production of calca- reous-shelled eggs, and with favorable winds and currents, I believe the presence of these lizards in the Lesser Antilles can best be accounted for by waif dispersal through rafting. DISTRIBUTION Undefwood ( 1954a: 128) states thatthe present distribution of Sphaerodactylus in the Lesser Antilles suggests that its dispersal is southward from the Greater Antilles. The basis for this is the apparent absence of the genus from Grenada, which he feels has been ade- quately collected. This statement also hinges on his belief that Sphaerodactylus has only recently entered South America ( Under- wood, 1954a:123). If the lizards are dispersing southward in the Lesser Antilles, they are being carried against the ocean cufrents, against the prevailing winds, and against the direction taken by cycl6nic st6rms. If the geckos are being dispersed by means of wind 1962 KING: . ANTILLEAN SPHAERODACTYLUS 41 or water, it is more reasonable to assume that they are being moved northward from the South American continent. Barbour ( 1980,1985, 1937 ) lists several reptilian genera whose distributions suggest this northwafd immigration„Bothrops, Constrictor, Boa, Pseudoboa, Ken- tropyx, Iguana, and Thecadactylus. These genera have a wide dis- tribution on the mainland, but reach the West Indies only in the Lesser Antilles. The little knowledge that we have of the Pleistocene herpetofauna of the Lesser Antilles suggests the same northward movements. Dur- ing the Pleistocene, Thecadactylus, Pseudoboa, and a now extinct species of frog, Hyla barbudensis, related to the South American forms, occurred as far north in the island chain as Barbuda ( Auffen- berg, 1958). As Sphaerodactylus is unrecorded as a fossil, nothing can be stated positively about its movements to or from the mainland, north or south through the islands. Barbour ( 1921:218') lists several morphological characters that he considers primitive or advanced for the genus Splmerodactylus. An example of a primitive character that he uses is the presence of numerous small granular dorsal scales, which presumably gave rise to small keeled granular scales, and then to keeled scales, and ulti- mately to large strongly keeled scales. In ofder te determine which characters might be considered primitive and which advanced, I examined other members of the family Sphaerodactylidae ( Under- wood, 1954b:476), which contains Bve genera of lizards-Coleodac- tglus. Gonatodes, Lepidoblepharis, Pseudogonatodes, and Sphaerodac- tylus. Noble ( 1921: 14) and Parker ( 1926.801) show that Sph£zero- dactylus may have been derived from an ancestral series of which Gonatodes represents the basal grade and Pseudogonatodes, Lepido- blepharis, and Coleodaetylus are intermediate grades in the order of development. Sphaerodactylus then is considered most advanced. Their work is based largely on osteological characters and scutellation of the ungual sheath. The present distribution of these more primitive members of the family is limited to Central America and northern South America (Underwood, 1954b:476). The most advanced a~nus, Sphaerodactylus, occurs in Central America, northern South America, and the Antilles. Examination of seyeral species of Sphaerodactylus from all parts of the range shows that the mainland species possess the greatest number of presumed primitive characters. Primitive characters are defined here as those which form a basal grade capable of giving rise to other, i.e. advanced, .characters. In addition, most primitive characters are close to the condition found in other members of the 42 BULLETIN FLORIDA STATE MUSEUM Vol. 7 family Sphaerodactylidae. Primitive characters used in this stitdy include the granular dorsal scales mentioned by Barbour, smooth gular scales, smooth round ventral scales, dorsal scales with large scale organs bearing numerous hairs, and the escutcheon of· the male long and lacking thigh extensions. Analysis of series of these presumed evolutionary grades reveals nothing of the dispersal routes of these animals. Figures 14 through 16 show the distribution of several characters of scutellation and col- oration. Such analyses do reveal that the most primitive species occur on the mainland of Central and South America, and the most ad- vanced in th'e Antilies. Nothing more definite can be stated about the point of origin of Sphaerodactglus at present. As mentioned above, no sphaerodactyl has yet been found in Grenada or the Grenadines. No sphaerodactyl is recorded from a number of additional islands, Sombrero, Redonda, Marie Galante, Iles des Saintes, and Barbados. Sombrero and Redonda are extremely small and remote: Sombrero is inhabited only by a crew of lighthouse tenders; Redonda is uninhabited. Few collectors have ever visited these islands, and chances are, if Sphaerodactyke exists on them, it was 6verlooked. Montserrat is a relatively large island and is in- habited. Its herpetofauna is supposedly well known ( see Sphaero- dact!/lus f. ligniservulus), but as late as 1958 it was possible to add two species as large and conspicuous as Iguana and Geochelone to the list of animals known to occur there ( these specimens are in the University of Florida Collections). Grenacla and the Grenadines are equally well known. For these islands-Sombrero, Redonda, Marie Galante, Iles des Saintes, Grenada, and the Crenadines-I believe sufficient collecting will demonstrate sphaerodactyls to be present. Barbados seems genuinely to laek the animal. This island is excep- tional among the southern Lesser Antilles because of the paucity of species in its reptile fauna ( Barbour, 1980, 1985, 1987; Feilden, 1889: 295; Grant, 1959b:97.). Extensive cultivation of sugarcane on the island has destroyed most of the original vegetation. This may account in part for the scarcity of reptiles. Grant ( 1959b:97) believes that most of the eight reptile genera present were introduced by man. He states that, "Barbados herpetology could be dismissed with the remark that in pre-columbian times the only reptiles were Dromicus per~uscus and possibly Kentropyx intermedius...." However, Iguana iguana is recorded as a Pleistocene fossil in Barbados ( Swinton, 1987:306). All things donsidered, Sphaerodact~/lus probably does not occur in Bar- bados. The island lies approximately 100 miles east of the main chain of islands, and in this p6sition is "upstream' in terms of ocean cur- 1962 KING : ANTILLEAN SPHAERODACTYLUS 43 25.9 34.6 48 -4 M. PARVUS ~9' SPUTATOR 55.5 5 27.5 38.0 , ~ 27.5 57.6 SAB'ANUS 0 ELEGANTULUS 45.2 30 . 7 %0 - e 27 . 9 37.0 • 33 :6 0. 4/GNISERVULUS ~ 25 .7 655 F FANTASTICUS 44 .3 32.6 · 29.3 47.3 U 58.4 V. FESTUS 47.4 30.1 ~~ MIC·ROLEPIS 63.7 34.5 0 49.5 61.6 V. VI.NCEN·TJ /3 ' ~~ 32 , 1 V a 0 D 679 77.2 M.OLEI 38.0 - , )r, FIGURE 14. Geographic distribution of scale counts. Mean number of dorsal scales from axilla to groin is above the specific epithet; mean number of. ventral scales, below the epithet; ·mean number of scale rows around midbody, beside the epithet. 44 BULLETIN FLORIDA STATE MUSEUM Vol. 7 MEEK S. C m7,71 M. PAR .VUS ~,~ <5? ~ SPUTATOR /.4 Pt....Y V ~ ELEGANTULUS 0 SABANUS 11814 0. F. LIGNISERY[/LUS ~ fh~:rs, ~Im ' 'ANTAST,cus 28* 1 3- I "WN~0 V. FEST, US ~ 61* V * ~ MICROLEPIS v. VI'NCENTI ~*1~ Or 0 V .0 O lill MOLEI %»6--%\Ik FIGURE 15. Geographic distribution of color patterns. The patterns are simplified presentations of the juvenile or young adult patterns. 1962 KING: ANTILLEAN SPHAERODACTYLUS 45 ~ M. PARVUS .0 SPUTATOR ~ % 4 ELEGANT.ULUS ~ ~.~ SABANUS 0 CS· F. LIGNISERVULUS G b Tw F. FANTASTICUS Ad.7 D *a..A., v. FESTUS ~~ MICROLEPIS ~ K VINCENT.1 0 5 0 0 a . MOLEI FIGURE 16. Geographic distribution of scale 6rgans. The sizes of the organs are exaggerated for purposes of illustration. 46 BULLETIN FLORIDA STATE MUSEUM Vol. 7 SOMBRERO . DOG ISL. < ~1| ANGUILLA ~·'67'ST.·MAR'IN f ,-·z~I r.z:CH' BARBUDASABA /,I 1\ 2 ST. KITTS _fl ANTIGUA REDONDA ==940 DES,RADE GUADELOUPE ,lk- ' / V 4 MARIE GALANTE ILES DES SAINTES \./61 V D DOMINICA MARTINIQUE ~ 7 ST. LUCIA ~ ~ ST. VINCENT Frli BARBADOS ~7 - THE GRENADINES O GRENADA TOBAGO ~ ~ 9 -3--UZ355 do-»\C YRINIDAD 1 VENIZUELA FIGURE 17. Geographic distribution of the races of Sphaerodactylus in the Lesser Antilles. (1) S. macrolepis parous, 12) S. sputator; (3) S. sabanus; .( 4) S. elegantulus; (5)S. fantasticus ligniservulus; (6) S.- fantasticus fantasticus; (7) S. vincenti festus and S. microlepis,(8) S. uincenti vincenti, (9) S. motei. 1962 KING: ANTILLEAN SPHAERODACTYLUS 47 rents and wind directions and thus isolated from the main route of dispersal. A raft leaving the mainland of South America would have to cross a major part of the North Equatorial current to reach Barba- dos. This necessitates beginning its trip farther south on the coast of South America, possibly in the vicinity of Surinam, French Guiana, or Brazil. Even then the wind directi6n is unfavorable. Thus the -chances of a raft reaching Barbados are slight indeed, which may account for its depauperate reptile fauna. Figure 17 plots the known distributions of the races of sphaerodac- tyls in the Lesser Antilles and Trinidad. SUMMARY AND CONCLUSIONS Eight species of Sphaerodactylus are known from the Lesser Antilles and Trinidad-S. elegantulus, S. fantasticus, S. macrolepis, S. micro- lepis, S. molei.. S. sabanus, S. sputator, and S. vincenti. Each of these · species is described and illustrated. Sparkman's ( 1784) type series of S. sputator is shown to consist of two species. A lectotype, based on the largest of his specimens, is designated and redescribed. A new race, S. macrolepis paruus, is described from Anguilla, Dog Island, St. Barts, St. Martin, and, Tintamarre. It is distinguished from S. macrolepis macrolepis by its keeled chest 5cales: A second race, S. fantasticus ligniservulus, is described from Mont- serrat. It is separated from S. fantasticus fantastictis by the fact that its dorsal scales are smaller and more numerous, and its ventral scales are never keeled. S. monilifer is reduced to a synonym of S. festus. The color pattern and scutellation thought to be distinctive of monilifer are shown to be within the normal variation of festus, s, festus is itself considered to be conspecific with S. uincenU. The two forms are treated as sub- species, S. u. oincenti and S. u. festus. They are distinguishable by the number of dorsal scales from axilla to groin ( 36-49 in festus; 47-53 in vincenti), and slight differences in edlor pattern. Because the LessBr Antilles have never been connected to the main- , land of South America or to the Greater Antilles. waif dispersal is the only method of immigration open to these lizards. Rafting is believed ~ to be the principal method of dispersal used by Sphaerodactylus because of adaptations for communal nesting and the laying of cal- careou5-shelled eggs impervious to sea water. The direction of immi- gration-in the Lesser Antilles-is thought to be northward into the islands from the South American mainland. Belief in dispersal by 48 BULLETIN FLORIDA STATE MUSEUM Vol. 7 means of rafting dictates this direction-ocean currents flow porth- west across the islands, trade winds blow from an easterly or north- easterly direction, and cyclonic storms move west-northwest. However, paleontological evidence is unavailable to support or deny this pro- posed direction of dispersal. Distribution of primitive and advanced characters of morphology and coloration fails to reveal any clines that might reflect direction of immigration. The evidence at hand does not support any definite conclusion regarding the place of origin of the Lesser Antillean sphaerodactyls. The species in the Lesser Antilles are closely related to and possibly conspecific with species widespread in the Greater Antilles ( S. macro- lepis with S. di#icilis, S. grandisquamis, and S. notatus) and on the mainland ( S. molei with S. lineolatus ). LITERATURE CITED Andersson, L. G. 1900. Catalogue of Linnean type-specimens of Linnaeus's Reptilia in the Royal Museum in Stockholm. Bihang till Kongliga Svenska Vetenskaps- Akademiens Handlingar, vol. 26, sec. 4, no. 1, pp. 1-29. Auffenberg, W. 1958. A small fossil herpetofauna from Barbuda, Leeward Islands, with the description of a new species of Hyla. Quart. Jour. Florida Acad. Sci., vol. 21, no. 8, pp. 248.254. Barbour, T. 1914. A contribution to the zoogeography of the West Indies, with special reference to amphibians and reptiles. Mem. Mus. Comp. Zool., vol. 44, no. 2, pp. 209-359. 1915. Recent notes regarding West Indian reptiles and amphibians. Proc. Biol. Soc. Washington, vol. 28, pp. 71-78. 1916. Some remarks upon Matthew'5 "Climate and Evolution," with supple- mentary note by W. D. Matthew. Ann. New York Acad. Sci., vol. 27, PP. 1-15 1917a. Notes on the herpetology of the Virgin Islands. Proc. Biol. Soc. Wash- ington, vol. 80, pA 97-104. - 1917b. A new Antillean Sphaerodactvlus. ibid., vol. 80, pp. 168-164. 1921. Sphaerodactylus. Mem. Mus. Comp. Zool; vol. 47, pp. 217-278. 1928. West Indian investigations of 1922. Occ. Pap. Mus. Zool. Univ. Michi- gan, no. 182, pp. 1-9. 1929. List of reptiles and amphibians to be found in this presidency-St. Christopher and Nevis. St. Kitts and Nevis Omcial Gazette, vol. 26, no. 80, p. 81. 1980. A list of Antillean reptiles and amphibians. 'Zoologica, vol. 11, no. 4, pp. 61-116. 1962 KING: ANTILLEAN SPHAERODACTYLUS 49 1985. Second list of Antillean reptiles and amphibians. Ibid., vol. 19, no. 8, pp. 77-142. 1937. Third list of Antillean reptiles and amphibians. Bull. Mus. Comp. Zool., vol. 82, pp. 77-166. Beebe, W. 1944. Field notes on the lizards of Kartabo, British Guiana, and Caripito, Venezuela. I Gekkonidae. Zoologica, vol. 29, no. 3, pp. 145-160. Boettgdr, 0. 1894. [Description of S. mold.] In R. R. Mole and F. W. Urich. 1894. A preliminary list of the reptiles and batrachians of the island of Trini- dad. Jbur. Trinidad Field Nat. Club, vol. 2, pA 77-99. Boulenger, G. A. 1885. Catalogue of the lizards in the British Museum ( Natural History). London. 2nd ed. vol. 1, pp. 1-497. 1891. On reptiles, batrachians, and fishes from the Lesser West Indies. Proc. Zool. Soc., London, pp. 351-357. Brown, W. C., and A. C. Alcala 1957. Viability of lizard eggs exposed to sea water. Copeia, 1957, pp. 89-41. Butterlin, 1 1957. La Constitution Geologique et la Structure des Antilles. Centre Na- tional de la Recherche Scientiftque, pp. 1-453. Cochran, D. M. 1988. Reptiles and amphibians from the Le5ser Antilles collected by Dr. S. T. Danforth. Proc. Biol. Soc. Washington, vol. 51, pp. 147-156. Darlington, F. J.. Jr. 1988. The origin of the fauna of the Greater Antilles, with discussion of dispersal of animals over water and through the air. Quart. Rev. Biol., vol. 13, pp. 274-300. 1957. Zoogeography: the geographical distribution of animals. New York: John Wiley and Sons, pp. 1-675. Duellman, W. E., and A. Schwartz 1958. Amphibians and reptiles of southern Florida. Bull. Florida State Mus., vol. 8, no. 5, pp. 181-324. Dum6ril, A. H. A. 1855. Description des reptiles nouveaux ou imparfaitement connus de la collection du Mudum d'Histoire Naturelle et· remarques sur la classi- fication et las caractdres des reptiles. Deuxidme M6moire. Archives du Mus6um National d'Histoire Naturelle, Paris, vol. 8, pp. 487-588 Dum6ril, A. M. C., and G. Bibron 1886. Erpdtologie g6n6rale ou histoire naturelle complate des reptiles. Paris: Librairie Encyclop6dique De Roret, vol. 8, Pp. 1-517. 50 BULLETIN FLORIDA STATE MUSEUM Vol. 7 Feilden, H. W. 1889, Notes on the reptiles of Barbados. Zoolbgist, vol. 8, no. 3, pp 295-298. Carman, S. 1887. On West Indian Geckonidae and Anguidae. Bull. Essex Inst., vol. 19, pp. 17-24. Graijt, C 1931. The sphaerodactyls of Porto Rico, Culebra, and Mona Islands. J6ur. Dept. Agric. Porto Rico, vol. 15, no. 3, pp. 199-213. -- 1932. Herpetological notes. Ibid., vol. 16, no. 4, pp. 405-407. 1959a. Identification and description of sphaerodactyls. Herpetologica, vol. 15, pp. 56-57. I959b. Herpetology of Barbados, B. W. I. Ibid., vol. 15, pp. 97-101. Gudger, E. W. 1921. Rains of fishes. Nat. Hist., vol. 21, pp. 607„619. Guilding, E. W. 1828. Observations on the zooldgy of the Caribbean Islands. Zool. Jour., vol. 8, pp. 408-408. Gunther, A 1859. On the reptiles from St. Croix, West Indies, collected by Messrs. A. and E. Newton. Ann. Mag. Nat. Hist., vol. 4, ser. 8, no. 21, pp. 269-217. 1888. Notes on reptiles and frogs from Dominica, West Indies. Ibid.,vol. 2, ser. 6, no. 48, pp. 862-866. King, W. 1960. The status 6f Sphaerodactulus pictus, with comments on the distribu- tion of.S. sputator and S. sabanus. Brev'iora, no. 132, PA 1-5. McAtee, W. L. 1917. Showers of organic·matter. Month. Weather Rev., vol. 45, pp. 217-224. Myers, G. S. 1988. Fresh-water Bshes and West indian zoogeography. Smithsonian Rept., 1987, pp. 389-364. Noble, G. K. 1921. The bony structure, and phyletic relations of Sphaerodact!/lus and al« lied genera, with the description of a new genus. Amer. Mus. Novitates, no. 4, PE· 1-16. Nbble, G. K., and G. C. Klingel 1932. The reptiles of Great Inagua Island. British West Indies. Amer. Mus. Novitates, no. 549, pp. 1-25. Parker, H. W. 1926. The neotropical lizards of'the genera Lepidoblepharis, Pseudogonatodes, Lathrogecko,' and Sphaerodactylus with the description of a new genus. Ann. Mag. Nat. Hist., vol. 17, ser. 9, no. 99, art. 40, pp. 291-301. 1962 KING: ANTILLEAN SPHAERODACTYLUS 51 1933. Some amphibians ·and reptiles from the Lesser Antilles. Ibid., vol. 11, ser. 10, no. 61, art. 18, pp. 151-158. Questial, A. 1941. The flora of St=Bartholomew ( French Wast Indies) and its origin. Basseterre, Guadeloupe: Imprimerie Catholique, .pp, :1„224. Reinhardt, J. and C. F. Lutken 1862. Bidrag til det vestindiske 6riges og navnligen til de dansk-vestindiske 6ers Herpetoiogie. Videnskabelige Meddelelser fra den naturhist6riBke F6rening i Kj6benhavn, pp. 153-291. Roux, J. 1927. Contribution A, l'erp6tologie du V6nizudla: Verhandlungen de 'Natur- forschenden Gesellschaft, vol. 38, pp. 252-261. Scharff, R. F. 1922. On the Origin of the West Indjan fauna. Bijdragen tot de dierkunde, Amsterdam, pp. 65-72. Schmidt, W. J. 1920. Einiges ubet die Hauptsinhesofgane der Agamiden, inhesondere von Calotes, nebst Bemerkungen uber diese Organe bei Geckonideh und Iguaniden. Anatomischer Andeger, vol: 53, pp. 413-139. Schuchert, C. 1935. Historical geology of the Antillean-Caribbean region. New York: John ~ Wiley and Sons, pp. 1-811. Shreve, B. 1947. On Venezutlan reptiles and amphibians collected by Dr. G. Kugler. Bull. Mus. Comp. Zool., vol. 99, no. 5, pp. 517-537. Simpson, G. G, 1956. Zoogeography of West Indian land mammals. Amer.. Mus. Novitates, no. 1759, pA 1-28: Sparrman, A. 1784. Lacerta *utator och .Lacerta bimaculata, tvA nya Odlor frdn Anferica. Kongliga Svenska Vetenskaps-Akademiens Handlingar, yol. 5, pA 164- 167. Steindachner, F. 1867. Reptilien. In Reise der 65terreichischen 'Fregatte Novarra um die Erde, in den Jahren 1857, 1858, 1859, unter den Befehlen des Commodore B.. von Wiillersdorf-Urbair. Zoologischer Theil, erster Band, Wirbely theire, pp. 1-98. · Stefneger, L. 1904. The herpetology of Poljo Rico. Ann. Rept. U.S. Nati. Museum, 1962, pp 553-724. 52 BULLETIN FLORIDA STATE MUSEUM Vol. 7 Sverdrup, H. U., M. W. Johnson, and R. H. Fleming 1942. The oceans, their physics, chemistry, and general biology. New York: Prentice-Hall pp. 1-1087. Swinton, W. E. 1937. Iguana remains from Barbados: Ann. Mag. Nat. Hist., vol. 19, ser. 10, no. 110, art. 24, pp. 306-307. Tannehill, I. R. 1938. Hurricanes, their nature and history. Princeton: Princeton Univ. Press, pp. 1-257. Underwood, G, 1954a. The distribution of Antillean reptiles. Nat. Hist. Notes Nat. Hist. Soc. Jamaica, no. 67, pp. 121,129. 1954b. On the classification and evolution of geckos. Proc. Zool. Soc. London, vol. 124, no. 3, pp. 469-492. 1957. On lizards of the family Pygopodidae. A contribution to the mor- phology and phylogeny of the Squamata. Jour. M6rph., vol. 100, no. 2, pp. 207-268. Werner, F. 1900. Ueber Reptilien und Batrachier aus Columbien und Trinidad II. Ver- handlungen der Zoologischen-botanischen Gesellschaft, Wien, vol. 50, pp. 262-272. Williams, E. E., and K. Koopman 1952. West Iridian fossil monkeys: Amer. Mus. Novitates, no. -1546, PIi. 1-16. Contributions to the BULLETIN OF THE FLORIDA STATE MUSEUM may be in any field of biology. Manuscripts dealing with natural history or systematic problems involving the southeastern United States or the Caribbean area are solicited especially. Manuscripts should be of medium length-50 to 200 pages. 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